Ennominae II (Geometrid Moths of Europe) [Annotated] 9004252223, 9789004252226

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The Geometrid Moths of Europe

Volume 6 Part 1

Bernd Müller, Sven Erlacher, Axel Hausmann, ­Hossein Rajaei, Pasi Sihvonen & Peder Skou

The Geometrid Moths of Europe Axel Hausmann, Pasi Sihvonen, Hossein Rajaei & Peder Skou (eds) Volume 6 Part 1

Bernd Müller, Sven Erlacher, Axel Hausmann, Hossein Rajaei, Pasi Sihvonen & Peder Skou Subfamily Ennominae II (Boarmiini, Gnophini, additions to previous volumes)

leiden

| boston

Editors: Axel Hausmann, Zoologische Staatssammlung München, Germany; Pasi Sihvonen, Finnish Museum of Natural History, Helsinki, Finland; Hossein Rajaei, State Museum of Natural History Stuttgar, Germany; Peder Skou, Ollerup, Denmark. Linguistic editor: Martin Corley, Faringdon, United Kingdom. Colour plates photographed and compiled by: Igor Kostjuk, Kiev, Ukraine. This publication should be cited as: Müller, B., Erlacher, S., Hausmann, A., Rajaei, H., Sihvonen, P. & Skou, P. (2019): Ennominae II. – In A. Hausmann, P. Sihvonen, H. Rajaei & P. Skou (eds): The Geometrid Moths of Europe 6: 1–906. Brill, Leiden.

Date of publication: 15.05.2019

Authors’ addresses: Bernd Müller, Weissdornallee 13, D-13158 Berlin, Germany, [email protected] Sven Erlacher, Museum of Natural History, Moritzstr. 20, D-09111 Chemnitz, Germany, [email protected] Axel Hausmann, SNSB – Bavarian State Museum of Zoology, Münchhausenstr. 21, D-81247 Munich, Germany, [email protected] Hossein Rajaei, State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany, [email protected] Pasi Sihvonen, Finnish Museum of Natural History, Zoology Unit, P. O. Box 17, FI-00014 University of Helsinki, Finland, [email protected] Peder Skou, Aamosen 1, Ollerup, DK-5762, Vester Skerninge, Denmark, [email protected]

Front cover: Erannis defoliaria (Clerck, 1759). Denmark: Fyn, Aamosen, 31 October 2018. Photo by Peder Skou, Denmark. Numerous photos of differential external features shown in Text-figs modified from Igor Kostjuk’s plate photographs. The work of Axel Hausmann was supported by grants of the Bavarian ministry of science (project BFB) and by the German ministry of Science (BMBF; project GBOL). The genetic analyses have received considerable support from Paul D. N. Hebert and his competent teams at the Centre for Biodiversity Genomics (CBG) and the Canadian Centre for DNA Barcoding (CCDB University of Guelph). The data management & analysis system BOLD was provided by Sujeevan Ratnasingham. The work was financially supported by Genome Canada (­Ontario Genomics Institute) in the framework of the iBOL I and II programs. Library of Congress Control Number: 2019938865 ISBN: 978-90-04-25222-6 (hardback set) ISBN: 978-90-04-40783-1 (part 1) ISBN: 978-90-04-40784-8 (part 2) ISBN: 978-90-04-38748-5 (e-book) Copyright 2019 by Koninklijke Brill NV, Leiden, The Netherlands. Koninklijke Brill NV incorporates the imprints Brill, Brill Hes & De Graaf, Brill Nijhoff, Brill Rodopi, Brill Sense, Hotei Publishing, mentis Verlag, Verlag Ferdinand Schöningh and Wilhelm Fink Verlag. All rights reserved. No part of this publication may be reproduced, translated, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by Koninklijke Brill NV provided that the appropriate fees are paid directly to The Copyright Clearance Center, 222 Rosewood Drive, Suite 910, Danvers, MA 01923, USA. Fees are subject to change. This book is printed on acid-free paper and produced in a sustainable manner.

Contents Part 1 Preface........................................................................................................................7 Abstract.......................................................................................................................9 Acknowledgements....................................................................................................15 Introduction...............................................................................................................21 Methodological notes................................................................................................23 Abbreviations............................................................................................................27 Systematic Account...................................................................................................29 Tribus Abraxini (part): Addition............................................................................29 Tribus Cassymini (part): Addition.........................................................................40 Tribus Macariini (part): Addition...........................................................................49 Tribus Prosopolophini Warren, 1894 (part): Addition...........................................52 Tribus Gnophini Duponchel, 1845........................................................................58 Tribus Boarmiini Duponchel, 1845.....................................................................253 New species for the fauna of Europe after publication of the previous volumes.......519 Index to scientific names..........................................................................................559

Part 2 Colour plates...........................................................................................................567 Genitalia plates........................................................................................................629 Appendix 1. Data of the microscopic preparations shown in male genitalia plates...... 761 Appendix 2. Data of the microscopic preparations shown in female genitalia plates.... 779 Revised, annotated systematic checklist of the ­Geometridae of Europe and adjacent areas, Vols 1–6.....................................................................................795 References...............................................................................................................873 Index to scientific names..........................................................................................903

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Preface Today, scientists agree that earth is suffering the sixth mass extinction with a rate of species extinction as much as 100 times the ‘normal rate’ (Ceballos & Ehrlich 2018) which may lead to the extinction of 40% of the world’s insect species over the next few decades (Sánchez-Bayo & Wyckhuys 2019). Together with many other alarming signals, as e.g. the severe loss of insect biomass (Hallman et al. 2017; Lister & Garcia 2018) and considering that 70–80% of our animal diversity are insects (Hausmann 2017) it is an imperative to monitor insect biodiversity on earth. In this context, the GME book series provides a modern baseline of knowledge for species identification, distribution, abundance etc. in Europe, which can be used to monitor biodiversity in its various forms. With this volume we have finished the publication of the Euro­pean inventory covering approx. 1000 species of geometrid moths, providing an integrative taxonomic revision based on both morphological, ecological, zoogeographical and mole­ cular data. This treatment deals with one of the largest animal families comprising approx. 10% of the European Lepidoptera fauna and close to 1% of the European insect fauna. Ennominae are the most species-rich geometrid subfamily with some 10,700 species having been described worldwide, so far (Scoble & Hausmann 2007), and the number of undescribed species will amount to at least 7,000 additional species, judging from the numerous unnamed genetic clusters in the Global Campaign of DNA Barcoding (iBOL Lepidoptera). The present sixth volume of The Geometrid Moths of Europe, providing an updated, valuable, modern tool for species identification for roughly the second half of European Ennominae (181 species), is a product of team-work. The division of tasks and responsibilities of the different authors are listed in the introduction. This volume also presents information on 21 species belonging to groups which were subject of the five previous volumes but which were recorded as new for the European fauna after publication of the related volumes. At the present stage (February 2019), we have generated, in the framework of the Global Geometrid Campaign of the Barcode of Life project, DNA barcodes of 988 European geo­ metrid species (99%; cf. Hausmann et al. 2011b; Hausmann et al. 2013). Tissues of the following eleven species are missing for completion: Anthometra plumularia, Idaea luteolaria, Idaea nexata, Idaea dromikos, Asthena lactularia, Eupithecia sardoa, Eupithecia karadaghensis, Eupithecia moecha, Isturgia homochromata, Psodos pyrenaica, Psodos werneri. As in the case of the two previous volumes we are happy that we could include this large amount of genetic information into our analyses and we thank Paul Hebert, Sujeevan Ratnasingham and their teams (Centre for Biodiversity Genomics, Guelph, Canada; cf. acknowledgements) for providing analysis tools and financial support. We hope that this book series will stimulate the re-examination of all existing collections, in order to improve quality of identification. As a continuation of the GME volumes, an update of the European checklist is envisaged for the future (www.faunaeur.org; www.zsm.mwn.de/lep/ gme.htm). The site of the Geometridae research initiative Forum Herbulot (www.herbulot.de) provides a list of family-group names, a global checklist of geometrid species and further information on geometridology. We would like to apologize to our readers that so much patience was needed in waiting for the publication of this last volume and we hope that the product will please and repay the long waiting time. Munich, 1.3.2019

Axel Hausmann (editor-in-chief)

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Abstract Volume 6 of the book series The Geometrid Moths of Europe presents the second and concluding part of the revision of the European taxa of the subfamily Ennominae, covering 202 species. The current volume focuses on Gnophini (part, 68 species) and Boarmiini (100 species), but it also treats parts of Ennominae tribes Abraxini (3 species), Cassymini (2 species), Macariini (1 species) and Prosopolophini (2 species). 5 species are of uncertain tribus association within Ennominae. We also present new species for the fauna of Europe after publication of the previous volumes (vols. 1–5), altogether 21 species. The species are presented with their full name, reference to the original description, ­fully referenced junior synonyms, (potentially) unavailable names, description, distribution, phenology, biology including food-plants, habitat, differential features from similar species, genetic data and remarks if needed. Further, for each species the following information is provided: colour photographs of the adults, greyscale photographs of the male genitalia, including in many instances the everted vesica, greyscale photographs of the female genitalia, and distribution map for Europe and adjacent countries. 219 text-figures illustrate and explain diagnostic characters and other morphological structures. Systematic and annotated checklist of the Geometridae of Europe and adjacent areas is provided for volumes 1–6, which covers the entire fauna. It includes new described taxa and taxonomic changes since the publication of earlier volumes: GME1: 2001 (Archiearinae, Oenochrominae, Geometrinae); GME2: 2004 (Sterrhinae); GME3: 2013 (Larentiinae I); GME4: 2003 (Larentiinae II); GME5: 2015 (Ennominae I). Total of 999 species are reported to occur in Europe and species diversity can be broken down to subfamily level as follows: Archiearinae 5 species, putative Orthostixinae 1 species, Desmobathrinae 4 species, Geo­ metrinae 32 species, Ennominae 327 species, Sterrhinae 206 species and Larentiinae 424 species. Four new species are described in this volume: Afriberina salemae Skou & Sihvonen sp. n. from Portugal: Algarve; Dicrognophos siciliana Skou & Sihvonen sp. n. from Italy: Sicily; Peribatodes omalensis Rajaei, Hausmann & Skou, sp. n. from Greece: Crete; Tephronia theo­ philaria Hausmann sp. n. from Italy: Abruzzo. One genus is raised from synonymy to genus rank: Phyllometra Boisduval, 1840, stat. rev., raised from synonymy of Tephronia Hübner, 1825. Four new genus level synonymies are proposed: Bellachima Kemal et al. 2018 is downgraded to synonymy with Apochima Agassiz, 1847 syn. n.; Contropis Wiltshire, 1967 is downgraded to synonymy with Megametopon Alphéraky, 1892 syn. n.; Glacies Millière, [1874] 1869 is downgraded to synonymy with Psodos Treitschke, 1825 syn. n.; Rhopalognophos Wehrli, 1951 is downgraded from subgenus rank of Charissa Curtis, 1826 to synonymy of Charissa subgenus Neognophina Wehrli, 1946 syn. n. 111 new species and subspecies level synonymies, new statuses and new combinations are proposed: Gnophos faroulti Rothschild, 1914 is raised from subspecies of Odontognophos dumetata (Treitschke, 1827) to species rank, Odontognophos faroulti (Rothschild, 1914) stat. rev.; Agriopis aurantiaria subsp. lariciaria (Scholz, 1947) is downgraded to synonymy with Agriopis aurantiaria (Hübner, 1799) syn. n.; Agriopis marginaria subsp. pallidata (­Turati, 1911) is downgraded to synonymy with Agriopis marginaria (Fabricius, 1776) syn. n.; Apocheima hispidaria subsp. cottei (Oberthür, 1913) is downgraded to synonymy with 9

Apocheima hispidaria (Denis & Schiffermüller, 1775) syn. n.; Apocheima hispidaria subsp. popovi Vojnits, 1972 is downgraded to synonymy with Apocheima hispidaria (Denis & Schiffermüller, 1775) syn. n.; Apochima flabellaria subsp. marmararia (Buresch, 1915) is downgraded to synonymy with Apochima flabellaria (Heeger, 1838) syn. n.; Ascotis selenaria subsp. dianaria (Hübner, 1817) is downgraded to synonymy with Ascotis selenaria (Denis & Schiffermüller, 1775) syn. n.; Biston cognataria alexandrina Wehrli, 1941 is upgraded from synonymy under Biston betularia (Linnaeus, 1758) to a subspecies as Biston betularia alexandrina Wehrli, 1941 stat. n.; Biston strataria subsp. meridionalis (Oberthür, 1913) is downgraded to synonymy with Biston strataria (Hufnagel, 1767) syn. n.; Cala­ modes occitanaria subsp. submelanaria (Zerny, 1927) is downgraded to synonymy with Calamodes occitanaria (Duponchel, 1829) syn. n.; Gnophos obfuscata subsp. myrtillata Thunberg, 1788 is downgraded to synonymy with Gnophos obfuscata (Denis & Schiffermüller, 1775) syn. n.; Gnophos obfuscata subsp. androgynus Reisser, 1936 is downgraded to synonymy with Gnophos obfuscata (Denis & Schiffermüller, 1775) syn. n.; Gnophos obfuscata subsp. lutescentaria Staudinger, 1871 is downgraded to synonymy with Gnophos obfuscata (Denis & Schiffermüller, 1775) syn. n.; Gnophos obfuscata subsp. marsicaria Dannehl, 1933 is downgraded to synonymy with Gnophos obfuscata (Denis & Schiffermüller, 1775) syn. n.; Gnophos obfuscata subsp. nivea Schawerda, 1913 is downgraded to synonymy with Gnophos obfuscata (Denis & Schiffermüller, 1775) syn. n.; Psodos alticolaria subsp. chalybaeus Zerny, 1916 is downgraded to synonymy with Psodos alticolaria Mann, 1853 syn. n.; Psodos schwingenschussi Wehrli, 1919 is downgraded to subspecific rank of Psodos canaliculata (Hochenwarth, 1785) stat. n.; Psodos dioszeghyi Schmidt, 1930 is downgraded to subspecific rank of Psodos coracina (Esper, 1805) stat. n.; Psodos coracina subsp. transiens Wehrli, 1921 is downgraded to synonymy with Psodos coracina (Esper, 1805) syn. n.; Psodos coracina subsp. tundranoides Malicky, 1966 is downgraded to synonymy with Psodos coracina (Esper, 1805) syn. n.; Psodos noricana subsp. kusdasi Wehrli, 1945 is downgraded to synonymy with Psodos noricana Wagner, 1898 syn. n.; Psodos trepidaria werneri Schawerda, 1916 is raised from subspecies of Psodos canaliculata (Hochenw ­ arth, 1785) to species rank, Psodos werneri Schawerda, 1916 stat. n.; Charissa avilarius (Reisser, 1936) is downgraded to subspecific rank of Charissa (Costignophos) cre­ nulata (Staudinger, 1871) stat. n.; Charissa crenulata subsp. tormoi (Expósito Hermosa, 1996) is downgraded to synonymy with Charissa crenulata (Staudinger, 1871) subsp. ara­ gonensis (Zerny, 1927) syn. n.; Charissa dubitaria subsp. staudingeri (Wnukowsky, 1929) is upgraded from subspecies rank to species rank as Charissa (Eucrognophos) staudingeri (Wnukowsky, 1929) stat. n.; Charissa herbuloti Leraut, 2009 is downgraded to subspecies rank under Charissa (Kemtrognophos) ambiguata (Duponchel, 1830) stat. n.; Charissa muci­daria subsp. grisearia (Prout, 1915) is downgraded to synonymy with Charissa (Eucro­ gnophos) mucidaria (Hübner, 1799) syn. n.; Charissa mucidaria subsp. lusitanica (Mendes, 1903) is downgraded to synonymy with Charissa (Eucrognophos) mucidaria (Hübner, 1799) syn. n.; Charissa mucidaria subsp. ochracearia (Staudinger, 1901) is downgraded to synonymy with Charissa (Eucrognophos) mucidaria (Hübner, 1799) syn. n.; Charissa obscurata subsp. argillacearia (Staudinger, 1871) is downgraded to synonymy with Charissa (Charissa) obscurata (Denis & Schiffermüller, 1775) syn. n.; Charissa obscurata subsp. calceata (Staudinger, 1861) is downgraded to synonymy with Charissa (Charissa) obscurata (Denis & Schiffermüller, 1775) syn. n.; Charissa obscurata subsp. lafauryata (Oberthür, 1913) is downgraded to synonymy with Charissa (Charissa) obscurata (Denis & Schiffermüller, 1775) syn. n.; Charissa obscurata subsp. pullaria (Hübner, 1799) is downgraded to synonymy with Charissa (Charissa) obscurata (Denis & Schiffermüller, 1775) syn. n.; 10

Charissa onustaria subsp. eugonia (Wehrli, 1953) is downgraded to synonymy with Cha­ rissa (Kemtrognophos) onustaria (Herrich-Schäffer, 1852) syn. n.; Charissa pentheri subsp. petrina (Thurner, 1936) is downgraded to synonymy with Charissa (Cnestrognophos) pentheri (Rebel, 1901) syn. n.; Charissa pullata subsp. albarinata (Millière, 1859) is downgraded to synonymy with Charissa (Costignophos) pullata (Denis & Schiffermüller, 1775) syn. n.; Charissa pullata subsp. confertata (Staudinger, 1871) is downgraded to synonymy with Charissa (Costignophos) pullata (Denis & Schiffermüller, 1775) syn. n.; Charissa pullata subsp. impectinata (Guenée, 1858) is downgraded to synonymy with Charissa (Costi­ gnophos) pullata (Denis & Schiffermüller, 1775) syn. n.; Charissa pullata subsp. kovacsi (­Vojnits, 1967) is downgraded to synonymy with Charissa (Costignophos) pullata (Denis & Schiffermüller, 1775) syn. n.; Charissa pullata subsp. nubilata (Fuchs, 1872) is d ­ owngraded to synonymy with Charissa (Costignophos) pullata (Denis & Schiffermüller, 1775) syn. n.; Charissa difficilis subsp. uniformis (Staudinger, 1892) is downgraded to synonymy with Charissa (Dysgnophos) difficilis (Alphéraky, 1883) syn. n.; Charissa subsignaria (Staudinger, 1892) is downgraded to synonymy with Charissa (Eucrognophos) mucidaria (Hübner, 1799) syn. n.; Charissa ­supinaria subsp. benestriata Kovács, 1954 is downgraded to synonymy with Charissa supi­naria (Mann, 1854) syn. n.; Charissa supinaria subsp. budensis (Kovács, 1954) is downgraded to synonymy with Charissa supinaria (Mann, 1854) syn. n.; Charissa supinaria subsp. dio­szeghyi (Kovács, 1954) is downgraded to synonymy with Charissa supinaria (Mann, 1854) syn. n.; Charissa supinaria subsp. gulsensis (Wolfsberger, 1959) is downgraded to synonymy with Charissa supinaria (Mann, 1854) syn. n.; Charissa supinaria subsp. francothuringiaca (Bergmann, 1955) is downgraded to synonymy with Charissa supinaria (Mann, 1854) syn. n.; Charissa variegata subsp. cava (Vojnits, 1968) is downgraded to synonymy with Charissa (Eucrognophos) variegata (Duponchel, 1830) syn. n.; Charissa variegata subsp. rothschildi (Prout, 1929) is upgraded from subspecies rank to species rank as Charissa (Eucrognophos) rothschildi (Prout, 1929) stat. n., comb. n.; Elophos caelibaria subsp. senilaria (Fuchs, 1901) is downgraded to synonymy with Elophos caelibaria (Heydenreich, 1851) syn. n.; Chemerina caliginearia subsp. adriatica (Schwingenschuss & Wagner, 1927) is downgraded to synonymy with Chemerina caliginearia (Rambur, 1833) syn. n.; Ectropis crepuscularia subsp. laricaria (Doubleday, 1847) is downgraded to synonymy with Ectropis crepuscularia (Denis & Schiffermüller, 1775) syn. n.; Ematurga atomaria subsp. isoscelata (Scopoli, 1763) is downgraded to synonymy with Ematurga atomaria (Linnaeus, 1758) syn. n.; Ematurga atomaria subsp. alpicolaria Vorbrodt & MüllerRutz, 1917 is downgraded to synonymy with Ematurga atomaria (Linnaeus, 1758) syn. n.; Ematurga atomaria subsp. minuta Heydemann, 1925 is downgraded to synonymy with Ematurga atomaria (Linnaeus, 1758) syn. n.; Ematurga atomaria subsp. arenaria Candèze, 1926 is downgraded to synonymy with Ematurga atomaria (Linnaeus, 1758) syn. n.; Erannis ankeraria subsp. bervaensis Jablonkay, 1965 is downgraded to synonymy with Erannis ankeraria (Staudinger, 1861) syn. n.; Gnophos ambiguata subsp. tatrensis Vojnits, 1969 is downgraded to synonymy of Charissa (Kemtrognophos) ambiguata subsp. pullularia (Herrich-­Schäffer, 1852) syn. n.; Gnophos macguffini Smiles 1979 is downgraded to synonymy with Charissa (Dysgnophos) turfosaria (Wehrli, 1922) syn. n.; Gnophos certhiatus Rebel & Zerny, 1931 is downgraded to synonymy with Charissa (Trilobignophos) graecaria (Staudinger, 1871) syn. n.; Gnophos certhiatus subsp. minorasiaticus Wehrli, 1936 is downgraded to synonymy with Charissa (Trilobignophos) graecaria (Staudinger, 1871) syn. n.; Gnophos ophthalmica Lederer, 1853 is upgraded from synonymy under Charissa ambigu­ ata (­Duponchel, 1830) to a subspecies of Charissa (Kemtrognophos) ambiguata ophthalmicata (Lederer, 1853) stat. n.; Gnophos sheljuzhkoi Schawerda, 1924 is downgraded to 11

subspecies of Charissa (Kemtrognophos ambiguata subsp. sheljuzhkoi (Schawerda, 1924) stat. n.); Gnophos urmensis (Wehrli, 1953) is downgraded to subspecies of Charissa (Dysgnophos) difficilis subsp. urmensis (Wehrli, 1953) stat. rev.; Hemerophila strictaria subsp. variegata Djakonov, 1936 is downgraded to synonymy with Phthonandria strictaria (­Lederer, 1853) syn. n.; Hypomecis punctinalis subsp. grisearia (Bastelberger, 1902) is downgraded to synonymy with Hypomecis punctinalis (Scopoli, 1763) syn. n.; Kemtrognophos ambiguatus [sic!] viidaleppi Vojnits, 1975 is downgraded to synonymy of Charissa (Kemtrognophos) subsp. ophthalmicata (Lederer, 1853) syn. n.; L ­ ycia hirtaria subsp. istriana Galvagni, 1901 is downgraded to synonymy with Lycia hirtaria (Clerck, 1759) syn. n.; Lycia hirtaria subsp. uralaria Krulikovski, 1909 is downgraded to synonymy with Lycia hirtaria (Clerck, 1759) syn. n.; Lycia hirtaria subsp. diniensis O ­ berthür, 1913 is downgraded to synonymy with Lycia hirtaria (Clerck, 1759) syn. n.; Lycia hirtaria subsp. pusztae Vojnits, 1971 is downgraded to synonymy with Lycia hirtaria (Clerck, 1759) syn. n.; Lycia bulgariensis Vojnits & Mészáros, 1973 is downgraded to synonymy with Lycia hirtaria (Clerck, 1759) syn. n.; Lycia graecarius subsp. istrianus (­Staudinger, 1901) is downgraded to synonymy with Lycia graecarius (Staudinger, 1861) syn. n.; Lycia graecarius subsp. adrinarius (Buresch, 1915) is downgraded to synonymy with Lycia graecarius (Staudinger, 1861) syn. n.; Lycia isabellae subsp. strobinoi Dujardin, 1965 is downgraded to synonymy with Lycia isabellae (Harrison, 1914) syn. n.; Lycia necessaria subsp. grupchei (Carnelutti, 1986) is downgraded to synonymy with Lycia necessaria (Zeller, 1849) syn. n.; Lycia zonaria subsp. britannica (Harrison, 1912) is downgraded to synonymy with Lycia zonaria (Denis & Schiffermüller, 1775) syn. n.; Lycia zonaria subsp. rossica (Harrison, 1912) is downgraded to synonymy with Lycia zonaria (Denis & Schiffermüller, 1775) syn. n.; Lycia zonaria subsp. atlantica (Harrison, 1912) is downgraded to synonymy with Lycia zonaria (Denis & Schiffermüller, 1775) syn. n.; Menophra nycthemeraria subsp. dinicola (Wehrli, 1941) is downgraded to synonymy with Menophra nycthemeraria (Geyer, 1831) syn. n.; Nychiodes amygdalaria subsp. almensis Wehrli, 1941 is downgraded to synonymy with Nychiodes amygdalaria (Herrich-­ Schäffer, 1848) syn. n.; Nychiodes amygdalaria subsp. malatyaca Wehrli, 1941 is downgraded to synonymy with Nychiodes amygdalaria (Herrich-­Schäffer, 1848) syn. n.; Nychiodes atlanticaria Schwingenschuss, 1936 is downgraded to a subspecies of Nychiodes hispanica Wehrli, 1929 as Nychiodes hispanica atlanticaria Schwingenschuss, 1936 stat.n.; Nychiodes cuencaensis Leraut, 2009 is downgraded to synonymy with Nychiodes waltheri Wagner, 1919 syn. n.; Nychiodes waltheri subsp. osthelderi Wehrli, 1929 is downgraded to synonymy with Nychiodes waltheri Wagner, 1919 syn. n.; Nychiodes torrevinagrensis ­Expósito, 1984 is upgraded from synonymy under Nychiodes atlanticaria Schwingenschuss, 1936 to a subspecies as Nychiodes hispanica torre­vinagrensis Expósito, 1984 stat. n. and comb. n.; Nychiodes waltheri subsp. osthelderi Wehrli, 1929 is downgraded to synonymy with Nychiodes waltheri Wagner, 1919 syn. n.; Nychiodes saerdabica Wehrli, 1938 is upgraded from synonymy under Nychiodes waltheri Wagner, 1919 to a subspecies as Nychiodes waltheri saerdabica Wehrli, 1938 stat. n.; Nychiodes waltheri subsp. syriaca Wehrli, 1929 is downgraded to synonymy with Nychiodes waltheri Wagner, 1919 syn. n.; Peribatodes abstersaria (Boisduval, 1840) is downgraded to synonymy with Peribatodes perversaria (Boisduval, 1840) syn. n.; Peribatodes abstersaria subsp. montserrata (Wehrli, 1943) is downgraded to synonymy with Peribatodes perversaria (Boisduval, 1840) syn. n.; Peribatodes rhomboidaria subsp. ichnusaria (Ghiliani, 1852) is downgraded to synonymy with Peribatodes rhomboidaria (Denis & Schiffermüller, 1775) syn. n.; Peribatodes secundaria subsp. occidentaria Lempke, 1952 is downgraded to synonymy with Peribatodes ­secundaria (Denis & Schiffermüller, 1775) syn. n.; Peribatodes subflavaria (Millière, 1876) 12

is downgraded to synonymy with Peribatodes perversaria (Boisduval, 1840) syn. n.; Peri­ batodes subflavaria (Millière, 1876) is downgraded to synonymy with Peribatodes perversaria (Boisduval, 1840) syn. n.; Peribatodes umbraria subsp. syrirana (Wehrli, 1943) is downgraded to synonymy with Peribatodes umbraria (Hübner, 1809) syn. n.; Peribatodes umbraria subsp. perumbraria Leraut, 2009 is downgraded to synonymy with Peribatodes umbraria (Hübner, 1809) syn. n.; Phigalia pilosaria subsp. meridionalis (Constantini, 1916) is downgraded to synonymy with Phigalia pilosaria (Denis & Schiffermüller, 1775) syn. n.; Phigalia pilosaria subsp. prostae Leraut, 1996 is downgraded to synonymy with Phigalia pilosaria (Denis & Schiffermüller, 1775) syn. n.; Sciadia zelleraria (new combination, see below) subsp. robusta Wehrli, 1922 is downgraded to synonymy with Sciadia zelleraria (Freyer, 1836) syn. n.; Selidosema brunnearia subsp. dilucescens Wehrli, 1924 is downgraded to synonymy with Selidosema brunnearia (Villers, 1789) syn. n.; Selidosema pyrenaearia subsp. oliveireata Mabille, 1876 is downgraded to synonymy with Selidosema pyrenaearia (Boisduval, 1840) syn. n.; Selidosema parenzani Hausmann, 1993 is considered a hybrid and is downgraded to synonymy with Selidosema brunnearia (Villers, 1789) syn. n.; Selidosema taeniolaria subsp. extensaria Turati, 1919 is downgraded to synonymy with Selidosema taeniolaria (Hübner, 1813) syn. n.; Synopsia sociaria subsp. propinquaria (­Boisduval, 1840) is downgraded to synonymy with Synopsia sociaria (Hübner, 1799) syn. n.; Synopsia sociaria subsp. luridaria (Freyer, 1845) is downgraded to synonymy with Synopsia sociaria (Hübner, 1799) syn. n.; Synopsia sociaria subsp. almasa Schawerda, 1912 is downgraded to synonymy with Synopsia sociaria (Hübner, 1799) syn. n; Synopsia sociaria subsp. unitaria Prout, 1915 is downgraded to synonymy with Synopsia sociaria (Hübner, 1799) syn. n.; Tephronia gerini Leraut, 2009 is downgraded to subspecies of Tephronia codetaria subsp. gerini Leraut, 2009 stat. n.; Tephronia sicula Wehrli, 1933 is downgraded from species rank to subspecies rank as Tephronia codetaria sicula (Wehrli, 1933) stat. rev.; Tephronia tonnaria Tautel, 2015 is downgraded to synonymy with Tephronia cyrnea (Schawerda, 1932) syn. n.; Tephronia oranaria hubertcleui Leraut, 2009 is down­graded to synonmy with Eumannia oranaria subsp. cebennaria (Chrétien, 1898) syn. n. Eight genera are transferred from one tribe to another: Crocota Hübner, 1823 is transferred from Boarmiini Duponchel, 1845 to Gnophini Duponchel, 1845; Euchlaena Hübner, 1823 is transferred from Angeronini Forbes, 1948 to Gnophini Duponchel, 1845; Lytrosis Hulst, 1896 is transferred from Angeronini Forbes, 1948 to Gnophini Duponchel, 1845; Xanthotype Warren, 1894 is transferred from Angeronini Forbes, 1948 to Gnophini Duponchel, 1845; Dicrognophos Wehrli, 1951 is transferred from Gnophini to Abraxini + Cassymini + Eutoeini group, tentatively in Cassymini; Odontognophos Wehrli, 1951 is transferred from Gnophini to Abraxini + Cassymini + Eutoeini group, tentatively in Abraxini; Chondrosoma fiduciaria Anker, 1854 is transferred from Bistonini to tentative Prosopolophini Warren, 1894; Dorsispina furcicornaria Nupponen & Sihvonen, 2013 is transferred from unassigned (Ennominae) to Prosopolophini Warren, 1894. Twelve new genus combinations are proposed: Enconista unicoloraria Staudinger, 1871 is transferred from Elophos Boisduval, 1840 to Sciadia Hübner, 1822, comb. n.; Gnophos perspersata Treitschke, 1827 is transferred from Odontognophos Wehrli, 1951 (Gnophini) to Ortaliella Hausmann, 1993 (Macariini), comb. n.; Gnophos zelleraria Freyer, 1836 is transferred from Elophos Boisduval, 1840 to Sciadia Hübner, 1822, comb. n.; Selidosema modestaria Püngeler, 1914 is transferred from Contropis Wiltshire, 1967 to Megametopon Alphéraky, 1892 comb. n.; Contropis tagana Wiltshire, 1967 is transferred from Contropis Wiltshire, 1967 to Megametopon Alphéraky, 1892 and downgraded to subspecies rank as 13

Megametopon modestaria tagana (Wiltshire, 1967) comb. n., stat. n.; Synopsia ­strictaria Lederer, 1853 is transferred from Megalycinia Wehrli, 1939 to Phthonandria Warren, 1894, comb. n.; Tephronia lepraria Rebel, 1909 is transferred from Eumannia Fleticher, 1979 to Tephronia Hübner, 1825, comb. n.; Mannia psyloritaria Reisser, 1958 is transferred from Eumannia Fletcher, 1979 to Tephronia Hübner, 1825, comb. n.; Tephronia oranaria Staudinger, 1892 is transferred from Tephronia Hübner, 1825 to Eumannia Fletcher, 1979, comb. n.; Idaea culminaria Eversmann, 1843 is transferred from Tephronia Hübner, 1825 to Phyllometra Boisduval, 1840 comb. rev.; Phyllometra gracilaria Boisduval, 1840 is transferred from Tephronia Hübner, 1825 to Phyllometra Boisduval, 1840 comb. rev.; Geometra aceraria Boisduval, 1840 is transferred from Inurois Butler, 1879 to Alsophila Hübner, 1825 comb. rev. (see checklist). The following 18 name-bearing type specimens are selected, all designations being essential for an unambiguous usage of valid names of the species group for the European fauna. One neotype is designated (valid genus in parentheses): Geometra operaria Hübner, 1813 (Elophos). 17 lectotypes are designated (valid genus in parentheses): Gnophos exilis Wehrli, 1922 (Charissa (Dysgnophos)); Gnophos onustaria var. eugonia Wehrli, 1953 (Charissa (Kemtrognophos)); Gnophos obscurata bellieri Oberthür, 1913 (Charissa (Charissa)); Gnophos obscuraria (sic!) var. zeitunaria Staudinger, 1901 (Charissa (Kemtrognophos)); Gnophos pullata pyrenaica (Oberthür, 1913) (Charissa (Costignophos)); Psodos alpmaritima Wehrli, 1924 (Psodos); Psodos bentelii panticosea Wehrli, 1945 (Psodos); Psodos chalybaeus Zerny, 1916 (Psodos); Glacies coracina dioszeghyi Schmidt, 1930 (Psodos); Psodos coracina lappona Wehrli, 1921 (Psodos); Psodos coracina f. pseudonoricana Wehrli, 1921 (Psodos); Psodos coracina transiens Wehrli, 1921 (Psodos); Psodos noricana belzebuth ­Praviel, 1938 (Psodos); Psodos noricana kusdasi Wehrli, 1945 (Psodos); Psodos trepidaria f. schwingenschussi Wehrli, 1919 (Psodos); Psodos wehrlii Vorbrodt, 1918 (Psodos); Psodos trepidaria werneri Schawerda, 1916 (Psodos). Data from papers published up to 15.02.2019, that have come to our knowledge, are included in the manuscript.

14

Acknowledgements The GME book series is founded upon the knowledge of numerous background contributors in a number of countries. Their assistance is greatly appreciated, even if only those, who were directly involved in the contents of this Volume 6 can be named here. A number of colleagues in lepidopterology granted valuable help in the preparation, or were involved in the editorial process, of Volume 6. We owe them sincere thanks for constructive advice and useful comments. Especially we enjoyed the kind cooperation of: Carl Ahner, Germany: Preparation of genitalia slides, photography, graphic editing, literature search. Paula Banza, Portugal: Provision of biological and faunistic data. Dr. Evgeny Beljaev, Russia: Provision of literature, donation of material (Larerannis). Prof. Dr. Stoyan Beshkov, Bulgaria: Provision of intensive biological and intensive faunistic data, loan and donation of material. Cooperation in DNA barcoding. Julian Bittermann, Germany: Loan of material, provision of faunistic data (e.g. Kuchleria menadiara). Daniel Bolt, Switzerland: Provision of intensive faunstic data. Rolf Blaesius, Germany: Donation of material, provision of faunistic and biological data. Dr. Gunnar Brehm, Germany: Provision of photographs and taxonomic information. Dr. Yuri I. Budashkin, Crimea: Provision of biological data. Rainer Busse, Germany: Provision of faunistic data, donation of material, cooperation in DNA barcoding. Dr. Feza Can, Turkey: Provision of intensive faunistic and biological data. Provision of literature, loan of specimens. Cooperation in DNA barcoding. Dr. Omar Cao, Italy: Provision of faunistic data, literature. Martin Česánek, Slovakia: Provision of material for DNA barcoding (Psodos). John Chainey, United Kingdom: Access to material, taxonomic discussions and support. Type investigations at the NHMUK. Dr. Sei-Woong Choi, South Korea: Access to material, provision of information and support. Julian H. Clarke, United Kingdom: Provision of photographs and information, loan of material and support. Martin Corley, United Kingdom: Provision of literature and intensive faunistic and biological data (Portugal). Linguistic editor, constructive revision of the manuscript with many annotations and additions to the manuscript, validity check of botanical and parasitoid names. Dr. Jorge Dantart, Spain: Provision of taxonomic and distribution information, advice and support. Helmut Deutsch, Austria: Providing material (Psodos). Clarissa Dieterle, Germany: Preparation of genitalia slides, photography, graphic editing, literature search, substantial assistance in preparing the text on Charissa, subgenus Kemtrognophos. Thomas Drechsel, Germany: Provision of faunistic data, food-plant records and rearing experience. Josepha Erlacher, Germany: Assistance in writing the text on Gnophos, preparation of genitalia slides, photography, graphic editing, discussions, inspirations and accompaniment on numerous field excursions. Dr. Marianne Espeland, Germany: Access to collections and support. 15

Andrès Expósito Hermosa, Spain: Provision of faunistic data (Spain), loan of material, taxonomic discussions (Myinodes, Kuchleria). Dr. Ralf Fiebig, Germany: Provision of biological and distribution information. Heinz Fischer, Germany: Provision of records and pictures of Selidosema erebaria from the island of Pantelleria, donation of material and literature. Gabriele Fiumi, Italy: Provision of faunistic data, literature, discussions. Dr. Claudio Flamigni, Italy: Provision of faunistic and taxonomic data, literature, discussions. Loan of material. Egbert Friedrich, Germany: Loan and donation of material, provision of intensive faunistic and biological data. Cooperation in DNA barcoding. Dieter Fritsch, Germany: Loan and donation of material, provision of intensive biological and faunistic data (Samos etc), cooperation in DNA barcoding, discussions (e.g. on Idaea textaria). Sabine Gaal, Austria: Provision of information, advice and support. Javier Gastón, Spain: Provision of faunistic data (Spain), literature, taxonomic discussions (e.g. on Idaea humiliata davidi). Dr. Jörg Gelbrecht, Germany: Provision of many faunistic data (e.g. Turkey), of numerous food-plant records, and of literature, loan of material, taxonomic discussions, extensive and constructive comments on the manuscript, revision of maps. Cooperation in DNA barcoding. Alessandro Giusti, United Kingdom: Access to material, provision of photographs, information and support. Barry Goater, United Kingdom: Discussion of taxonomic matters. Dr. Stanislav Gomboc, Slovenia: Provision of intensive biological and faunistic data, donation of material (Hypomecis). Guido Govi, Italy: Provision of faunistic data, literature, discussions. David Grundy, United Kingdom: Provision of taxonomic and distribution information. Penny Hale, Spain: Provision of faunistic and biological data (southern Spain; e.g. on Scopula donovani), cooperation in DNA barcoding. Jahne Harmeling, Germany: Preparation of genitalia slides, photography, graphic editing, literature search. Prof. Dr. Gerhard Haszprunar, Germany: Support and encouragement of research on the GME series as the director of the ZSM. Dr. Paul Hebert, Canada: Kind cooperation in DNA barcoding, initiation of the global geometrid campaign of DNA barcoding, sequencing of >21,000 geometrid species, covering almost all species of this volume. Bob Heckford, United Kingdom: Access to material, provision of information, literature and support. Dr. Maria Heikkilä, Finland: Provision of taxonomic information and advice. Claude Herbulot †, France: Kind support during the author’s visits (AH), release of his entire collection (with many types), provision of literature and faunistic data, taxonomic discussions. Dr. Axel Hille, Germany: Technical support in DNA barcoding (Charissa, Psodos, Gnophos). Chantal Holzhause, Germany: Photography of several preparations. Martin Honey, United Kingdom: Kind support during the author’s visits (AH) at the NHMUK, loan of material, provision of biological and faunistic data (Balearics). Dr. Mike Hull, United Kingdom: Provision of faunistic data. 16

Dr. Peter Huemer, Austria: Provision of faunistic data, literature, scientific discussions, provision of literature, loan of material. Extensive cooperation in DNA barcoding. Dr. Ties Huigens, The Netherlands: Providing permission to use Dutch distribution data gathered by De Vlinderstichting. Dr. Marco Infusino, Italy: Kind support during the author’s visits (AH), provision of faunistic data (Sicily), loan and donation of material. Cooperation in DNA barcoding. Urmas Jürivete, Estonia: Provision of intensive faunistic data. Claudia Junghans, Germany: Preparation of genitalia slides, photography, graphic editing, literature search, assistance in preparing the text on Psodos. Jari Junnilainen, Finland: Provision of distributional data, access to material, donation of material, advice and support. Dr. Lauri Kaila, Finland: Access to material, loan of material, advice and support. Ole Karsholt, Denmark: Numerous discussions related to Ennominae, access to the ZMUC collection, loan of material, provision of taxonomic information and advice. Sung-Soo Kim, South Korea: Access to material, provision of information and support. Dr. Gareth King, Spain: Provision of intensive food-plant data from Spain. Donation of material. Ulrike Kleinkamp, Germany: Access to collections and support. Igor Kostjuk, Ukraine: Provision of many literature data (e.g. all records from Ukraine, revision of maps for eastern Europe), loan of material, faunistic and taxonomic discussions. Photography and preparation of colour plates. Donation of material. Dr. Bernard Landry, Switzerland: Provision of taxonomic information. Thomas Lange, Germany: Provision of material (Dasypteroma thaumasia). Dr. Zdeneˇk Las˘tu˚ vka, Czech Republic: Provision of material, literature, intensive faunistic and biological data. Michael Leipnitz, Germany: Provision of intensive biological and distributional data, loan and donation of material, provision of literature and photographs. Dr. Antoine Lévêque, France: provision of faunistic data, cooperation in DNA barcoding, provision of photographs of Idaea barbuti. Dr. Kyra von der Lippe, Germany: Accompaniment on numerous field excursions (Psodos). Tina Llera Pérez, Germany: Photography, graphic editing, literature search. Dr. Hans Löbel, Germany: Provision of faunistic and biological data. Kind phototechnical help. Taxonomic discussions. Maurizio Lupi, Italy: Provision of intensive faunistic data. Dr. Adam Malkiewicz, Poland: Provision of intensive faunistic data. Tobias Malm, Sweden: Provision of taxonomic information and photos. Eduardo Marabuto, Portugal: Provision of biological and distributional data. Geoff Martin, United Kingdom: Loan of material and provision of access to collections under his care. Dr. Reinhard Maschler, Germany: Provision of faunistic data and food-plant records (NW Germany). Josephine Matzig, Germany: Technical support in preparation of DNA barcoding (Charissa). Dr. Robert Mazel, France: Provision of literature and faunistic data, donation of his collection. Paolo Mazzei, Italy: Provision of biological data. Dr. Wolfram Mey, Germany: Kind support during the author’s visit (BM and AH) to the MNHU, loan of material including type specimens. Dr. Iva Mihoci, Croatia: Provision of faunistic data (Croatia). Cooperation in DNA bar­coding. 17

Dr. Joël Minet, France: Access to collections and support. Dr. Vladimir Mironov, Russia: Provision of intensive faunistic and biological data (mainly Russia), loan of material including type specimens, taxonomic discussions. Control of the manuscript concerning new Perizoma and Eupithecia species for Europe. Tomas Molina, Spain: Provision of faunistic and biological data. Rando Müller, Germany: Provision of faunistic data and material (Charissa). Leidys Murillo-Ramos, Colombia/Sweden: Provision of phylogenetic information. Dr. Marko Mutanen, Finland: Provision of faunistic data, literature, scientific discussions (e.g. on the status of Ectropis crepuscularia). Extensive cooperation in DNA barcoding. Dr. Hideo Nakajima, Japan: Access to material, provision of information and support. Kari Nupponen, Finland: Provision of intensive biological, faunistic data and taxonomic and distribution information, loan of material and scientific discussions. Dr. Zuhal Okyar, Turkey: Provision of literature and faunistic data (European Turkey). Prof. Dr. Antonio Ortiz Cervantes, Spain: Kind cooperation in DNA barcoding, provision of faunistic data (Spain; e.g. on Idaea lobaria). Prof. Dr. Paolo Parenzan, Italy: Provision of faunistic data, and literature (e.g. on Kuchleria menadiara), discussion of faunistic and taxonomical matters. Dr. Mark Parsons, United Kingdom: Provision of faunistic and biological data and literature. Gergely Petrány, Hungary: Provision of faunistic data, cooperation in DNA barcoding. Dr. Willy de Prins, Belgium: Providing access to Belgian distributional data, provision of literature. Norbert Pöll, Austria: Provision of intensive biological and faunistic data, loan of material. Cooperation in DNA barcoding. Prof. Dr. László Rákosy, Romania: Taxonomic and faunistic discussions and support. Victor Redondo, Spain: Provision of literature and faunistic data, taxonomic discussions (e.g. on Idaea humiliata davidi), advise and support. Dr. Ladislaus Reser, Switzerland: Provision of faunistic data and literature, loan of specimens, discussion of taxonomic problems. Dr. Hartmut Rietz, Germany: Provision of intensive faunistic data, loan and donation of material, provision of faunistic and biological data (Spain, Italy; e.g. Perizoma barrassoi). Dr. Laszlo Ronkay, Hungary: Loan of material, provision of faunistic data (Tephronia lepraria). Dr. Frank Rosenbauer, Germany: Provision of faunistic data (Biston), Cooperation in DNA barcoding. Dr. Walter Ruckdeschel, Germany: Provision of extensive faunistic data from Crete, donation of material. Marcello Romano, Italy: Provision of biological data. Prof. Dr. Nils Ryrholm, Sweden: Provision of biological and intensive faunistic data. Heinz Salpeter, Germany: Provision of faunistic data. Kind help in computer problems. Lena Sassenberg, Germany: Photography, graphic editing and literature search. Vladimir Savchuk, Crimea: Provision of biological and faunistic data, provision of photographs and specimens of Ithysia pravata. Dr. Nikolay Savenkovs, Latvia: Provision of intensive faunistic data. Dr. Stefano Scalercio, Italy: Kind support during the author’s visit (AH), provision of many faunistic data (Italy), loan and donation of material. Provision of photographs for Nothocasis rosariae. Cooperation in DNA barcoding. Bernd Schacht, Germany: Provision of material (Charissa).

18

Franziska Schäfer, Germany: Preparation of genitalia slides, photography, graphic editing, literature search. Dr. B. Christian Schmidt, Canada: Provision of material (Charissa macguffini). Dr. Andreas H. Segerer, Germany: Provision of faunistic data (Germany), extensive cooperation in DNA barcoding. Christian Siegel, Italy: Provision of biological data. Harri Sihvonen, Finland: Provision of distribution information. Guy Sircoulomb, France: Provision of material (Charissa). Bjarne Skule, Denmark: Provision of faunistic data. Manfred Sommerer, Germany: Provision of faunistic data, loan and donation of extensive material. Much advice on many details. Cooperation in DNA barcoding. Adrian Spalding, United Kingdom: Provision of literature. Dirk Stadie, Germany: Provision of biological, taxonomic and faunistic data, loan and donation of material, advice and support. Hermann Staude, South Africa: Provision of taxonomic and distribution information, advice and support. Andreas Stübner †, Germany: Provision of faunistic data. Dr. Dieter Stüning, Germany: Extensive and in-depth advice on numerous taxonomic questions, particularly on genus systematics of Synopsia, Megalycinia, Phthonandria, ­Wehrliola and Dasypteroma, and data sharing and support. Kind support during our visits (BM and AH) at the ZFMK, information on type material in ZFMK, loan of material, provision of literature. Cooperation in DNA barcoding. Reinhard Sutter, Germany: Donation of material. Gidrius Svitra, Lithuania: Providing access to Lithuanian distribution data. Provision of intensive faunistic data for Belarus. Dr. Levente Székely, Romania: Provision of biological and intensive faunistic data. Dr. Kálmán Szeôke, Hungary: Donation of material (Peribatodes). Kornél Takáts, Hungary: Provision of biological data. Prof. Dr. Gerhard Tarmann, Austria: Providing financial support for DNA barcoding of Psodos. Prof. Dr. Toomas Tammaru, Estonia: Provision of literature, taxonomic discussions. Claude Tautel, France: Provision of biological and intensive faunistical data, loan and donation of material, provision of literature, taxonomic discussion (e.g. on Tephronia tonnara and Perizoma juracolaria). Mike Taylor, United Kingdom: Provision of faunistic data. Balázs Tóth, Hungary: Provision of biological and faunistical data. Dr. Robert Trusch, Germany: Loan of material, taxonomic discussions, provision of literature. Kind support during visits (BM) at the SMNK, type search in the SMNK. Dr. Josep Ylla i Ullastre, Spain: Provision of intensive faunistic data and biological data, discussion of faunistic and taxonomical matters, loan and donation of material. Thierry Varenne, France: Provision of biological data. Dr. Dragan Vajgand, Serbia: Provision of intensive faunistic data. Francesc Vallhonrat, Spain: Provision of literature and faunistic data (Spain: Catalonia) incl. provision of photos of type specimens of Peribatodes ebusaria. Dr. Jaan Viidalepp, Estonia: Provision of intensive faunistic data. Taxonomic discussions. Dominic Wanke, Germany: Photography of several genitalia slides. Prof. Dr. Niklas Wahlberg, Sweden: Provision of phylogenetic information.

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Hartmut Wegner, Germany: Provision of intensive faunistic and biological data. Provision of literature. Donation and provision of literature. Dr. Michael Weidlich, Germany: Provision of literature, donation of type specimens (Eupithecia pindosata). Friedrich Weisert †, Wien (A): Provision of faunistic data, discussions (e.g. on Tephronia from Cyrenaica) Benjamin Wiesmair, Austria: Loan of material, taxonomic and distribution information. Werner Wolf, Germany: Provision of faunistic data, discussion of taxonomic problems (Charissa). Mandy Wolfram, Germany: Database management (Psodos), literature search. Dr. Norbert Zahm, Germany: Provision of biological and intensive faunistic data (Germany, central Italy). Cooperation in DNA barcoding (e.g. Perizoma barrassoi). Bernd Müller owes his wife Margit and his son Ronny sincere thanks for their great understanding and support during the many years of his time-consuming work on this book. Sven Erlacher would like to thank his wife Josepha and his children Vera, Robert, Clara, Meta and Matti for their understanding, patience and all kinds of friendliness during the work on this book. Axel Hausmann thanks his wife Silvia and the children, who encouraged and supported him, though suffering from the 25 years-long period of the preparation of this book series. Hossein Rajaei deeply thanks his wife Maryam Boroomandi, for her emotional and mental support, encouragement and thoughtfulness not only during this project, but in whole last thirteen years. Pasi Sihvonen wishes to thank his wife Karoliina Reunanen, and our children Eero and Siiri, for their lasting understanding and patience during evenings, weekends and holidays when this book was prepared. Peder Skou wishes to thank his companion Anne-Grete Klausen for showing a great understanding for his deep passion for the Geometrids and for spending so much time on these creatures.

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Introduction Volume 6 of the book series The Geometrid Moths of Europe is the second and concluding part of the revision of the European taxa of the subfamily Ennominae, covering the Gno­ phini (part) and the Boarmiini. Furthermore, it deals with the taxa discovered in Europe after publication of previous volumes and presents a complete checklist of the European and adjacent regions’ Geometridae fauna. Volume 5 (Skou & Sihvonen 2015) presented the first part of the revision, covering 141 Ennominae species in the tribes Abraxini, Apeirini, Baptini, Caberini, Campaeini, Cassymini, Colotoini (currently considered a junior synonym of Prosopolophini), Ennomini, Epionini, Gnophini (part), Hypochrosini, Lithinini, Macariini, Theriini, and 34 species of uncertain tribal association. Introduction to the subfamily Ennominae, including a review of its internal taxonomy and review of tribes, monophyly, its systematic position within the Geometridae, diversity and ecology are provided in vol. 5 (Skou & Sihvonen 2015). An extensive multi-gene phylogenetic study of the Geometroidea is under preparation (Murillo-Ramos et al. 2019). Results of this analysis have been largely included and commented in the Checklist, including those that concern the Ennominae. Molecular studies have brought great advances to numerous disciplines in biology, including phylogenetic systematics and taxonomy. The latter has been revolutionized by the incorporation of DNA barcodes (a 658bp part of the mitochondrial COI gene) into the taxonomic process (Hebert et al. 2003) and the related analytical tools as well, available on Barcode of Life Data Systems database (BOLD, http://www.boldsystems.org/). This is true also for the Geometrid Moths of Europe series, where a separate Genetic Data chapter has been included for each species in the last two volumes (Hausmann & Viidalepp 2012; Skou & Sihvonen 2015). For the species of Volumes 1 and 2 the DNA barcoding data were subsequently provided by Hausmann et al. (2013). In the context of European Geometridae, molecular advances have largely focused on species-level research questions (for instance Huemer & Hausmann 2009; Sihvonen et al. 2014; Huemer & Mayr 2015), or on analyses of massive data sets (for instance Hausmann et al. 2011b; Hausmann et al. 2013; Mutanen et al. 2012), but the classification of species into genera and tribes and their relationships have received less attention. The need for such studies, and integrative approaches, was pronounced by Skou & Sihvonen (2015). We were still facing a similar problem, when this volume was prepared. Taxonomy is becoming more accurate than ever, but combination of species into higher taxonomical categories is still somewhat arbitrary and descriptive. Inclusion of extensive DNA materials (including nuclear DNA data) into phylogenetic studies is in great demand, with the ultimate aim of creating more rigorous taxonomical hypotheses, eventually leading to more stable classifications. Volume 6 deals with several taxonomically difficult genera and species complexes. These include for instance Charissa Curtis, 1826, Lycia Hübner, 1825, Nychiodes Lederer, 1843, Peribatodes Wehrli, 1943, Psodos Treitschke, 1825, Sciadia Hübner, 1822, Selidosema Hübner, 1823 and Tephronia Hübner, 1825. The study of these, together with almost all taxa treated in this volume, has benefited from the inclusion of DNA barcodes as an additional source of taxonomic information.

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Methodological notes The general methodological aspects have been given in The Geometrid Moths of Europe, vol. 1 (Hausmann 2001) and for the Ennominae in vol. 5 (Skou & Sihvonen 2015). Terminology of the wing pattern follows Hausmann (2001, see also Skou & Sihvonen 2015: Text-fig. 10) and wing venation (see Skou & Sihvonen 2015: Text-figs 11, 12) largely follows that of McGuffin (1977). Terminology for the male and female genitalia follows Klots (1970) and Kristensen (2003) and in cases, where homologies of structures were difficult to identify, descriptive terms were used instead. As was explained in vol. 5 of The Geometrid Moths of Europe (Skou & Sihvonen 2015), genus and tribe level classifications of the Geometridae are currently yet incompletely resolved and most taxa are not analyzed in a phylogenetic context. This concerns the European fauna also. Against this background, we have made the decision to arrange the higher taxa largely, but not solely, according to conventional views. The order, therefore, does not necessarily reflect the phylogenetic relationships of the taxa. As a basis for the arrangement of genera and tribes in this monograph, the Distributional Checklist of European Geometridae by Müller (1996) has been used. This work is complemented with our findings observed during preparation of the book, and extensive consultation of recent country and regional check-lists, taxon-specific publications and publications such as Holloway (1994; 1996; 2011), Viidalepp et al. (2007), Pitkin (2002), Ounap et al. (2011), Beljaev (2006a; 2006b; 2008), Redondo et al. (2009), Sihvonen et al. (2011), Flamigni et al. (2007; 2016), Jiang et al. (2017), and particularly the extensive multi-gene analysis, which was available to us prior to its publication (Murillo-Ramos et al. 2019). The specimens shown in the adult plates were photographed individually, removed from the background and edited in Adobe Photoshop. The single images were compiled into plates with Photoshop and drop shadows were added to final plates. Numerous photos of differential external features shown in text figures were modified from Igor Kostjuk’s plate photographs. Photos in text figures are often not in scale, because the objective was to illustrate diagnostic features. The genitalia and abdomen were prepared following standard methods, see for instance Hardwick (1950). In numerous specimens the male aedeagus is shown both as vesica uneverted, to allow comparison with earlier literature, and as vesica everted. The vesica of species with small aedeagus was everted via the caecum that was cut open by placing the aedeagus inside a hypodermic syringe (Sihvonen 2001), or in species with large aedeagus, by following the method described by Lafontaine & Mikkola (1998). Numerous structures shown in the genitalia plates were photographed in 2–6 images of different depth of ­focus and combined into single images using image-stacking software such as CombineZP (­Hadley 2012), Helicon Focus and Photoshop. Original images were cleaned, removed from the background, edited and compiled into plates with Adobe Photoshop (CS6) or with Corel Draw. Magnified parts of genitalia figures are not in scale. For the DNA analyses, one or two legs were removed from each dried specimen and stored in an individual tube, which in most cases contained ethanol. DNA extraction, amplification, and sequencing of the barcode region of the mitochondrial cytochrome c ­oxidase I (COI) gene (658 base pairs) were carried out in the Canadian Centre for DNA Bar­coding, Ontario, Canada, using standard high throughput protocols (Ivanova et al. 2006; deWaard et al. 2008), those are described at CCDB (2013). Sequence divergence within and 23

­between species was calculated using the Kimura 2–parameter model (Kimura 1980) and the neighbour-joining algorithm (Saitou & Nei 1987), as implemented in BOLD (http:// www.boldsystems.org/). The Barcode Index Number (BIN), an algorithm of BOLD database (­Ratnasingham & Hebert 2007), was used to verify (sub-)species identifications since the clusters have been shown to perform high concordance with (sub-)species when combined with a morphological analysis (Ratnasingham & Hebert 2013; Hausmann et al. 2013). The genus and species-group names largely follow Scoble (1999) and more recent literature as indicated in appropriate chapters. The genus concepts and diagnoses are largely based on Palaearctic species unless otherwise indicated. For each species a full synonymy of species-group names is given including the original combination and reference. As in the Geometrid Moths of Europe, vol. 3 (Hausmann & Viidalepp 2012), less emphasis is given to nomenclaturally unavailable names at infrasubspecific rank, as there are too many names in existence. Distribution maps were prepared by compiling data from collection specimens. In taxonomically challenging groups identities were verified further from the genitalia, DNA barcodes or other relevant information. These data were supplemented by adding data from recent faunistic inventories of United Kingdom (Hill et al. 2010), Norway (Aarvik et al. 2000), Finland (Huldén et al. 2000; Kullberg et al. 2001), the Netherlands (Anonymous 2018), Austria (Huemer et al. 2009; Kerschbaum & Pöll 2010), Spain (Redondo et al. 2009), Italy (Flamigni et al. 2007; 2016) and south-western Germany (Ebert 2001). These maps were further supplemented by sending out the blank maps of the majority of species to numerous experienced lepidopterists across Europe. Data from these sources were drawn together, quality controlled by the authors, and final maps were sent out one more time to the same lepidopterists for approval and for additional data. Distribution maps show the hypothetical resident distribution area (grey), combined with localities from which specimens have been examined (black dots). Unless otherwise mentioned in the maps, old records are marked with open dots, when the species has probably disappeared from that locality. Doubtful records are marked with question marks and explained in the text. Information on larval food-plants and on habitats is based on the authors’ own original data, literature data, and personal communications from numerous experienced lepido­ pterists. For the first-hand verified feeding observations from nature, we use the expression “found on” while for literature data we use “recorded on” and for feeding observations in captivity we used “reared on”. Vertical distribution refers mainly to the analysis of collection label data, with personal communications and literature data used as an additional source. Phenological information is mainly based on collection label data, while literature data are used as an additional source. It is important to note that the flight period may vary greatly annually and geographically. Therefore single outlying records are usually omitted, while we tried to consider and present regional differences in the flight period and number of generations. For practical reasons, and to avoid duplication during preparation, the following division of labour was established. –– Bernd Müller covered all genera of this volume except those mentioned under the sections of the other co-authors. For these genera, he also dissected the genitalia, prepared the text-figures and tables and edited distribution maps. –– Sven Erlacher covered genera Charissa (partly), Gnophos, and Psodos (with Hossein Rajaei). For these genera, he also dissected and photographed the genitalia, prepared 24

––

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(with Hossein Rajaei) the text-figures and tables and edited distribution maps. In addition he provided, for the above mentioned genera, photographs of the wing pattern and genitalia for the plates. Axel Hausmann covered genera Tephronia, Eumannia, Phyllometra, Ithysia and Charissa subgenus Kemtrognophos (based on information provided by Sven Erlacher) as well as all new species recorded for Europe after publication of the first five volumes of the series. For these genera and species, he also dissected the genitalia, prepared the text-­ figures and tables, prepared and edited distribution maps and selected specimens for adult plates. In addition he reviewed, corrected, supplemented, unified and edited the whole text, performing linguistic and stylistic streamlining. He analyzed the DNA data and wrote the Genetic Data texts for the whole volume, and compiled the complete checklist of European Geometridae (Vols 1–6). Hossein Rajaei covered genera Nychiodes, Peribatodes and Psodos (with Sven ­Erlacher). For these genera, he also dissected and photographed the genitalia, prepared the text-­ figures and tables and edited distribution maps. In addition he edited the distribution maps for the parts of Bernd Müller and Sven Erlacher. Rajaei photographed the genitalia slides for the parts of Bernd Müller and Axel Hausmann, and compiled the male and female genitalia plates for the whole volume. In addition he prepared or edited text-figures for the whole volume, reviewed, corrected and edited several parts of the manuscript. Pasi Sihvonen covered genera (Afriberina, Crocota, Dicrognophos, Ekboarmia, Elophos, Megalycinia, Odontognophos, Ortaliella, Phthonandria, Pseudocoremia, Sciadia and Yezognophos, including the genus chapters and under species accounts paragraphs on taxonomy, external characters and abdomen, variation, male genitalia, female genitalia, similar species and remarks where relevant. For these genera, he also dissected the genitalia, prepared the genitalia plates, text-figures and tables and edited distribution maps. In addition, Sihvonen wrote the text on Charissa subgenus Neognophina (based on information provided by Sven Erlacher), the Introduction, Methodological notes and under Systematic Account the tribus chapters and prepared the associated text-figures. He also compiled the Abstract, References, Appendices 1 and 2 and reviewed, corrected and edited several parts of the manuscript and helped to compile the checklist of European Ennominae (Vol. 6). Peder Skou wrote the paragraphs on distribution and abundance, phenology, biology and habitat for all genera covered by Bernd Müller and for genera Afriberina, Crocota, Dicrognophos, Ekboarmia, Elophos, Megalycinia, Odontognophos, Ortaliella, Phthonandria, Pseudocoremia, Sciadia, Charissa subgenus Neognophina and Yezognophos. Skou also selected specimens for adult plates, made the disposition for and helped to compile colour plates, compiled Acknowledgments, Abbreviations, compiled distributional data and extrapolated the distribution areas for all distribution maps except the taxa covered by Axel Hausmann.

Igor Kostjuk photographed nearly all adult specimens and arranged the numerous individual photographs into the final adult colour plates.

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Abbreviations Beshkov Collection of Stoyan Beshkov in National Museum of Natural History, ­Sofia, Bulgaria (NMNH) Clarke Private collection of Julian Clarke, Crawley Down, United Kingdom Colomb Private collection of Claude Colomb, Saint-Etienne, France CREA-SAM Collection of Consiglio per la ricerca in agricoltura e l’analisi dell’economia agraria – Unità di ricerca per la selvicoltura in ambiente mediterranea, Rende, Italy Erlacher Private collection of Sven Erlacher, Chemnitz, Germany Expósito Private collection of Andrés Expósito Hermosa, Madrid, Spain Fiebig Private collection of Ralf Fiebig, Roßleben, Germany Fischer Private collection of Heinz Fischer, Weissach, Germany Flamigni Private collection of Claudio Flamigni, Forlì, Italy Friedrich Private collection of Egbert Friedrich, Jena, Germany Fritsch Private collection of Dieter Fritsch, Lörrach, Germany Gaston Private collection of Javier Gaston, Getxo Bizkaia, Spain Gelbrecht Private collection of Jörg Gelbrecht, Königs Wusterhausen, Germany Govi Private collection of Guido Govi, Forlì, Italy Junnilainen Private collection of Jari Junnilainen, Vantaa, Finland Kaitila Private collection of Jari Kaitila, Vantaa, Finland Leipnitz Private collection of Michael Leipnitz, Stuttgart, Germany Löbel Private collection of Hans Löbel, Sonderhausen, Germany May Private collection of Bernhard May, München, Germany MNHN Musée National d’Histoire Naturelle, Paris, France MNHU Museum für Naturkunde – Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin, Germany B. Müller Private collection of Bernd Müller, Berlin, Germany R. Müller Private collection of Rando Müller, Jena, Germany NHMUK Natural History Museum, London, United Kingdom NHMW Naturhistorisches Museum, Wien, Austria NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden NML Natur-Museum Luzern, Switzerland NSMW Naturhistorische Sammlungen Museum Wiesbaden, Germany Nupponen Private collection of Kari and Timo Nupponen, Espoo, Finland Pöll Private collection of Norbert Pöll, Ischl, Austria Schacht Private collection of Bernd Schacht, Dahlewitz, Germany Sihvonen Private collection of Pasi Sihvonen, Veikkola, Finland Sircoulomb Private collection of Guy Sircoulomb, Anquetierville, France Skou Private collection of Peder Skou, Vester Skerninge, Denmark Sommerer Private collection of Manfred Sommerer, München, Germany SMNH State Museum of Natural History, Lviv, Ukraine SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany Stadie Private collection of Dirk Stadie, Lutherstadt Eisleben, Germany Tautel Private collection of Claude Tautel, Paris, France TLMF Tiroler Landesmuseum Ferdinandeum, Naturwissenschaftliche Samm­ lungen, Hall, Austria TMB Hungarian Natural History Museum, Budapest, Hungary 27

Wegner Weidlich ZISP ZFMK ZMH ZMKU ZMUC ZSM

Private collection of Hartmut Wegner, Adendorf, Germany Private collection of Michael Weidlich, Neißemünde, Germany Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn, Germany Finnish Museum of Natural History, University of Helsinki, Finland Zoological Museum of Kiev national Shevchenko University, Ukraine Zoological Museum of the University, Copenhagen, Denmark Zoologische Staatssammlung, München (SNSB), Germany

‡ name not nomenclaturally available under the ICZN ab. aberration BIN Barcode Index Number BOLD Barcode of Life Data Systems coll. collection e.o. ex ovo (reared from egg) e.l. ex larva (reared from larva) f. form leg. legit (collected by) pers. comm. personal communication var. variation

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Systematic Account Subfamily Ennominae Duponchel, 1845 (continued from volume 5) Abraxini + Cassymini + Eutoeini group of tribes (part): Addition (other species treated in volume 5) The tribes Abraxini, Cassymini and Eutoeini are morphologically similar (see Skou & Sih­ vonen 2015 for details) and their close relationship is supported by molecular data also (Sihvonen et al. 2011). An extensive molecular study on the Geometroidea phylogeny is under preparation and preliminary results continue to support the relationships of the three mentioned Ennominae groups including the genus Odontognophos Wehrli, 1951 (­Murillo-Ramos et al. 2019) which is tentatively placed here in the tribe Abraxini. Morphology of the genus Dicrognophos Wehrli, 1951 also supports association with the mentioned group, tribe Cassymini (it has thus far not been included in a molecular study). Both genera are transferred here from the Gnophini (e.g. Hausmann et al. 2004; 2011a) to the Abraxini + Cassymini + Eutoeini group. Odontognophos had been included in the broad concept of Macariini in Beljaev (2016). In volume 5 of the Geometrid Moths of Europe (Skou & Sihv­ onen 2015) the following genera of the mentioned tribal group were treated: Abraxas Leach, 1815, Ligdia Guenée, 1858, Lomaspilis Hübner, 1825 and Stegania Guenée, 1845. The group is diagnosed by the following characters (Holloway 1994; Skou & Sihvonen 2015): Valva divided, its dorsal arm often narrow and curved, setose apically. Male 8th sternite unmodified. Fovea in male forewing often present. Chaetosemata present. Transverse setal comb often present on male 3rd abdominal sternite. Forewing radial veins often reduced in number. Female structures variable, diagnostic characters not yet identified.

Tribus Abraxini (part): Addition (other species treated in volume 5)

Odontognophos Wehrli, 1951 (new tribus combination) Odontognophos Wehrli, 1951, Lambillionea 51: 8, 29. Type species: Gnophos dumetata T ­ reitschke, 1827. Unavailable names (misspelling): Odontignophos: Wehrli (1951: 8), Odonthognophos: Vives Moreno (1994).

Diversity and distribution: Odontognophos contains three species in the western Palaearctic region. External characters and abdomen: Rather large moths, wingspan 33−39 mm. Wings light grey to brown with faint blackish medial lines. Underside of wings almost devoid of markings. Wing termen undulating, particularly on hindwings. Male and female antennae filiform. Hindlegs in both sexes with 2+2 spurs, male hindleg slightly swollen with narrow hair pencil. Abdominal sternites unmodified. 29

Male genitalia: Uncus setose. Socii absent. Gnathos well-developed, medial element upturned. Valva divided, ventral arm with spinose apex, dorsal arm setose. Ventral part of ­juxta simple, elongated, sclerotized. Dorsal part of juxta with two shallow concavities. Saccus round. Aedeagus with sclerotized ridges. Vesica base covered with microcornuti. Female genitalia: Papillae anales elongated. Lamella postvaginalis and antevaginalis fused, massive, strongly sclerotized. Corpus bursae elongated, membranous. Signum elongated plate with horizontal ridge, located at anterior end of corpus bursae. Biology, habitat, phenology: Larva of O. dumetata recorded on Rhamnus spp., including R. alaternus, R. saxatilis, reared on R. catharticus (Elliot & Skinner 1995; Flamigni et al. 2016). Erroneously recorded on Phillyrea latifolia Millière (1858–1864) (Flamigni et al. 2016). Pupation in a fine loose web above ground (M. Leipnitz pers. comm.). O. dumetata inhabits xerothermic, rocky and bushy habitats, preferring limestone substrate (Patočka & Turčani 2005), clearings of karst woods and dry oak forests (Vojnits 1967), occurring from 70 up to 1850 m (Flamigni et al. 2016). Adults from August to October overwintering as egg (Patočka & Turčani 2005; Redondo et al. 2009; Flamigni et al. 2016). Immature stages: Larva of O. dumetata light to dark fleshy grey with vague longitudinal striations, dorsal line prominent and blackish on segments 1−3 and 9−11. Well camouflaged on its resting background, a Rhamnus twig (Elliot & Skinner 1995). Pupa cremaster of O. dumetata elongated and bifurcate (Patočka & Turčani 2005). Remarks: Earlier studies did not reach a consensus which species should be combined with genus Odontognophos. Scoble (1999) included three species: O. dumetata, O. perspersata and O. zacharia. Other authors have combined some of the above-mentioned species with Gnopharmia Staudinger, 1892 (Leraut 2009), Odontognophos subgenus Dicrognophos (Vives Moreno 1994), and Gnophos subgenus Dicrognophos (Gómez de Aizpúrua 2002). Leraut (2009) included one additional species in the genus: O. sartata. Recently Odontognophos has been classified in the Gnophini (Vives Moreno 1994; Viidalepp 1996; Hausmann et al. 2004; 2011a), in the Boarmiini (Patočka 2003) and in the broad concept of Macariini in Beljaev (2016). The divided valva, setose uncus in the male genitalia and bifurcate pupa cremaster are similar to the equivalents in the Abraxini + Cassymini + Eutoeini group (Patočka & Turčani 2005), but elongated signum with horizontal ridge is similar to the Boarmiini, which is sister group of the Abraxini + Cassymini + Eutoeini group (see for instance Aethalura McDunnough, 1920 and Peribatodes Wehrli, 1943 in this volume). In an extensive molecular phylogeny analysis genus Odontognophos groups within the Abraxini + Cassymini + Eutoeini group (Murillo-Ramos et al. 2019). Systematic position: Abraxini + Cassymini + Eutoeini group, tentatively in Abraxini.

1. Odontognophos dumetata (Treitschke, 1827) Gnophos dumetata Treitschke, 1827: Schmett. Eur. 6 (1): 163 ([Yugoslavia]: Dalmatia). Lecto­ type ♂, paralectotypes 2♀ (designated by Vojnits (1967), in Hungarian Natural History Museum, Budapest, Hungary, examined externally). Treitschke (1827) reports that he has received this species from Dalmatia and probably from southern France. The latter location is doubtful, therefore any material from this locality should be excluded from the type series (ICZN 1999, Article 72.4.1). Ennomos daubearia Boisduval, 1840: Genera Index meth. eur. Lepid: 183 ([France: perhaps Montpellier]). Syntypes. Scoble (1999) gives Corsica as type locality, but according to current

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knowledge dumetata does not occur on that island. Valid at subspecific rank (Scoble 1999) or considered a form of dumetata Treitschke, 1827 (Leraut 2009). See Variation. Ennomos temperata Eversmann, 1846: Bull. Soc. imp. Nat. Moscou 19 (2): 89, pl. 2, fig. 1 (Russia: Simbirsciensi). Syntypes. Junior synonym of dumetata Treitschke, 1827 (Scoble 1999). Gnophos dumetata var. saturata Fuchs, 1902: Jb. Nassau. Ver. Naturk. 55: 73 (Hungary). Syntypes. Junior synonym of dumetata Treitschke, 1827 (Vojnits 1967). Gnophos dumetata var. scopulata Fuchs, 1889: Jb. Nassau. Ver. Naturk. 42: 221 ([probably Switzerland/ Germany]: Rhine mountains). Syntypes 2♀. Treated earlier either as a form of dumetata (for instance Prout (1912−1916) or as a subspecies (Scoble 1999), but synonymized under taxon daubearia Boisduval, 1840 by Flamigni et al. (2016). See Variation. Gnophos dumetata vindobonica Vojnits, 1967: Annls hist.-nat. Mus. natn. Hung. 59: 379, figs 5, 6 (Austria: Vienna, Dornbach). Holotype ♂ (Hungarian Natural History Museum, Budapest, Hungary, examined externally). Valid at subspecific rank (Scoble 1999). See Variation. Odontognophos dumetata hibernica Forder, 1993: Entomologist’s Rec. J. Var. 105 (9–10): 202, figured (Ireland: County Clare). Holotype ♂ (private coll. P. Forder, West Yorkshire, England, examined externally). Valid at subspecific rank (Scoble 1999; Bond & O’Connor 2012). Unavailable names (infrasubspecific): costata: Leraut (2009) (f.).

External characters and abdomen: Wingspan 32−40 mm. Wings brown, grey-brown to whitish grey, markings black. Antemedial and medial lines weakly developed, postmedial line strongest, often discontinuous, its outer margin bordered with narrow shadow. Medial lines widest on forewing costa. Hindwing base irrorated or weakly striated with blackish. Terminal area darker (brown). Terminal line blackish. Wing termen distinctly undulating. Discal spots weak or absent. Underside of wings almost without markings, except rather large discal spots and diffuse postmedial line. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Male and female antennae filiform. Tympanal organs medium-sized. Hindlegs in both sexes with 2+2 spurs, male hindleg slightly swollen with narrow hair pencil. Abdominal sternites unmodified. Variation: Odontognophos dumetata is a widespread, rather variable species, with a strong tendency towards producing local phenotypic variations, apparently being the reason for numerous putative subspecies. These need to be evaluated critically, and investigations must include extensive genetic data. As such data is not available at the moment, we do not give descriptive accounts (apart from subsp. hibernica), and do not mark them on the distribution map. The variation can be summarized as follows: forewings chocolate brown with shadow on postmedial line (taxon dumetata); forewings sand-grey with olive hue and shadow on postmedial line absent (taxon vindobonica); forewings ash-grey, weakly tinged with brownish to greyish white (taxon daubearia); forewings ash-white (taxon scopulata); forewings blue-grey with rather distinct medial lines (taxon hibernica). Terminal area often darker (brown), but sometimes barely noticeable. Subsp. hibernica has blue-grey forewings with rather distinct medial lines. Male cornuti are smaller and less dentate than in nominate subspecies (Text-figs 1−6). Corpus bursae in the female genitalia resemble O. margaritata, but the membranous structures are difficult to compare. Known from western Ireland: County Clare only. The origin of the isolated Irish population is uncertain, but it has been speculated to be an interglacial relict from the late pleistocene. The County Clare locality hosts isolated populations of other species also, including species such as Calamia tridens (Hufnagel, 1766) (Noctuidae: Xyleninae) (Elliot & Skinner 1995) and Aspitates gilvaria (Denis & Schiffermüller, 1775) (Skou & Sihvonen 2015).

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Male genitalia: Uncus narrowing evenly, setose dorsally. Socii absent. Gnathos well-developed, medial element upturned, weakly granulate. Valva divided. Ventral arm of valva with round and spinose apex, dorsal arm of valva heavily setose. Ventral part of juxta simple, elongated, apex granulate and often rounded. Dorsal part of juxta with two shallow concavities. Saccus wide, weakly elongated. Aedeagus with two sclerotized, dentate ridges. Vesica base weakly expanded, covered with microcornuti. Vesica opens between 45−90 degree angle (variable). Female genitalia: Papillae anales elongated, setose. Lamella postvaginalis and antevaginalis fused, massive and strongly sclerotized. Posterior margin with narrow extension medially, anterior margin round. Corpus bursae very large, posterior part long, anterior part expanded sac. Signum elongated plate with horizontal ridge, located at anterior end. Distribution and abundance: South European-West Asiatic. In Europe from south-eastern European Russia through southern central Europe and parts of southern Europe to southern France and north-easternmost Spain and with a very isolated occurrence in the Burren, western Ireland (subsp. hibernica). – Somewhat local, usually found in limited numbers. – Outside Europe in Dagestan, Armenia and Turkmenistan.

Phenology: Univoltine. From early August to early October. In Hungary from late August to mid-September, with only few specimens before or after that, and with the earliest record 20th July and the latest 2nd October (Vojnits 1967). In Ireland from late July to mid-August. Larva from April or May to June or July. Hibernation as egg (M. Leipnitz pers. comm.). Adults active at night, males attracted to light, females more rarely. Biology: Larva possibly monophagous on Rhamnus. Larvae of the isolated Irish population found on Rhamnus cathartica (Skinner 2009; Woodrow et al. 2018). Recorded on Rhamnus alaternus and R. saxatilis (Flamigni et al. 2016). Erroneously recorded on Phillyrea latifolia 32

(Millère 1858–1864) (Flamigni et al. 2016). Reared on Rhamnus cathartica (M. Leipnitz pers. comm.). Pupation in a fine loose web on the ground among moss and plant litter (M. Leipnitz pers. comm.; rearing observation) down to 3–5 cm in the ground (Z. Las˘tu˚ vka pers. comm.).

Text-figs 1−6. Diagnostic characters (indicated) of Odontognophos dumetata subsp. ­dumetata (Treitschke, 1827) and subsp. hibernica Forder, 1993. The position of cornuti is not fully comparable in all photos, because some slides have been prepared by earlier authors without support for cover glass. The male genitalia capsules are similar in both taxa. Text-figs 1−3. O. ­dumetata subsp. dumetata. Text-fig. 1. Croatia: Dubrovnik-Neretna district, 215 m, 5.x.2015, slide Sihvonen 2302 (Skou). Text-fig. 2. France: Dept. Aude, 6 km N. of Maury, 400 m, 8.ix.2002, slide Sihvonen 2301 (coll. Skou). Text-fig. 3. Spain: Prov. Barcelona, 2 km NW of Gurb, 600 m, 11.ix.2002, slide Sihvonen 2717 (coll. Skou). Text-figs 4−6. O. dumetata subsp. hibernica. Text-fig. 4. Ireland: Co. Clare, Burren, without date, slide Goater 482 (coll. ZMUC). Text-fig. 5. Ireland: Co. Clare, Burren, 28.vii.1993, slide Goater 478 (coll. ZMUC). Text-fig. 6. Ireland: Co. Clare, Burren, without date, slide Goater 483 (coll. ZMUC).

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Habitat: Xerothermophilous. In open, warm scrubland. In Hungary in the clearings of karst woods and scrubland (in general, of dry oakwood), on slopes of dolomite rubble, sparsely grassy rocky slopes and on meadows on loose sand substrate (Vojnits 1967). In Bulgaria associated with open limestone rocky areas with petrophilous vegetation (Beshkov & Zlatov 2009). In Ireland on sparsely vegetated limestone pavement that may be subject to seasonal flooding (Woodrow et al. 2018). From sea-level up to 1500 m, sometimes up to 1850 m (Flamigni et al. 2016: Aosta Valley, Italy). Similar species: Odontognophos margaritata is very similar, and so far reliable external dia­ gnostic characters have not been found. Aedeagus apically bearing one sclerotized row in O. margaritata (two sclerotized rows in O. dumetata), see Text-figs 1−6, and valva apex is usually elongated in O. margaritata (usually round in O. dumetata). Shape of valva apex is somewhat variable, but the number of aedeagus sclerotizations has so far proven diagnostic. Female genitalia considerably smaller, margin of lamella antevaginalis irregular, sclerotization of lamella postvaginalis wide and anterior end of corpus bursae weakly expanded in O. margaritata (genitalia large, margin of lamella antevaginalis round, sclerotization of lamella postvaginalis narrow and anterior end of corpus bursae expanded in O. dumetata). O. zacharia also similar, but with dark blotches on postmedial line, uncus wide, aedeagus apically with three sclerotizations, and sclerotization of lamella postvaginalis curved.

Text-figs 7−9. Diagnostic male genitalia characters (indicated) of Odontognophos Wehrli, 1951 species. Text-fig. 7. O. dumetata (Treitschke, 1827). Croatia: Dubrovnik-Neretna district, Oslje, 215 m, 5.x.2015, slide Sihvonen 2302 (coll. Skou). Text-fig. 8. O. margaritata (Zerny, 1927). Spain: Prov. Teruel, Albarracin, 1200 m, 18.ix.2001, slide Sihvonen 2299 (coll. Skou). Text-fig. 9. O. zacharia (Staudinger, 1879). Greece: Crete, Nom. Iraklion, 1450 m, 29.ix.2016, slide Sihvonen 2306 (coll. Skou).

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Genetic data: BIN: BOLD:AAU3550 (n=8 from Ireland, Italy, north-easternmost Spain, eastern Spain and Croatia). The barcode from Ireland (subsp. hibernica) only diverging by 0.15% (1bp) from those of Italian and Catalonian populations. Nearest species: O. margaritata 0.9% from a Croatian specimen, and 1.6% from Italian and Catalonian populations. COI data not suggesting any relationship with other Gnophos species, but a potential position between genera Asovia and Erannis, while multi-gene analysis clearly shows a position in the Abraxini + Cassymini + Eutoeini group, tentatively in Abraxini (Murillo-Ramos et al. 2019).

2. Odontognophos margaritata (Zerny, 1927) Gnophos dumetata margaritata Zerny, 1927: Eos. Madr. 3 (4): 425 (Spain: Aragon, Albarracin). Syntypes 3♂1♀ (Naturhistorisches Museum, Vienna, Austria, examined externally).

External characters and abdomen: Generally as in O. dumetata, see description above, but with following quantitative differences: wingspan on average 35.6 mm (n=10), wings greybrown, postmedial line distinct on upperside, weakly visible on underside in O. margaritata (wingspan on average 39.5 mm (n=30), and wing colour variable, but in closest populations in north-eastern Spain greyish white with brown terminal area, postmedial line broken into spots on upperside, and more visible on underside in O. dumetata). Variation: Rather little. Wing colour varies from grey-brown to light grey, brown irrorations can be weak or intense. Specimens taken from the type locality (Spain: Albarracin) are grey-brown, finely irrorated with brown or blackish brown. Zerny (1927a) described them as “pearl grey” with brownish irroration. Postmedial line often complete, distinct. Male genitalia: Generally as in O. dumetata, see description above, and in O. zacharia, but with following differences (see Text-figs 7−9): aedeagus apically ornamented with one row of sclerotized teeth and valva apex pointed in O. margaritata (two sclerotized rows and ­valva apex round in O. dumetata; three sclerotized rows and valva apex pointed and curved ventrally in O. zacharia). Female genitalia: Generally as in O. dumetata, see description above, and in O. zacharia, but with following differences: Corpus bursae small, posterior margin of lamella postvaginalis with wide extension, ductus bursae funnel-shaped, margin of lamella antevaginalis irregular and anterior end of corpus bursae weakly expanded in O. margaritata (corpus bursae large, posterior margin of lamella postvaginalis with narrow extension, ductus bursae funnel-shaped, margin of lamella antevaginalis round and anterior end of corpus bursae expanded in O. dumetata; posterior margin of lamella postvaginalis with curved extension, ductus bursae wide in O. zacharia). The mentioned differences are tentative, due to small number of specimens available for study (O. dumetata 4 specimens, O. margaritata 1 specimen, O. zacharia 1 specimen). Distribution and abundance: An Iberian endemic. In northern central Spain. – Somewhat local, usually found in limited numbers. Phenology: Univoltine. So far recorded from late August to mid-October. Adults active at night, males attracted to light, rarely also the females. Biology: Unknown, but probably much like that of O. dumetata. Habitat: Xerothermophilous. In open, warm scrubland. From 200 up to 1250 m. Similar species: O. margaritata and O. zacharia, see text under O. dumetata. 35

Genetic data: BIN: BOLD:AAU3550 (n=2 from eastern Spain). Genetically homogeneous. Nearest species: O. dumetata (0.9%). Remarks: Taxon margaritata was raised to species rank by Leraut (2009), based on the male genitalia characters. The mentioned aedeagus and valva characters appear diagnostic, in addition to the female characters diagnosed in the current volume, and we consider O. margaritata valid at species level, as did Vives Moreno (2014) also. So far it has been possible to classify all examined specimens in the Iberian peninsula either as O. dumetata or O. margaritata, but we highlight that more material needs to be examined in the Pyrenees and north-eastern Spain to exclude the possibility of clinal variation. Interestingly, these taxa have not yet been found to occur sympatrically. The sclerotized rows of teeth at the aedeagus apex (two in O. dumetata and one in O. margaritata) have so far been diagnostic, whereas the shape of valva apex (round in O. dumetata, elongated in O. margaritata) is more variable. Gnophos faroulti Rothschild, 1914 (Novit. zool. 21: 351, Algeria: Guelt-Es-Stel, syntypes 7♂2♀ in NHMUK, examined, including male genitalia) has recently been classified as a subspecies of Odontognophos dumetata (Treitschke, 1827) (Scoble 1999). This non-European taxon is structurally similar to O. margaritata and we consider it valid at species-level as ­Odontognophos faroulti (Rothschild, 1914) stat. rev., based on diagnostic male genitalia structures (Text-fig. 10, diagnostic structures of European species are illustrated in Text-figs 7–9). Male genitalia capsule of O. faroulti is as in O. margaritata, but aedeagus apex is curved distinctly, cornuti teeth are distinct and base of cornutus is round in O. faroulti (apex weakly curved or straight, cornutus teeth smaller and base of cornutus not round in O. ­margaritata). O. faroulti has wings weakly irrorated with blackish, almost without markings except interrupted postmedial line. Wingspan (n=13) is similar to that of O. margaritata. NHMUK has two syntype females of O. faroulti only, and both of those have been dissected by an anonymous person, but the slides are missing. No other females were available to us, therefore 36

examination of female structures and DNA barcodes are needed to understand whether this North-African allopatric taxon is better classified as separate species (as here) or as a subspecies of O. margaritata (from the nomenclature point of view O. faroulti is the senior name). More research is needed.

Text-fig. 10. Diagnostic male characters (indicated) of non-European Odontognophos ­faroulti (Rothschild, 1914) stat. rev. Aedeagus apex holds diagnostic features in this species-group (compare against other text-figures), so upper right hand corner shows the structures enlarged. Aedeagus shown non-everted and everted. Female genitalia were not available for study, see text for details. Wingspan. 31 mm. Algeria: Guelt-es-Stel, 7.x.1912, slide Sihvonen 2777, syntype (coll. NHMUK).

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3. Odontognophos zacharia (Staudinger, 1879) Gnophos zacharia Staudinger, 1879: Horae Soc. ent. ross. 14: 454 (Turkey: Amasia). Holotype ♀. Odontognophos zacharia dagestanensis Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 573, pl. 47: h (Russia: Dagestan). Syntypes (Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia). Validated at subspecific rank (Scoble 1999), awaiting confirmation.

External characters and abdomen: Wingspan 31−37 mm. Wings grey to pale beige-grey, weak markings and irrorations brown to blackish. Forewing postmedial line with two diffuse, dark grey blotches. Antemedial and postmedial lines weak, postmedial line strongest yet diffuse and broken into spots. All postmedial lines expanded and distinct on forewing costa. Subterminal line blackish grey, most distinct near forewing costa. Terminal area darker. Terminal line weak, blackish. Fringes weakly chequered. Underside of wingsalmost without markings, subterminal line occasionally present as diffuse dark fascia. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Male and female antennae filiform. Tympanal organs medium-sized. Hindlegs in both sexes with 2+2 spurs, male hindleg weakly swollen with narrow hair pencil. Abdominal sternites unmodified. Variation: Rather little. Wing colour and size varies geographically. Specimens are grey and small in Greece: Crete and Turkey: Antalya (wingspan 31−33 mm). Specimens are grey and large in mainland Greece (wingspan 36−37 mm), and those from eastern Turkey are brown and large (wingspan 35−36 mm). Subsp. dagestanensis is large (wingspan up to 37 mm), dark grey-brown with distinct, blackish medial lines, and hindwings equally dark (Wehrli 1953). Subspecific rank (Scoble 1999) awaiting confirmation. We have examined two males from Turkey: Erzurum (coll. Stadie), and those are brown, approaching in appearance subsp. dagestanensis. Male genitalia: Generally as in O. dumetata, see description above, and in O. margaritata, but with following differences: uncus wide, aedeagus apically with three sclerotized rows (Text-figs 11−13) and valva apex pointed and curved ventrally in O. zacharia (uncus narrow, aedeagus apically with two sclerotized rows and valva apex round in O. dumetata; uncus narrow, aedeagus apically with one sclerotized row and valva apex elongated but not curved ventrally in O. margaritata). The number of teeth on sclerotizations on aedeagus is variable, see Text-figs 11−13. Female genitalia: Generally as in O. dumetata, see description above, and in O. margaritata, but with following differences: corpus bursae small, posterior margin of lamella postvaginalis with curved extension and ductus bursae wide in O. zacharia (corpus bursae large, posterior margin of lamella postvaginalis with narrow extension, ductus bursae funnel-shaped, margin of lamella antevaginalis is round and anterior end of corpus bursae is expanded in O. dumetata; posterior margin of lamella postvaginalis with wide extension, ductus bursae funnel-shaped in O. margaritata). The mentioned differences are tentative, due to small number of specimens available for study (O. dumetata 4 exx., O. margaritata 1 ex., O. zacharia 1 ex.). Distribution and abundance: East Mediterranean-Caucasian. In Europe only known from Crete and Mount Parnassos in mainland Greece. – A poorly known and rare species with only a few specimens known from Europe. In Crete only in few places and very rare (Ruckdeschel 2007). – Outside Europe in Turkey, Georgia, Armenia, Dagestan (tentative subspecies dagestanensis), Azerbaijan and Turkmenistan. 38

Text-figs 11−13. Number of teeth varies in aedeagus apex in Odontognophos zacharia (­Staudinger, 1879). Text-fig. 11. Greece: Crete, Nom. Iraklion, Ida Mts., 1450 m, 29 ix.2016, slide Sihvonen 2306 (coll. Skou). Text-fig. 12. Turkey: Erzurum, near Ispir Westl. Ortsrand, 1100 m, 30.ix-8.x.2000, slide Sihvonen 2716 (coll. Stadie). Text-fig. 13. Greece: Fthiótida, Parnassos Ski Centre, 1750 m, 6.ix.2008, slide Sihvonen 2307 (coll. Skou).

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Phenology: Univoltine. Scarce data from early and late September, early and mid-October and early November. Natural larval time unknown. Hibernation as egg (M. Leipnitz pers. comm.; rearing observation). Adults active at night, attracted to light. Biology: Larva apparently monophagous on Rhamnus. Reared on Rhamnus cathartica and R. saxatilis (M. Leipnitz pers. comm.). Pupation in a fine loose web on the ground among moss and plant litter (M. Leipnitz pers. comm.; rearing observation). Habitat: Xerothermophilous. In mainland Greece in open mountain forest. In Crete in open land with low growing bushes. In Turkey recorded from sea-level up to 1000 m, in Crete from 1450 to 1600 m and in Greece at 1750 m, close to the treeline. Similar species: Odontognophos dumetata and O. margaritata, see text under O. dumetata above. European specimens of O. zacharia are diagnosable by grey colour and two diffuse, dark grey blotches in forewing postmedial line. These blotches are weak or absent and specimens are brown in eastern Turkey and in Caucasus. Male and female genitalia are diagnostic. Genetic data: BIN: BOLD:AAU3550 (n=2 from Turkey). Both sequences identical. BIN-­ sharing with O. dumetata but diverging by 1.4%. Remarks: Wehrli (1934: plate VI, fig. 44) illustrates an early photograph of the O. zacharia male genitalia from Turkey: Aksehir, matching well with the European and Turkish specimens we have studied.

Tribus Cassymini (part): Addition (other species treated in volume 5)

Dicrognophos Wehrli, 1951 (new tribus combination) Dicrognophos Wehrli, 1951, Lambillionea 51: 10. Type species: Gnophos orthogonia Wehrli, 1939. Originally proposed as a subgenus of Gnophos Treitschke, 1825, and was again so proposed by Wehrli, 1953 (in Seitz, Gross-Schmett. Erde 4 (Suppl.): 578).

Diversity and distribution: Dicrognophos contains about 15 species (Wehrli 1951), main area of diversity is in Middle East, particularly in Iran. External characters and abdomen: Rather large moths, wingspan 28–45 mm. Wing coloration in different shades of brown, yellowish brown or grey-brown, often heavily irrorated with dark brown, particularly in terminal area. Medial lines enlarged on forewing costa, appearing as two or three dots, medial line often strongest, not sharply defined. Terminal area with pale blotch halfway in each wing, additional blotch on fore – and hindwing apex but weaker. Underside of wings darker, particularly in terminal area, pale blotches more distinct, including those on wing apex. Male antennae ciliate-fasciculate, base swollen, cilia long or short. Female antennae filiform. Hindlegs in both sexes with 2+2 spurs, male hindleg slightly swollen with narrow hair pencil. Abdominal sternites unmodified. Male genitalia: Uncus narrow, setose. Socii absent. Gnathos weakly developed or absent. Tegumen elongated, triangular. Valva divided into sclerotized ventral sacculus and free, curved dorsal ampulla. Juxta simple, elongated, sclerotized. Saccus round. Aedeagus with one large cornutus. 40

Female genitalia (based on European species only): Papillae anales very large. Lamella postvaginalis membranous or weakly sclerotized. Lamella antevaginalis loosely rectangular in shape, forming deep cavity. Ductus bursae short, sclerotized. Corpus bursae elongated, posterior part sclerotized and grooved. Signum absent. Biology, habitat, phenology: Larva of D. sartata recorded on Rhamnus alaternus, Rosa, Prunus and Polygonum (see Flamigni et al. 2016 and references therein). Dicrognophos sartata occurring in the plains and hills, up to 800 meters in Italy (Flamigni et al. 2016), living in garrigue, scrub, oak (especially Quercus ilex) and pine forests and in cultivated areas (Flamigni et al. 2016). The Iranian species assumedly inhabit semi-arid or even arid areas. D. culminata (Brandt, 1938) has been found at 3750 meters in Iran. Both sexes are attracted to light. Immature stages: Larva of D. sartata pale brown with grey-blackish markings, and two small, horn-shaped dorsal projections on 11th segment (Lepiforum 2018). Pupal cremaster with four pairs of setae, D2 setae being the largest (Lepiforum 2018). Remarks: Association of Dicrognophos species with species in the Abraxini + Cassymini + Eutoeini group supported for instance by divided valva with dorsal arm narrow and curved, setose apically; male 8th sternite unmodified and chaetosemata present, bushshaped. Some structures uncharacteristic for that group, being potentially homoplasious: lack of fovea in forewing, well-developed and sclerotized juxta, gnathos weakly developed, transverse setal comb absent on male 3rd abdominal sternite, forewing radial veins developed normally (five veins present), pupal cremaster with four pairs of setae, stellate female signum absent. Recently Dicrognophos has been classified in the Gnophini (e.g. Viidalepp 1996; Hausmann et al. 2004; 2011a). Systematic position: Abraxini + Cassymini + Eutoeini group, tentatively Cassymini.

4. Dicrognophos sartata (Treitschke, 1827) Gnophos sartata Treitschke, 1827: Schmett. Eur. 6 (1): 175 ([Croatia] Dalmatia). Syntypes, at least 20 specimens. Scotopterix immundata Warren, 1899: Novit. zool. 6: 347 (Lebanon: Beirut). Syntypes 2♂1♀ (NHMUK, examined externally).

External characters and abdomen: Male wingspan 28–43 mm (on average 33.3 mm), female 32–45 mm (on average 38.5 mm). Wings pale yellow, markings and extensive irrorations brown. Medial line not sharply defined, wide. Postmedial line as row of blackish brown dots. Subterminal line pale yellow, most distinct near forewing costa. Terminal area with pale blotch halfway on each wing. Discal spots large, faint. Terminal line fine. Fringes concolorous with wings. Hindwing termen undulating. Underside of wings darker, particularly on terminal area making terminal area blotches more distinct, additional blotches on fore – and hindwing apices also visible. Discal spots more distinct on underside. Frons and labial palpi dark brownish black. Collar, thorax and abdomen concolorous with wings. Male antennae ciliate-fasciculate, with very short cilia. Bases swollen and fused with adjoining segments. Female antennae filiform. Hindlegs in both sexes with 2+2 spurs, male hindleg slightly swollen, hair pencil present. Male tympanal organs large, touching each other. Female tympanal organs medium-sized. Abdominal sternites unmodified.

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Variation: Wingspan varies a lot (compare to D. siciliana), smallest males can be almost 2 cm smaller than largest females. Wingspan varies in males between 28–43 mm (spring generation) and 28–41 mm (autumn generation); in females between 32–45 mm (spring generation) and 33–42 mm (autumn generation). Wing patterns varying little but with variable degree of brown to grey-brown irrorations on wings. Terminal area on underside of wing almost blackish, in such case the characteristic pale blotches standing out very distinctly. Male genitalia: Uncus narrow, triangular, heavily setose. Tegumen elongated, triangular, with distinct mid-dorsal suture. Socii absent. Gnathos wide, but weakly sclerotized band. Valva divided. Ventral arm of valva (sacculus) sclerotized, apex free and setose. Dorsal arm of valva (ampulla) narrow, curved, apex heavily setose, base with sclerotized tooth. Centre of valva membranous. Juxta wide, concave, with two pairs of lateral extentions (Text-figs 14–15). Anterior arms of juxta absent. Saccus round. Caecum enlarged, apex dorsoventrally flattened, with distinct tooth. Vesica weakly expanded sac, base with single large cornutus, its apex elongated. With lateral row of small cornuti (Text-figs 18–23). Female genitalia: Papillae anales very large, setose. Apophyses posteriores slightly longer than apophyses anteriores. Lamella postvaginalis membranous, horizontally striated. Lamella antevaginalis membranous, loosely rectangular in shape, forming deep cavity. Posterior margin of ductus bursae sclerotized, half-ringed. Ductus bursae short, wide, partly sclerotized. Corpus bursae elongated, right margin (in ventral view) without angle, posterior part sclerotized and with vertical grooves, anterior part membranous and often slightly expanded. Signum absent. Accessory glands sometimes massive. Distribution and abundance: (South-east) European-Anatolian. In Europe only in southern Balkan peninsula including Crete, along the coast east of the Adriatic Sea and through the main part of the Italian peninsula. In ZFMK one specimen from south-eastern France in 1967 (C. Tautel pers. comm.). Previously also recorded in Sicily (Flamigni et al. 2016),

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Text-figs 14–15. Juxta of Dicrognophos sartata (Treitschke, 1827) in dorso-lateral view, removed from genitalia capsule. Juxta concave, with two pairs of lateral extentions. Text-fig. 14. Italy: Puglia, 7 km N Altamura, 475 m, 29.v.2016, slide Sihvonen 2202 (coll. Skou). Text-fig. 15. Greece: Epiros, Parga, 17–31.v.1999, slide Sihvonen 2200 (coll. Sihvonen).

requiring examination and probably referring to D. siciliana. A very old and doubtful record from Ukraine (Belke 1859; cf. Kostjuk 2004), mentioned from Crimea by Viidalepp (1996), but no specimens known from there (Savchuk pers. comm.). – Common, often occurring abundantly. – Outside Europe known from Turkey, Cyprus, Lebanon, Israel, Jordan, Caucasus and Iran. Phenology: Bivoltine. Generally from late April to late June and from late August to mid-­ October. In Greece recorded in mid-February, in Samos in early April, in central Turkey in mid-July, in Cyprus in early November and in Israel both in mid-April and throughout November. A last instar larva found in late April (D. Fritsch pers. comm.). Hibernation as a small or medium-sized larva (M. Leipnitz pers. comm.). Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Calicotome (M. Leipnitz pers. comm.) and Pyrus spinosa (D.  Fritsch pers. comm.). Recorded on Rhamnus and low plants (Fajčik 2003). Reared on Lonicera tatarica (M. Leipnitz pers. comm.) and Genista (D. Fritsch pers. comm.). Pupation in a light silken web among mosses, plant litter and soil particles on the ground (M. Leipnitz pers. comm.; rearing observation).

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Habitat: Xerothermophilous. In a large variety of habitat types, both open and places with scattered trees and bushes including gardens. From sea-level up to at least 1200 m. In ­Cyprus up to 1950 m. Similar species: Dicrognophos siciliana, see that species for diagnostic characters. Dicro­ gnophos sartata and D. siciliana clearly differing from other European species by their pale blotches in terminal area, one on each wing. These blotches more distinct on underside of wings. Genetic data: BIN: BOLD:AAB3861. Genetically rather homogeneous, but in Europe (n=32), splitting into two major lineages at a genetic distance of 1.55% (lineage 1: central and southern Italy: Puglia, Bulgaria, Greece including Crete, Turkey, Lebanon and lineage 2: southern Italy: Basilicata, Croatia, Cyprus). Nearest species: D. siciliana Skou & Sihvonen, sp. n. (5.26% from lineage 1 and 6.27% from lineage 2). Remarks: Leraut (2009) combined D. sartata with genus Odontognophos Wehrli, 1951 (see text under Odontognophos), and László et al. (2018) with genus Gnophos Treitschke, 1825. One of the examined specimens, a female from Bulgaria (Rhodopi Mts, Assenovgrad, 14.ix.1985, genitalia dissected, slide Sihvonen 2073, in coll. Skou) has small mites on underside of wings. These might be Cheletomorpha lepidopterorum (Shaw, 1794) (long-legged cannibal mite), or something close to it (see Enghoff 1988).

5. Dicrognophos siciliana Skou & Sihvonen sp. n. Holotype: ♂, ‘Italy, Sicily, 5,8 km ESE Santo Stefano Quisquina, 1000 m, 2.vi.2016, Peder Skou leg.’; Dicrognophos siciliana, Skou & Sihvonen, holotype [red rectangle label]; Pasi Sihvonen, prep. number 2208. [blue rectangle label]’, in coll. Skou (Denmark), to be deposited at the Zoological Museum, University of Copenhagen (Denmark). Paratypes (all with red rectangle label: Dicrognophos siciliana, Skou & Sihvonen, paratype): total 31♂12♀. – In coll. Skou, Denmark: 2♂: Italy, Sicily, Piano Zucchi, Rifugio Orestano, 1065 m, 5.vi.2016, Peder Skou leg. 2♂: Sicily, 5 km SE of Castelluzzo, 300 m, 12.v.2003, Morten Molgaard leg (1♂ dissected, slide Sihvonen 2092). 3♂2♀: Italy, Sicily, 5.8 km ESE Santo Stefano Quisquina, 1000 m, 2.vi.2016, Peder Skou leg. 2♂1♀: Italy, Sicily, 5.5 km ESE Collesano, ­Rifugio Orestano, 1100 m, 17.ix.2014, Peder Skou leg (1♂ dissected, slide Sihvonen 2210; 1♀ dissected, slide Sihvonen 2068). 4♂4♀: Italy, Sicily, 5.8 km E Santo Stefano Quisquina, near Pizzo della Rondine, 1000 m, 23.–24.ix.2014, Peder Skou leg. (1♂ dissected, slide Sihvonen 2069; 1♀ dissected, slide Sihvonen 2211; 1♂1♀ DNA barcode BMB Lep 00157, 00158). 1♂1♀, Italy, Sicily S., Maginelli river, 15 km N Noto, 400 m, 13.ix.2002, leg. M. ­Fibiger & G. Jeppesen (♀ DNA barcode BMB Lep 00156). – In coll. Nupponen, Finland: 10♂: Italy, Sicily, Ragusa 12 km NE, 470 m a.s.l., 9.ix.2015, T. Nupponen leg. 4♂1♀: Italy, Sicily, Mistretta 3 km S, 1060–1100 m a.s.l., 10.ix.2015, T. Nupponen leg. 1♀: Italy, Sicily, Ficuzza 7 km SE, Alpe Cucco, 1150 m a.s.l., 11.ix.2015, T. Nupponen leg. – In coll. CREASAM: 2♂, Italy, Sicily, Campo Italia, 476 m, 30.v.2010, leg. M. Infusino (♀ DNA barcode BC MI 0139, 145). – In coll. ZSM: 1♀, Sicilia, Madonie, 1200 m, 5.viii.[no year], leg. Dannehl. 1♂1♀, Sicilia, Palermo, S. Martino d. Scale, 20.–31.v.1954, leg. J. Klimesch. Etymology: The name of the species refers to the type locality.

Diagnosis: Dicrognophos siciliana is similar to D. sartata. Diagnostic characters are summarized in Table 1.

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Table 1. Diagnostic characters of D. sartata (Treitschke) and D. siciliana Skou & Sihvonen sp. n. See also the relevant text-figs. The minimum genetic distance (DNA barcode) supports diversification at the species-level. Character

D. sartata

D. siciliana

Wings Wings markings Wingspan Saccus Aedeagus apex Cornutus Caecum Corpus bursae margin

pale yellow brown 28–45 mm elongated with distinct spike base narrow and apex long enlarged without angle

grey-white dark grey to blackish 27–40 mm round with tiny spike base wide and apex short not enlarged with angle

Minimum genetic distance (DNA barcode) between D. sartata and D. siciliana: 5.26%

External characters and abdomen: Females on average slightly larger than males. Wingspan males 31–36 mm (spring generation) or 27–31 mm (autumn generation); wingspan females 39–40 mm (spring generation) or 29–33 mm (autumn generation). Generally similar to D. sartata, but with following differences: Wings grey-white and markings dark grey to blackish in D. siciliana (pale yellow and markings brown in D. sartata). Male has distinct, paired structures on the 8th segment laterally, located between sternites and tergites (Text-fig. 16). Such structures are present in D. sartata also, but they are very small or rudimentary. Variation: Very little. Amount of dark grey irrorations somewhat varying, in most specimens grey-white wing colour prevailing. Wingspan rather uniform (compare D. sartata), varying 5 mm in males (n=26), 4 mm in females (n=8).

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Text-figs 16–17. Structures on the 8th lateral segment of males in Dicrognophos siciliana Skou & Sihvonen sp. n. and Yezognophos serotinaria subsp. dognini (Thierry-Mieg, 1910). Function of these gland-like structures is unknown, but potentially they are part of the respiratory system because the structures on the 8th segment are located in a similar position to the openings on segments 2–7. The difference is, however, that the gland-like structures on the 8th segment are concealed within the abdomen and they do not have an opening like spiracles. The illustrated structures are absent or rudimentary in many Geometridae, but they are large for instance in genus Problepsis Lederer, 1853 (Sterrhinae: Scopulini). Text-fig. 16. Dicrognophos siciliana. ­Italy: Sicily, Santo Stefano Quisquina, 1000 m, 23–24.ix.2014, slide Sihvonen 2069 (coll. Skou). Textfig. 17. Yezognophos serotinaria subsp. dognini. Spain: Lerida, Guils, 1460 m, 16.vii.2003, slide Sihvonen 2187 (coll. Skou).

Male genitalia: Generally as D. sartata, but with following differences: Saccus round, aedeagus apically with tiny spike, cornutus base wide and apex short, caecum not enlarged in D. siciliana (saccus elongated, aedeagus apically with distinct spike, cornutus base narrow and apex long, caecum enlarged in D. sartata). See Text-figs 18–23. Female genitalia: Generally as D. sartata, but, with following difference: corpus bursae marginally with an angle in D. siciliana (corpus bursae margin without angle in D. sartata). Distribution and abundance: European, endemic to Italy: Sicily. Despite efforts, so far no specimens of D. siciliana have been found in mainland Italy, although special attention was given to specimens from southern Italy: Calabria (3 males and 1 female dissected), Basili­ cata (1 specimen barcoded) and Puglia (1 male and 1 female dissected). – Apparently rather common, found in numbers.

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Text-figs 18–23. Diagnostic characters (indicated) of Dicrognophos sartata (Treitschke) and D. ­siciliana Skou & Sihvonen sp. n. (figures are in scale). In addition to the indicated characters, the aedeagus is about 20% smaller in D. siciliana. Point of origin of ductus ejaculatorius is ­indicated by a circle. Text-fig. 18. Italy: Calabria, Monte Pollino, 800 m, 26.ix.2014, slide Sihvonen 2093 (coll. Skou). Text-fig. 19. Cyprus: Agia Napa, 50 m, 7.iv.1996., slide Sihvonen 2201 (coll. Skou). Text-fig. 20. Syria: Qual’aat Saam’ann, 535 m, 16.x.2000, slide Sihvonen 2206 (coll. Skou). Text-fig. 21. Italy: Sicily, 5.5 km ESE Collesano, 1100 m, 17.ix.2014, slide Sihvonen 2210 (coll. Skou). Text-fig. 22. Italy: Sicily, 5.8 km E Santo Stefano Quisquina, 1000 m, 23–24. ix.2014, slide Sihvonen 2069 (coll. Skou). Text-fig. 23. Italy: Sicily, 5 km SE of Castelluzzo, 300 m, 12.v.2003, slide Sihvonen 2092 (coll. Skou).

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Phenology: Bivoltine. Recorded from mid-May to mid-June and again in mid-September in Sicily, but probably with rather similar flight period to that of D. sartata. Adults nocturnal, readily attracted to light. Biology: Unknown. Habitat: Xerothermophilous (Text-fig. 24). Found in open forests with scattered trees, from 300 m up to 1150 m.

Text-fig. 24. Habitat of Dicrognophos siciliana Skou & Sihvonen sp. n. in Italy: Sicily, 5.8 km east of Santo Stefano Quisquina, 1000 m, 23.ix.2014. Photo: P. Skou.

Similar species: Dicrognophos sartata, see above for diagnostic characters. D. sartata and D. siciliana clearly differ from other European species by their pale blotches in terminal area, one on each wing, being more distinct on wing underside. Genetic data: Two BINs: BOLD:AAT9476 and BOLD:ACE9155 (n=5 from Sicily). Both BINs diverging by 1.4%. Nearest species in Europe: Dicrognophos sartata (Treitschke, 1827) (5.26% from D. sartata lineage 1 and 6.27% from D. sartata lineage 2). Remarks: The diagnostic corpus bursae margin (without an angle in D. sartata and with angle in D. siciliana) corresponds with orientation of the male cornuti (smaller angle in D. sartata and larger angle in D. siciliana), indicative of lock-and-key morphology.

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Tribus Macariini (part): Addition (other species treated in volume 5)

6. Ortaliella perspersata (Treitschke, 1827) (new tribus combination) Gnophos perspersata Treitschke, 1827: Schmett. Eur. 6 (1): 166. Replacement name for Geo­ metra respersaria Hübner, 1813. Geometra respersaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 79, fig. 406 (Europe). Syntypes. Junior primary homonym of Geometra respersaria Hübner, 1799, which is a junior synonym of Perconia strigillaria (Hübner, 1787), see Fletcher (1979) for further information. Gnophos perspersata chalcea Oberthür, 1913: Études Lépid. comp. 7: 296, pl. 177, fig. 1727 (Spain). Syntype(s). Treated as a form (Vives Moreno 2014; Wehrli 1939−1954), or as a subspecies (Scoble 1999), or not mentioned at all in several recent Spanish publications (for instance Vives Moreno 1994; Redondo & Gastón 1999; Redondo et al. 2009). We agree with Vives Moreno (2014), and consider it to fall within intraspecific variation. Gnophos perspersata ochrea Oberthür, 1913: Études Lépid. comp. 7: 296, pl. 177, fig. 1728 (Spain: Chiclana). Syntype(s). Treated as a form (Vives Moreno 2014; Wehrli 1939−1954), or as a subspecies (Scoble 1999), or not mentioned at all in several recent Spanish publications (for instance Vives Moreno 1994; Redondo & Gastón 1999; Redondo et al. 2009). We agree with Vives Moreno (2014), and consider it to fall within intraspecific variation. Potentially unavailable names (infrasubspecific): infuscatus: Schwingenschuss (1927) (ab.).

External characters and abdomen: Wingspan 30−39 mm. Wings brown to grey-brown, irror­ ated or striated with blackish brown and marked with blackish brown, weakly undulating medial lines. Forewing postmedial line angled inwards before costa. Terminal line narrow, continuous. Fringes concolorous with wings. Forewing discal spot small, sharp, hindwing spot often absent. Hindwing termen undulating. Underside of wings weakly marked, with small discal spots and evenly curved postmedial line, bordered with pale fascia. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Male and female antennae filiform. Tympanal organs medium-sized. Hindlegs in both sexes with 2+2 spurs, male hindleg slightly swollen with narrow hair pencil. Male sternites 1 and 2 with small but free lateral processes, male 8th sternite weakly concave, other abdominal sternites in both sexes without modifications. Variation: Little. Wings vary from grey-brown to sand-brown, and density of brown irroration or striation varies. All medial lines normally visible, but these can be weak, sometimes ante- and postmedial lines are almost absent. Male genitalia: Uncus round, densely setose, with few stout setae dorsally. Socii absent. Gnathos arms sclerotized, fused ventrally, forming granulate round plate. Juxta large, simple. Valva divided; dorsal arm of valva long; costa sclerotized and setose, its ventral margin terminated subapically. Ventral arm of valva short, sparsely setose, apex weakly elongated, medial part with small sclerotized flap. Transtilla wide, weakly dentate. Saccus wide, round. Aedeagus weakly curved, caecum long, apex spatulate Vesica opening at 90 degree angle. Basal part expanded, with one large cornutus, apex narrowing tube. Female genitalia: Papillae anales elongated, striated, setose. Apophyses anteriores, apophyses posteriores and 9th segment extremely long. Lamella postvaginalis large plate, posterior 49

margin concave. Lamella antevaginalis fused with ductus bursae. Ductus bursae short, wide, most sclerotized laterally. Posterior part of corpus bursae angled, weakly sclerotized and granulate, followed by long membranous section. Anterior part of corpus bursae expanded, ovoid, membranous, only granulate near signum (Text-fig. 25). Signum large, stellate. Distribution and abundance: West Mediterranean. In Europe only in the Iberian peninsula where the species is widely distributed. – Common, quite often found in numbers. – Outside Europe only two records in northern Morocco.

Phenology: Univoltine. Recorded from mid-April to late July. According to Redondo et al. (2009) from May to August. Continued moth trapping over a number of years near Este­ pona in southern Spain shows clearly, that the species has only one generation from first half of May to late June or the first days of July (P. Hale pers. comm.). The apparent longer flight period for the whole of the Iberian peninsula caused by latitude, altitude or c­ limatic reasons. Earliest records from Almeria province, the latest from mountains in northern Spain. Larva found in March and April (Gómez de Aizpúrua 2002), hibernation stage unclear. Adults active at night, males readily attracted to light, females less. Biology: Larva polyphagous. Found on Launaea arborescens (D.  Fritsch pers. comm.). Reared on Rhamnus alaternus (D. Stadie pers. comm.), and recorded on Rhamnus, Genista, Spartium (Redondo et al. 2009) and on Rhamnus lycioides (Gómez de Aizpúrua 2002), the latter illustrating photographs of caterpillar and pupa. Habitat: Xerothermophilous. In a wide range of habitat types such as dry, open scrubland or Mediterranean macchia. From sea-level up to 1600 m, according to Redondo et al. (2009) up to 2000 m in Sierra Nevada in southern Spain.

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Text-fig. 25. Signum and surface of corpus bursae of Ortaliella perspersata (Treitschke, 1827). The granulate surface of corpus bursae covers only areas adjacent to the signum, otherwise the surface is rather smooth. Similar granulate structure is found in congeneric O. gruneraria (Staudinger, 1862) also, see Skou & Sihvonen (2015).

Similar species: No similar species in Europe, but lack of distinct external features may lead to confusion, for instance with Gnophos furvata (Denis & Schiffermüller, 1775) and several species in Charissa Curtis, 1826. Small, distinct and solid discal spots and three medial lines of O. perspersata are characteristic. Other Ortaliella Hausmann, 1993 and closely related non-European Stueningia Hausmann, 1993 species are similar, but diagnosable by external and genitalia characters (Hausmann 1993b; Skou & Sihvonen 2015). Genetic data: BIN: BOLD:ABA9748 (n=3 from northern and eastern Spain, partly provided by P.  Huemer pers. comm.). Genetically homogeneous. Nearest species: Ortaliella palaestinensis (4.6%). Remarks: Systematic position of Gnophos perspersata Treitschke, 1827 has varied, demonstrating the difficulties that authors have faced while trying to classify it. Combinations include at least Gnophos (for instance Redondo & Gaston 1999; Corley 2015), Odonto­ gnophos (Redondo et al. 2009; Vives Moreno 2014), Odontognophos subg. Dicrognophos (Wehrli 1939−1954 who discusses its structures and affinities with different genera; Gómez de Aizpúrua 2002; Vives Moreno 1994) and Gnopharmia (Leraut 2009). Gnopharmia combination as suggested by Leraut (2009) is not supported by morphology (see illustrations of 51

structures in Rajaei 2010; Rajaei et al. 2012). External and structural features support a close relationship with the Macariini genera Ortaliella Hausmann, 1993 and Stueningia Hausmann, 1993 (see Hausmann 1993b; Skou & Sihvonen 2015 for descriptions and illustrations), accordingly we transfer Gnophos perspersata from Gnophini to Macariini. The proposed systematic position is supported among others by the setose uncus with several stout setae dorsally, gnathos structure, divided valva with a flap on medial part in the male genitalia; extremely long apophyses and 9th abdominal segment, stellate signum, similar corpus bursae shape and granulate surface on corpus bursae near signum in the female genitalia. Absence of fovea on forewing base in O. perspersata is an atypical feature for Macariini. Ortaliella perspersata specimens have been submitted to multi-gene phylogenetic analysis, in order to get further evidence on its systematic position.

Tribus Prosopolophini Warren, 1894 (part): Addition (other species treated in volume 5) Prosopolophini Warren, 1894 (Prosopolophinae). Type species: Geometra jourdanaria Serres, 1826, currently combined with genus Compsoptera Blanchard, 1845. Ligiini Guenée, 1858 (Ligidae), emended. Senior synonym, but based on a junior homonym of a genus group name outside Lepidoptera (Crustacea). Invalid according to Code art. 39 (ICZN 1999). Colotoini Wehrli, 1940 (Colotoinae). Type species: Phalaena pennaria Linnaeus, 1761, currently combined with genus Colotois Hübner, 1823. Subordinated under Prosopolophini in Beljaev (2016), here tentatively confirmed. Wilemanini Wehrli, 1941 (Wilemaninae). Type species: Gonodontis nitobei Nitobe, 1907, currently combined with genus Wilemania Prout, 1928. Subordinated under Prosopolophini in Beljaev (2016), here confirmed. Sclerotized structures of abdomen of Wilemania nitobei (­Nitobe, 1907) are illustrated in Nupponen & Sihvonen (2013). Male genitalia of W. nitobei are structurally similar to Apochima and Colotois (P. Sihvonen, pers. observation). Compsopterini Herbulot, 1963 (Compsopterini). Type species: Geometra jourdanaria Serres, 1826, currently combined with genus Compsoptera Blanchard, 1845. Subordidated under Prosopolophini in Beljaev (2016), here confirmed. ‡ Apochimini Viidalepp, 1989 (see Beljaev 2008 for details). ‡ Zamacrini Viidalepp, 1989 (see Beljaev 2008 for details).

Diversity and distribution: Potentially restricted to the Palaearctic region, currently with seven genera and about 15 species. Non-European Wilemania Prout, 1938 is known from Japan, China and Far-East Asia. External characters: Male antennae bipectinate, often with very long pectinations, female antennae filiform. Posterior margin of frons with lateral setae (see Skou & Sihvonen 2015, text-figs 8–9, all included species not examined). Collar with dorsally raised setae, giving mohican-like appearance in Compsoptera (see Skou & Sihvonen 2015, text-fig. 145). Wing shape and wing pattern very diverse. Resting position of Apochima is unusual: forewings rolled and often upright, resembling non-European Coenina Walker, 1860 (Ennominae). Male genitalia: Uncus often bilobed, asymmetrical. Valva wide, or partially divided. Male 8th sternite unmodified. Tergites in W. nitobei, D. furcicornaria and C. fiduciaria with sclerotized spines (Nupponen & Sihvonen 2013). Female genitalia: Papillae anales wide, stout. Ductus bursae often sclerotized, short, wide. Corpus bursae membranous. Signum small, weakly stellate or absent. 52

Biology: Polyphagy common, Colotois pennaria feeding on deciduous trees and shrubs (Ebert 2003), Dasycorsa modesta on Spartium junceum and Calicotome villosa, Compso­ ptera opacaria on Cytisus, Chondrosoma fiduciaria on Seseli tortuosum (Y. Budashkin & V. Savchuk pers. comm.) and Apochima flabellaria on Calendula arvensis, Anthemis (D. Fritsch pers. comm.). Flight period of many species in late autumn or even in winter. Immature stages: Eggs deposited in groups in Compsoptera jourdanaria, Chondrosoma fiduciaria and Apochima flabellaria. Larva of Compsoptera jourdanaria greyish brown, with two dark brown, undulating lines dorsally (Gómez de Aizpúrua 2002), that of Colotois pennaria is twig-mimic, smooth with two small, conical-shaped dorsal structures on A11 (Porter 2010) and that of Apochima flabellaria is covered with stout spikes. Pupa cremaster of Colotois pennaria is with two pairs of curved setae, apical pair wider (Patočka & Turčani 2005). Remarks: Pachycnemiini Kirby, 1903 has been considered a junior synonym of Prosopolophini (Forum Herbulot 2019 and references therein), but this is not supported by morphology (Skou & Sihvonen 2015) and not by molecular evidence (Sihvonen et al. 2011; Murillo-­Ramos et al. 2019). In multi-gene analysis, Prosopolophini (Compsoptera not analyzed) is closely related to Alsophila, and the entire complex is sister to Campaeini (Murillo-Ramos et al. 2019).

Apochima Agassiz, 1847 Apochima Agassiz, 1847, Nomencl. zool. (Index univl.): 29. Type species: Amphidasis flabellaria Heeger, 1838. [Replacement name]. Genus synonyms after Scoble (1999). ‡ Apocheima Herrich-Schäffer, 1838, in Panzer, Faunae Insect. Germ. initia (Dtl. Insect.) (Heft 163): wrapper (legend for fig. 11). Type species: Amphidasis flabellaria Heeger, 1838. [Junior homonym of Apocheima Hübner, [1825] 1816 (Geometridae, Ennominae)]. Zamacra Meyrick, 1892, Trans ent. Soc. Lond. 1892: 101 (key), 121. Type species: Amphidasis flabellaria Heeger, 1838. ‡ Acanthocampa Dyar, 1905, Proc. U. S. natn. Mus. 28: 951. Type species: Acanthocampa excavata Dyar, 1905. [Junior homonym of Acanthocampa Packard, 1902 (Lepidoptera, Satur­ niidae)]. Bellachima Kemal et al., 2018, Type spesies: Zamacra diaphanaria Püngeler, 1904. Since the molecular analysis shows several flaws (just based on the mtDNA COI gene for genus diagnosis; just based on bootstrap support and not considering the genetic distances from A. flabellaria (8.5%) and A. (d.) rjabovi; no comparison with genetic distances from other closely allied genera like Chondrosoma etc.), the validity of the genus is not accepted here.

Diversity and distribution: Apochima contains five species in the Palaearctic region. External characters and abdomen: Large species. Wingspan 24–43 mm. Wings narrow, reddish brown or greyish. Forewing with straight costa. Veins R1 and R2 fused, anastomosing with Sc. Apex of hindwings pointed. Abdomen, thorax, tegula appearing furry. Fovea absent from male forewings. Labial palpi bushy scaled, proboscis vestigial. Frons roughscaled. Male antennae bipectinate to tip with long branches, female antennae filiform. Male genitalia and female genitalia: Single species in Europe, see below, Remarks: Apochima is listed under Prosopolophini in Beljaev (2016), who regards Colotoini as synonym with Prosopolophini. See the checklist for a potential relationship to genus Coenina. Relationship between Colotois, Apochima and Chondrosoma is supported by shape of the uncus and suggested by DNA barcode similarity. Apochima has not been included in a multi-gene analysis so far. 53

7. Apochima flabellaria (Heeger, 1838) Amphidasis flabellaria Heeger, 1838: Beiträge zur Schmetterlingskunde: 6, pl. 1, figs 6–11 (Italy: Palermo). Syntype(s). Zamacra flabellaria var. marmararia Buresch, 1915: Spis. bulg. akad. nauk: 12 (Nat. Sci. & Math.) (5): 49 (Turkey: Silivri on Sea of Marmara). Holotype ♂. Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features.

External characters and abdomen: Wingspan ♂ 34–38 mm, ♀ 40–43 mm. Wings narrow, apex of hindwings pointed. Wings speckled with dark scales. Ground colour brownish, in females sometimes more whitish grey. Postmedial line straight, on forewing bent close to costa, on hindwing diffuse. Medial line thick, diffuse. Antemedial line bent. Terminal area of forewings sometimes lighter, divided from the rest of the wing by the distinct and lightly shadowed subterminal line. Apex usually with white streak. Discal spots diffuse, often indistinct. Fringes chequered. Male antennae bipectinate to tip with long, dark grey branches, female antennae filiform. Frons, collar, tegula and thorax densely covered with hairy scales, grey to brown. Abdomen greyish brown, weakly hairy, segments dorsally with black dot. Hindtibia of both sexes with two spurs. Variation: Little. Ground colour darker or lighter, degree of speckling varies. Male genitalia: Uncus wide, bilobed. Gnathos wide, arms ventrally not fused. Valva stout, simple, wide at base, granulate, apex pointed with setae. Costal area membranous. Juxta a plate with two arms half of juxta length. Saccus long, anteriorly concave. Aedeagus claviform, with dilatation at the anterior end. Without cornutus. Female genitalia: Papillae anales large, oval, setose. Apophyses posteriores long, robust, length on average 1.55 mm. Apophyses anteriores shorter, about half length of apophyses posteriores. Both ending hook-shaped. Lamella antevaginalis alate. Ductus bursae long, straight, weakly sclerotized. Corpus bursae membranous, posteriorly weakly striated. Without signum. Distribution and abundance: Central Asian-Mediterranean. In Europe in Crimea, Bulgaria, Greece incl. Crete and in Italy incl. Sardinia and Sicily; also a few records in south-western Spain. – Rather common, now and then occurring in numbers. – Outside Europe from Turkey and Cyprus to the Levant (Syria, Israel, Jordan), Transcaucasus (Azerbaijan, Armenia) Iran and Turkmenistan. Also in North Africa: Morocco, Algeria, Tunisia and Libya. Phenology: Univoltine: Generally from late November to early April. In Cyprus from late November to early March. In Crimea from late January to early March (V. Savchuk pers. comm.). In Morocco from December to February (Rungs 1981). In Bulgaria from late November to late March with a single record from late October (S. Beshkov pers. comm.). In Turkey in early April in Taurus Mountains (T. Drechsel pers. comm.). In Samos larvae from April (Fritsch et al. 2014) to early June, in Sardinia and Crimea to late May (C. Siegel, V. Savchuk pers. comm.), in central Turkey also to early June (J. Gelbrecht pers. comm.). Adults active at night, males readily attracted to light, females less often.

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Biology: Larva polyphagous. Found on Calendula arvensis, Anthemis (D.  Fritsch pers, comm.) Tanacetum parthenium, Anthemis chia (Fritsch et al. 2014), Ferula communis (C. Siegel pers. comm.), Medicago sativa, Vicia sativa (F. Can pers. comm.), Pastinaca clausii and Tragopogon dubius (V. Savchuk pers. comm.). Recorded on Calendula officinalis, Senecio, Chrysanthemum, Galium, Foeniculum vulgare, (Flamigni et al. 2007). Reared on Trigonella phoenum-graecum, Sonchus tenerrimus (Flamigni et al. 2007) and Tanacetum parthenium (M. Leipnitz pers. comm.). Pupation in the ground (V. Savchuk, M. Leipnitz pers. comm.). In rearings, pupation in a very firm cocoon of moss and soil (M. Leipnitz pers. comm.). Habitat: Xerothermophilous. In a variety of open habitats, in dry meadows and places with scattered trees and bushes. Ruderal areas and roadsides (D. Fritsch pers. comm.). In Bulgaria in open meadows and pastures from sea-level up to 400 m (S. Beshkov pers. comm.). In Crimea in steppe localities with clay soil (V. Savchuk pers. comm.). From sea-level, in southern Appenines in Italy up to 1400 m (Flamigni et al. 2007). Similar species: No similar species in Europe, identification easy based on the wing shape. Genetic data: BIN: BOLD:AAF8334 (n=10 from Bulgaria, Turkey, Israel, Kyrgyzstan). Genetically heterogeneous, maximum variation 3.1%; diverging into three allopatric, BIN-sharing clusters, one from Europe, another from the Levant, the third from Turkey and Kyrgyzstan. Nearest species (in the Palaearctic): A. diaphanaria rjabovi (Wehrli, 1936) from Transcaucasus and Turkey (8.2%). Remarks: Resting position of adults unusual, with wings rolled and forewings set in upright position.

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Chondrosoma Anker, 1854 Chondrosoma Anker, 1854, Verh. zool.-bot. Ver. Wien 4 (Sber.): 111. Type species: C ­ hondrosoma fiduciaria Anker, 1854. There are no genus synonyms.

Diversity and distribution: Chondrosoma is a monotypic genus, the only species occurs in the West Palaearctic region (South-East Europe). External characters, abdomen and Genitalia: See species description below. Systematic position: Tentatively classified as member of Prosopolophini (new classification). Relationship between Colotois, Apochima and Chondrosoma supported by shape of the uncus and suggested by DNA barcode similarity. In Viidalepp (1996) placed in Bistonini between Apocheima and Agriopis. Chondrosoma has not been included in a multi-gene analysis so far.

8. Chondrosoma fiduciaria Anker, 1854 Chondrosoma fiduciaria Anker, 1854: Verh. zool.-bot. Ver. Wien 4 (Sber.): 112 (Hungary: by Budapest (‘Ofen’)). Syntype(s).

External characters and abdomen: Wingspan ♂ 20–25 mm, ♀ wingless. Male: Wings covered with scattered creamy scales. Ground colour brownish grey, indistinctly irrorated with whitish scales, veins dark grey. Postmedial line on forewing distinct, creamy, thicker. Other lines reduced to three creamy blotches in basal area of forewing and to a creamy streak on basal area of hindwing. Frons, collar, tegula and thorax densely covered with creamy hairy scales, abdomen short, blackish with creamy hairy scales. Antennae bipectinate to tip, branches long, black. Female: Frons creamy. Collar, tegula and thorax with mixture of creamy and dark brown hairy scales. Abdomen cylindric, annulated reddish brown and grey. Antennae ciliate-fasciculate. Male abdominal tergites covered with stout sclerotized spines and setae (cf. Nupponen & Sihvonen 2013). Variation: Very little. Male genitalia: Simple structure. Uncus with two rounded tips. Valva setose, wide at base, apex narrower, round. Costa weakly sclerotized, setose subapically. Ventral margin concave, followed by an angle, at base sclerotized. Saccus round. Juxta with concave anterior margin, tip deeply rounded. Aedeagus short, slender, apex tapered, in the centre slightly constricted, caecum rounded. Without cornutus. Female genitalia: Compared with other European ennomine moths with unusual structure. Papillae anales short, wide, weakly sclerotized, setose. Length of apophyses posteriores 1 mm. Tergum A8 bent, strongly sclerotized, densely covered with setae. Apophyses anteriores very short, tapered. Lamellae ante- and postvaginalis inconspicuous. Ostium bursae small, funnel-shaped. Ductus bursae very long, narrow, membranous, twisted. Corpus bursae small, oval, without signum. Distribution and abundance: European-Turanian. Only known from a few localities in easternmost Austria, in southern Hungary and around Budapest, in one place in southern Slovakia and from Crimea. – Highly endangered due to destruction of its habitats. Usually found in limited numbers, despite some old records of abundant occurrences. Most records

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old, due to present day protection, but in Crimea still regularly recorded (V. Savchuk pers. comm.). In Hungary no observations from the last 30 years (B. Tóth pers. comm.). – Outside Europe only recorded from Tarbagatai mountains in eastern Kazakhstan (Kasy 1965).

Phenology: Univoltine: In eastern Austria from late October to early November with a few records from mid-October in autumns with especially cold weather conditions. In Crimea recorded in early and mid-November (Y. Budashkin & V. Savchuk pers. comm.). Flight period probably not lasting more than three weeks (Kasy 1965). In Crimea caterpillars found from mid-May to mid-June (Y. Budashkin & V. Savchuk pers. comm.). Hibernation as egg. Males flying by day until 2 o’clock in the afternoon, mainly from 10 o’clock (Y. Budashkin & V. Savchuk pers. comm.). Wingless females sitting in grass and other meadow vegetation. Biology: Larva polyphagous. Found on Seseli tortuosum (Y. Budashkin & V. Savchuk pers. comm.). Recorded on Centaurea jacea, C. scabiosa and the larger larvae also on Achillea asplenifolia (Kasy 1965). Reared on Inula and Tragopogon (Kasy 1965) and on Euphorbia virgata, E. stepposa, Centaurea diffusa and C. dealbata (Y. Budashkin & V. Savchuk pers. comm.). Eggs spirally placed around plant stems, similar to Malacosoma (Lasiocampidae) (see Text-fig. 17 in GME Vol. 1). Pupation in the ground in a silky cocoon (Y. Budashkin & V. Savchuk pers. comm.). Habitat: Xerothermophilous. In Austria in places with mixed dry and moist soil, in Hungary in a sandy area and a xerothermic slope. In Crimea on limestone steppe (V. Savchuk pers. comm.). In Europe a lowland species, in eastern Kazakhstan at about 1200 m altitude. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:ACL7591 (n=1 from Ukraine). Nearest species: North American Neoterpes ephelidaria (Hulst, 1886) (7.2%). 57

Tribus Gnophini Duponchel, 1845 Gnophini Duponchel, 1845 (Gnophites). Type species (of type genus Gnophos): Geometra furvata Denis & Schiffermüller, 1775. Aspilatini Duponchel, 1845 (Aspilatites). Type species (of type genus Aspilates): Geometra gilvaria Treitschke, 1825. Aspilatini is based on a misspelling of the genus-group name Aspitates Treitschke, 1825 (Aspilates is an unjustified emendation by Treitschke, 1827) as noted in Fletcher (1979). Aspilatini was included in the concept of the Angeronini (Holloway 1994). Morphology supports its inclusion in the Gnophini (our observation). Dasydini Duponchel, 1845 (Dasydites). Type species (of type genus Dasydia): Geometra torvaria Hübner, 1813. Tribe concept is based on Dasydia Guenée, 1845, a junior objective synonym of Sciadia Hübner, 1822. Sionini Duponchel, 1845 (Sionites). Type species (of type genus Siona): Phalaena lineata Sco­ poli, 1763. Included in the Gnophini by Herbulot (1962–1963), confirmed by Viidalepp et al. (2007), Wahlberg et al. (2010) and Skou & Sihvonen (2015). Angeronini Forbes, 1948. Type species (of type genus Angerona): Phalaena (Geometra) prunaria Linnaeus, 1758. Considered valid at tribal rank (for instance McGuffin 1981; Viidalepp 1996; Hausmann et al. 2004; 2011a; Vives Moreno 2014), but included as a junior synonym of Gnophini by Beljaev (2016). Synonymy is here confirmed, quoting structural similarities (see Diagnosis below) of adults, including for instance wing venation, male genitalia, structure of pupal cremaster and results from molecular phylogenetic analyses in which Angerona Duponchel, 1829 and Chariaspilates Wehrli, 1953 (Yamamato & Sota 2007) and Angerona and Charissa Curtis, 1826 grouped together (Sihvonen et al. 2011). The synonymy is supported by recent multi-gene analysis also (Murillo-Ramos et al. 2019). ‡ Psodinae Povolný & Moucha, 1955. Type species: Psodos quadrifaria (Sulzer, 1776). Name is not nomenclaturally available (Beljaev 2016), included in the Gnophini by Beljaev (2016). Diaprepesillini Kuznetzov & Stekolnikov, 1982. Type species: Diaprepesilla flavomarginaria (Bremer, 1864). Included in the Gnophini by Viidalepp (1996) and Beljaev (2016).

The Gnophini contain about 350 species in about 30 genera, but taxonomy is partly poorly resolved and the number of included genera is somewhat uncertain. Widely distributed in the Palaearctic region, numerous species combined with genus Gnophos and allied genera, and a few species extending into the Oriental region (for instance Wehrli 1939–1954; Viidalepp 1996; Scoble 1999; Kim et al. 2001); about 25 species in the Nearctic region and a few species extending into the Neotropics (see Remarks); about 10 species in the Arabian peninsula (Wiltshire 1990; Hacker & Hausmann 1999; Hausmann et al. 2016) and less than ten species are described from southern Africa (Vári et al. 2002; Krüger 2005). The Gnophini had been recognized as a valid Ennominae tribe in the literature until Hollo­ way (1994) proposed a very broad concept of Boarmiini, synonymizing numerous tribes into it. As regards the Gnophini, Holloway (1994: p. 7) concluded: ‘Not investigated in detail but probably subsumed by the concept of Boarmiini in this volume.’ This view is not supported by the morphology (Viidalepp et al. 2007) nor by molecular phylogenetic analyses, where Boarmiini and Gnophini were not related (Viidalepp et al. 2007; Wahlberg et al. 2010; Sihvonen et al. 2011; Jiang et al. 2017; Murillo-Ramos et al. 2019). In Sihvonen et al. (2011) the Boarmiini grouped together with the Macariini + Cassymini + Abraxini lineage, the Gnophini grouped together with the Angeronini (synonymized here). For these reasons we consider the Gnophini a valid tribe. external characters and abdomen:

Viidalepp et al. (2007) proposed the following potential synapomorphic characters for the Gnophini: fovea usually lacking in male forewings, male abdominal sternite A3 without a comb of stiff setae and male antennae are often pectinated

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Text-figs 26–28. Selected diagnostic characters (indicated) of the Gnophini. See text for other diagnostic characters. Text-fig. 26. Valva with process in costa usually present, often bearing a spine or group of spines (see genitalia plates in the Geometrid Moths of Europe, Vol. 5 for additional examples), Gnophos obfuscata (Denis & Schiffermüller, 1775), Finland: Savonlinna, 5–8. viii.1995, slide Sihvonen 2084 (coll. P. Sihvonen). Text-fig. 27. Pupal cremaster with setae D2 ‘bicornute’ (ventral view), Gnophos obfuscata (Denis & Schiffermüller, 1775), Italy: Monte Baldo, reared by B. Müller in 1990 (coll. ZSM). Text-fig. 28. Pupal cremaster with setae D2 separate and setae L1, D1, SD1 present (dorsal view), Angerona prunaria (Linnaeus, 1758), Hungary: Pilis Mountain, reared by B. Müller in 1985–1986 (coll. ZSM).

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to the apical segments. Externally the Gnophini are a variable group and clear defining characters in the facies have not been identified so far. Gnophos and allied genera are often grey, grey-brown or whitish grey, discal spots have pale centre and postmedial line is distinctly dentate, bordered with pale area on underside. Females are brachypterous in several species. Angerona, Aspitates and allied genera often have yellow coloration on the wings and forewing postmedial line is oblique, whereas Dyscia species are characterized by distinct, elongated discal spots and robust build. Tympanal organs are often rather small, they do not meet medially. Abdominal sclerites are without structural modifications, posterior margin of male 8th sternite is occasionally narrowed. Male genitalia: Viidalepp et al. (2007) proposed the following potential synapomorphic characters for the Gnophini: Uncus short and bird’s head-shaped. Costal projection on valva usually present, bearing a spine or group of spines (Text-fig. 26). Juxta is often H-shaped with short basal (dorsal) and longer distal (ventral) appendages. Other characteristic features include (our observations): fused gnathos with upturned medial element. Valva base is wide in some taxa. Juxta margins are often serrate or dentate. Vesica often bears one large or several minute cornuti. Sclerotized juxta is fused to aedeagus in some groups, for instance in Crocota Hübner, 1823. Female genitalia: No definitive, synapomorphic characters have been identified. Papillae ­anales are often round, ovipositor is not telescopic. Lamella antevaginalis is often horizontally elongated and rather large, sclerotized, but may be small, roundish and weakly sclerotized. Corpus bursae bears sclerotized, parallel ridges in some groups and it may have an appendix bursae with several, often slightly triangular signa. Signum is often definitive, located on anterior part of corpus bursae, laterally elongate and concave medially (not stellate), or small, or absent. Immature stages: Viidalepp et al. (2007) proposed the following potential synapomorphic character for the Gnophini: cremaster with setae D1, SD1, L1 and D2 present, setae D2 separated (Text-fig. 28). The character is characteristic for the Gnophini but not unique, because, notably the genus Gnophos has this character as bicornute (illustrated in Patočka & Turčani 2005), see Text-fig. 27. The larvae of Gnophos, Elophos and Dyscia, where information exists, have two distinct, horn-shaped dorsal projections on 11th segment (Gómez de Aizpúrua 2002; Porter 2010; Lepiforum 2018). The larvae of Hypoxystis pluviaria, Perconia strigillaria and Siona lineata may rest in coiled position (Lepiforum 2018). Biology: The Gnophini as a group do not appear to show preference for certain food-plants. Larvae have been recorded for instance on Betula, Vaccinium, Rubus (Elophos vittaria; Mikkola et al. 1989), Sedum, Knautia, Vicia (Gnophos furvata, G. obfuscata; Ebert 2003), cushion plants, especially Silene acaulis (Sciadia; Huemer & Hausmann 2009) and Thymus spp. (Charissa glaucinaria; Gómez de Aizpúrua 2002). Several species overwinter as (young) larva, but for majority of species overwintering stage is unknown. The Gnophini includes forest and open habitat species, but numerous groups are found in rocky habitats and adults of Gnophos and allied genera commonly rest on rocks during daytime, and the wing patterns match well such surfaces, making them difficult to see. Several groups are active by day, or facultatively diurnal such as Sciadia (Huemer & Hausmann 2009). The Gnophini species are found from sea-level up to 3300 metres, many species are restricted to high mountain habitats or in high latitudes above the treeline. Included European genera (several genera were treated in the Geometrid Moths of Europe, vol. 5 (Skou & Sihvonen 2015)): Angerona, Aspitates, Chariaspilates, Charissa, Cleorodes, 60

Crocota, Dicrognophos, Dyscia, Elophos, Gnophos, Hypoxystis, Odontognophos, Menophra, Perconia, Phthonandria, Psodos, Sciadia, Siona, Synopsia, Yezognophos. We are currently listing tentatively as ‘Gnophini of uncertain association’ the following three European genera: Dasypteroma, Megalycinia, Zernyia. Remarks: Taxonomy of the Gnophini is partly poorly resolved and a number of included genera are somewhat uncertain. Morphologically the core Gnophini (Gnophos and related genera) are rather homogeneous, but genera such as non-European Rhipignophos Wehrli, 1951 are distinctly different (see illustrations in Viidalepp 1988b). The Angeronini, including the Aspilatini fall within the concept of Gnophini, the association is supported by the morphology (see Diagnosis above) and molecular phylogenetic analyses (Yamamoto & Sota 2007; Sihvonen et al. 2011). Three mainly Nearctic genera are also structurally similar (illustrated for instance in McGuffin 1981) and based on the examination of the type species of each genus, we propose new tribal combinations (systematic position of other included species in the genera needs confirmation): Euchlaena Hübner, 1823, Lytrosis Hulst, 1896 and Xanthotype Warren, 1894 are transferred from Angeronini (McGuffin 1981; Hodges et al. 1983) to the Gnophini. In addition to the above-mentioned genera, the Gnophini in North America include also one Gnophos species: ‘Gnophos’ macguffini Smiles, 1979 which is here synonymized with Charissa turfosaria (classified in the Boarmiini in Hodges et al. 1983). Viidalepp (1996) included 46 genera in the Gnophini. We have not examined the systematic position of the non-European genera. Wehrli (1951) revised the genus classification of Gnophos Treitschke, 1825 and introduced numerous new genera and subgenera. Some authors have synonymized many of these under Gnophos (Scoble 1999), while others have retained most of these as valid (Viidalepp 1996). Selected Gnophini literature includes for instance Viidalepp (1988b), Kaila et al. (1996), Erlacher & Erlacher (2016; 2017a; 2017b), Erlacher et al. (2017).

Gnophos Treitschke, 1825 Gnophos Treitschke, 1825: in Ochsenheimer, F.: Die Schmetterlinge von Europa (Fortsetzung des Ochsenheimer’schen Werks). Vol. 5 (2): 429. Leipzig. Type species: Geometra furvata Denis & Schiffermüller, 1775.

Diversity and distribution: In appearance similar to some species of Elophos Boisduval, 1840, Charissa Curtis, 1826, Dicrognophos Wehrli, 1951, and of other Geometrid moth genera with species being perfectly adapted to rocks. The tentative listing of Gnophos species in Scoble (1999) representing an assemblage of species belonging to different genera (e.g. Dicrognophos Wehrli, 1951 and Rhipignophos Wehrli, 1951). After revision, the genus will probably comprise only 15 to 20 species, mainly distributed in South East Asia. In any case, a worldwide revision of the genus Gnophos is required. External characters and abdomen: European species medium- to large-sized, often with distinct transverse lines and discal spots and highly variable wing coloration. Antennae of males filiform. Hindtibia slender, significantly longer than hindfemur (Sauter 1990). Male genitalia: Uncus and gnathos rounded and blunt ending. Proximal third of costa of valva bulging, bearing one to three long, slender and bent spines. Valva with smooth margin, narrowed towards apex, setose. Juxta approximately heart-shaped. Aedeagus tubular, apically claw-shaped, pointed.

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Female genitalia: Antrum small with sclerotized inner convolution. Corpus bursae pipeshaped with more or less narrow proximal and widened distal part. Biology: Larvae of European Gnophos species polyphagous, feeding on a variety of herbaceous plants, shrubs and broad-leaved trees. Larva nocturnal, hiding near food-plants during daytime. Adults nocturnal, resting on rocks or lumps of rock during daytime.

9. Gnophos furvata (Denis & Schiffermüller, 1775) Geometra furvata Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 108 (Austria, Vienna district). Syntype(s) (lost). ‡ Gnophos furvata [“f.”] cinerascens Turati, 1919: Il Naturalista Siciliano 23: 319, pl. 4, fig. 27 (Italy, Calabria, San Severino). Syntype(s) (lost?). Infrasubspecific. ‡ Gnophos furvata [“f.”] meridionalis Wehrli, 1924: Dt. ent. Z. Iris 38: 84, pl. 1, fig. 5 (Switzer­ land: Tessin, southern valleys of Graubünden, Lostallo, Grono, South Tyrol). Syntypes (ZFMK, not examined). Synonym of Geometra furvata Denis & Schiffermüller, 1775 in S ­ coble (1999) and Flamigni et al. (2016).

External characters and abdomen: Wingspan 38–52 mm. Wings broad with termens undulating. Ground colour usually grey or brownish in various shades, with scattered darker scales. Transverse lines denticulate, sometimes indistinct and blurred, medial band more or less darkened, discal spots dark, often indistinct. Underside of wings less marked, pale. Antennae filiform in both sexes. Frons, collar, thorax and abdomen concolorous with wings. Variation: The ground colour of the wings can vary between different geographical regions, however, according to Leraut (2009) and Flamigni et al. (2016) a distinction into separate subspecies is not justified. Even among local forms, there are always specimens resembling the nominotypical subspecies. Male genitalia: Uncus and gnathos rounded and blunt ending. Costa of valva with two long, slender and bent spines on a bulge in proximal third. Valva with smooth margin, narrowed towards apex, setose. Juxta approximately heart-shaped. Aedeagus tubular, apically clawshaped, pointed distal end, without cornuti. Female genitalia: Antrum small, twice as long as wide, with sclerotized inner convolution. Membranous part of ductus bursae short. Corpus bursae membranous, proximal part pipelike with fine longitudinal folding, distal part widened, bearing a small signum. Distribution and abundance: European. Widely distributed in central and southern Europe, in the north to the Netherlands and central Germany, in the east to Bulgaria and Ukraine, in the south to Italy, and in the west to the northern half of the Iberian peninsula as well as in the Sierra Nevada (Viidalepp 1996; Huemer et al. 2009; Leraut 2009; Redondo et al. 2009; Malkiewicz 2012; Flamigni et al. 2016). – Widespread, sometimes in numbers. Phenology: Univoltine. From late June to September, with main flight period from July to August (Redondo et al. 2009; Malkiewicz 2012; Flamigni et al. 2016). Overwintering as larva, last instar from late April to early June. Adults nocturnal, active at twilight. Biology: Larva polyphagous. Found among others on Clematis vitalba, Cornus sanguinea, Coronilla coronata, Helianthemum nummularium, Hippocrepis comosa, Ligustrum vulgare, Medicago falcata, Prunus spinosa, Sanguisorba minor and Thymus serpyllum (Bartsch 2003) as well as on Medicago falcata (S. Erlacher own observation). Found also under plants of Artemisia, Lathyrus, Oxyria, Sedum, and Viola (King & González-Estébanez 2015). 62

Recorded on Chenopodium, Clinopodium, Cotoneaster, Crataegus, Dorycnium, Fragaria, Genista, Hieracium, Hyoscyamus, Lonicera, Mespilus, Onobrychis, Origanum, Phillyrea, Plantago, Polygonum, Potentilla, Rubus, Rosa, Rumex, Stachys, Teucrium, Vaccinium, Viburnum and Vicia (Leraut 2009; Redondo et al. 2009; Flamigni et al. 2016). Larva found by night with flashlight after overwintering on coppices and small bushes of Prunus spinosa (south-western Germany: M. Leipnitz pers. comm.). Reared on Rumex crispus, R. acetosella (King & González-Estébanez 2015), on Polygonum and Taraxacum, and, after overwintering, on different species of young broad-leaved shoots, such as Carpinus, Corylus, Ulmus, Celtis and Symphoricarpos (Flamigni et al. 2016). Larva hiding on ground amongst low plants during daytime (Bartsch 2003; King & González-Estébanez 2015). Detailed description of larva in King & González-Estébanez (2015). Pupation in a very loose spinning under moss and withered plants on the soil (rearing observation: M. Leipnitz pers. comm.). Adults resting on rocks during daytime. Nectar-feeding observed on flowers of Echium, Eupa­ torium cannabinum, Knautia arvensis, Sedum, Sambucus ebulus (Bartsch 2003), Campanula trachelium (W. Wolf pers. comm.) and on Scabiosa (S. Erlacher own observation). Habitat: Xerothermophilous. Found in open dry hills, steep slopes and calcareous rock habitats with sparse vegetation. From 70 m up to 2100 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC6119 (n=20 from Germany, Austria, France, Italy, Croatia, Bulgaria). Genetically homogeneous. Nearest species: Gnophos obfuscata (5.0%).

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10. Gnophos obfuscata (Denis & Schiffermüller, 1775) Geometra obfuscata Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 108 (Austria: Vienna district). Syntype(s) lost. Phalaena myrtillata Thunberg, 1788: Museum naturalium Academiae Upsaliensis 6: 75 (Sweden). Lectotype ♂ (UZIU, not examined). Retained synonym by Scoble (1999), validated at subspecies rank by Skinner (2009), downgraded here to synonymy with Geometra obfuscata Denis & Schiffermüller, 1775 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos obfuscaria [sic!] [“v.”] lutescentaria Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1:168 (Caucasus,?Armenia). Syntypes, no type material mentioned in the original description. Validated at subspecies rank (Scoble 1999), downgraded here to synonymy with Geometra obfuscata Denis & Schiffermüller, 1775 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos myrtillata [“var.”] nivea Schawerda, 1913: Verh. zool.-bot. Ges. Wien 63: 164 (Bosnia-­ Herzegovina: Vucijabara, 1200 m). Syntypes 7♀ (NHMW, not examined). Validated at subspecies rank (Scoble 1999 downgraded here to synonymy with Geometra obfuscata Denis & Schiffermüller, 1775 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos myrtillata marsicaria Dannehl, 1933: Ent. Z., Frankf. a. M. 47 (17): 140 (Italy: southern Abruzzo, Monte Rotella, 1600 m). Holotype (not examined). Validated at subspecies rank (Scoble 1999) and Flamigni et al. (2016), downgraded here to synonymy with Geometra obfuscata Denis & Schiffermüller, 1775 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos myrtillatus [sic!] androgynus Reisser, 1936: Ent. Rdsch. 53 (10): 146, pl. 4, figs A3, A4, A5, A6, C3, C4, C5 (Spain: Sierra de Gredos). Syntypes 9♂3♀ (NHRS, not examined). Validated at subspecies rank (Scoble 1999), downgraded here to synonymy with Geometra obfuscata Denis & Schiffermüller, 1775 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation.

External characters and abdomen: Wingspan ♂ 39–42 mm, ♀ 33–36 mm. Ground colour varying from dark grey to pale brownish grey, sometimes yellowish sprinkled. Wing pattern with transverse lines and discal spot variably expressed, often distinct. Females much smaller and narrow-winged, lighter grey and more distinctly patterned than males. Wing termen slightly undulating. Underside of wings pale and dull. Frons, collar, thorax and abdomen concolorous with wings. Variation: Ground colour of adults varying from whitish to dark grey. Specimens from granite rocks (Sierra de Gredos, Spain) with a mosaic of yellowish specks on slightly variable ground, specimens on limestone (central Apennines) with very light, whitish grey ground colour with slight yellow tinge (Flamigni et al. 2016). In northern British Isles, ­Fennoscandia and parts of the Baltic, males tend to be smaller and darker, with less distinct pattern. ­Females lighter grey, hardly smaller than males (Wehrli 1953). Male genitalia: Uncus and gnathos rounded and blunt ending. Proximal third of costa of valva bulging, bearing one to three long, slender and bent spines. Valva with smooth margin, narrowed towards apex, setose and with thorn-shaped sclerotization. Juxta approximately heart-shaped. Aedeagus tubular, apically claw-shaped, pointed, bearing a variety of short and straight cornuti in spiral position. Female genitalia: Antrum small, about as long as wide, with sclerotized inner convolution. Corpus bursae proximally sclerotized, distal part widened, without signum.

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Distribution and abundance: Euro-Caucasian. Widely distributed in the European Alps, the Iberian peninsula, in central Apennines, the Balkan peninsula, the Caucasus, the north of the British Isles, in Fennoscandia and parts of the Baltic (Wehrli 1953; Skou 1986; Viidalepp 1996; Huemer et al. 2009; Redondo et al. 2009; Skinner 2009; Flamigni et al. 2016). – Widespread, sometimes in numbers.

Phenology: Univoltine. From June to late August, rarely September (Redondo et al. 2009; Flamigni et al. 2016). Larva from early August to early October, after hibernation (in L2–L3: M. Leipnitz pers. comm.) from late April to late June (Flamigni et al. 2016). Adults nocturnal, readily attracted to light, resting under overhanging plant cushions and on rocks near the ground during daytime (M. Leipnitz pers. comm.). Biology: Larva polyphagous on herbaceous plants and shrubs. Found under and in cushions of Dryas octopetala, Salix alpina, Salix herbacea, Potentilla spp. (western Austria: M. Leipnitz pers. comm.). Recorded among others on Calluna vulgaris, Campanula, Cytisus scoparius, Genista (especially on G. tinctoria, G. anglica), Gentiana (especially on G. lutea), Glyceria, Helianthemum, Lotus corniculatus, Plantago, Poa, Polygonum, Rubus, Rumex, Sedum telephium, Taraxacum, Vicia (especially on V. cracca), Viola (Redondo et al. 2009; Flamigni et al. 2016). Reared on Erica carnea (pulp), Erica tetralix, Genista hispanica, Vaccinium myrtillus (Flamigni et al. 2016), on Taraxacum, Salix myrtilloides, Salix, Alchemilla, Potentilla and Prunus spinosa (M. Leipnitz, pers. comm.). Oviposition in small groups of approx. 8–12 grey-green eggs on the underside of leaves, e.g. of Alchemilla (M. Leipnitz pers. comm.). Eggs found on leaves of Hippophae rhamnoides (Flamigni et al. 2016). According to Denis & Schiffermüller (1775) larva also feeding on Alnus glutinosa. Pupation in a very loose cocoon or under moss and withered plants (rearing observation: M. Leipnitz pers. comm.).

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Habitat: Found in open rocky areas and scree with short vegetation, dwarf shrub heaths with rich vegetation, stocks of Rhododendron hirsutum, R. ferrugineum, Erica herbacea and Pinus mugo, block and rock screes with Salix, Dryas octopetala, Valeriana tripteris, V. montana, Arabis, sunny and rocky places with scattered Picea, Abies and Larix, and with herbaceous, species-rich undergrowth (e.g., Vaccinium mytillus, V. vitis-idaea), alpine pastures with rich, tall, herbaceous vegetation (M. Leipnitz pers. comm.). Montane, up to 3000 m, occasionally below 1000 m (Redondo et al. 2009; Flamigni et al. 2016), in northern Alps usually up to 2200 m (M. Leipnitz pers. comm.). In the Inner Hebrides of Scotland occurring on sea-cliffs down almost to sea-level (M. Corley pers. comm.). Similar species: Females of G. obfuscata can be confused with Charissa ambiguata or Elophos dilucidaria. In some cases, an examination of the genitalia will be necessary. Genetic data: Five BINs: BOLD:ACE7309, BOLD:AAB8516, BOLD:AAB8517, BOLD:ABZ5475, BOLD:ACC5028 (n=41 from Finland, Germany, Austria, Switzerland, Italy, Macedonia, Bulgaria, Georgia, Turkey). Genetically very heterogeneous, BINs at minimum pairwise distances of 1.1–2.2%, a singleton from northern Spain (Cantabria; P. ­Huemer, pers. comm.) more strongly diverging (4.2%; BOLD:ACC5028), either not conspecific or referring to a NUMT. Genetic heterogeneity requiring further integrative analysis. Nearest species: Gnophos furvata (5.0%).

Charissa Curtis, 1826 Charissa Curtis, 1826: British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland. Vol. 3: object number: 105. Type species: Geo­ metra obscurata Denis & Schiffermüller, 1775, originally designated as “Geometra obscuraria Hüb.” (unjustified emendation). The case is yet to be decided by the ICZN. Until then, the name Charissa is considered protected. ‡ Hyposcotis Hübner, 1825: Verzeichniß bekannter Schmettlinge [sic!]: 314. Type species: Geo­ metra mucidaria Hübner, 1799, by subsequent designation by Prout (1904). See remarks under Charissa. The case is yet to be decided by the ICZN. Until then, the name Hyposcotis is considered suppressed.

Diversity and distribution: Charissa currently contains about 70 species in the Palaearctic and Oriental regions, one species also occurs in the Nearctic. In external appearance, species of Charissa are very similar to those of Gnophos Treitschke, 1825, Elophos Boisduval, 1840, Dicrognophos Wehrli, 1951, and of some other geometrid moth genera whose species are perfectly adapted to rocks. For the same reason, it is often difficult to distinguish (isolated) subspecies from other local adaptations (infrapopulational forms and local forms with clinal transitions towards neighbouring populations). Previous taxonomy, which validated about twice as many subspecies than species for the European fauna (Scoble 1999) is critically revised here. External characters and abdomen: Medium-sized, transverse lines and discal spots often distinct, postmedial line often dentate. Antennae of males ranging from filiform to shortly bipectinate, both character states even occurring in the same subgenus (e.g. Cnestrognophos, Wehrli, 1951 and Euchrognophos Wehrli, 1951). Hindtibia clavate, thickened, only slightly longer than hindfemur (Sauter 1990). Wing venation very variable, even within the same species, and not at all constant as suggested by Sauter (1990).

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Male genitalia: Juxta divided into two arms, without terminal spines. Valva costa more strongly sclerotized than rest of valva, setose but with not more than one or two spines. Female genitalia: Bursa copulatrix with sclerotized longitudinal folds. Biology: Larvae of European species feeding on several low growing plants, active at night, hiding near foodplants during daytime. Pupation on the ground. Adults active at night, both sexes attracted to light, resting on rocks or lumps of rock during daytime. Remarks: According to article 23.9.3. ICZN, the prevailing usage of the genus name Charissa Curtis, 1826 instead of Hyposcotis Hübner, 1825 is to be maintained while a case submitted by Erlacher & Erlacher (2017a) is under consideration (nomen protectum).

Subgenus Charissa Curtis, 1826 Charissa Curtis, 1826: British entomology; being illustrations and descriptions of the genera of insects found in Great Britain and Ireland. Vol. 3: object number 105. Type species: Geo­ metra obscurata Denis & Schiffermüller, 1775. Lycognophos Wehrli, 1951, Lambillionea 51: 11. Type species: Geometra obscurata Denis & Schiffermüller, 1775. Originally proposed as subgenus of Gnophos Treitschke, 1825. Junior objective synonym of Charissa Curtis, 1826. ‡ Lysognophos Wehrli, 1953, in Seitz, Gross-Schmett. Erde 4 (suppl.): 582. Incorrect subsequent spelling.

Diversity and distribution: Currently, Charissa s. str. is a monotypic subgenus with Euro-­ Caucasian distribution. Male and female genitalia: see below. Remarks: Wehrli (1951; 1953) treated ‘Gnophos’ graecaria Staudinger, 1871 (= ‘Gnophos’ certhiatus Rebel & Zerny, 1931; see below) and ‘Gnophos’ idrisarius Zerny, 1934 as members of Gnophos, subgenus Lycognophos Wehrli, 1951 which is a junior objective synonym of Charissa Curtis, 1826. The taxon idrisarius was still listed under Gnophos in Scoble (1999). Nevertheless, our study has shown that C. graecaria belongs to the subgenus Trilobignophos Wehrli, 1951, while the systematic position of C. idrisarius within the genus Charissa remains unclear.

11. Charissa obscurata (Denis & Schiffermüller, 1775) Geometra obscurata Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 108 (Austria, Vienna district). Syntype(s) (lost). Geometra obscurata pullaria Hübner, 1799: Samml. Eur. Schmett. 5, Geometrae (1): pl. 27, fig. 145 (Europe). Syntype(s) (not examined). Validated at subspecific rank (Scoble 1999), herewith downgraded to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos obscurata [“ab. (v)”] calceata Staudinger, 1861: In: Staudinger & Wocke: Catalog der Lepidopteren Europa‘s und der angrenzenden Länder 1: 72 (Europe). Syntype(s) (MNHU, not examined). Validated at subspecific rank (Scoble 1999), herewith downgraded to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos obscurata [“v. (et ab.?)”] argillacearia Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 166 (southern France). Syntype(s) (MNHU, not examined). Validated at subspecific rank (Scoble 1999), herewith downgraded) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.).

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Gnophos obscurata lafauryata Oberthür, 1913: Études Lépid. comp. 7: 308, pl. 180, figs 1760, 1761 (south-western France: Dax and Bordeaux). Syntypes (ZFMK, not examined). Validated at subspecific rank (Scoble 1999), herewith downgraded to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos obscurata bellieri Oberthür, 1913: Études Lépid. comp. 7: 308, pl. 180, fig. 1759 (Corsica). Lectotype ♂ (pl. 2, figs 11h, 11i, hereby designated) (ZFMK, examined, label data: Corsica). Validated at species rank in Scoble (1999) and Leraut (2009), validated at subspecific rank by Flamigni et al. (2016). Here, we accept subspecies status, based on differences in wing pattern and divergences in the COI gene (see Genetic data).

External characters and abdomen: Wingspan 25–33 mm. Ground colour of wings usually brown, but ranging from blackish to whitish. Fringes slightly lighter. Transverse lines on all wings dark and clearly dentate, discal spots dark. Hindwing margin undulate. Underside of wings less distinctly marked, lighter, with uniform ground colour. Frons, collar, thorax and abdomen concolorous with wings. Antennae filiform in both sexes. Variation: The ground colour of the adults often varies with the habitat and the colour of the rocks, on which they rest during daytime, even within very short geographical distances and often with clinal transitions, without representing isolated subspecies (see Warnecke 1939). Even among local forms, there are always specimens resembling the nominotypical subspecies. Subsp. bellieri Oberthür, 1913 (Corsica and Sardinia): more sharply lined and contrasting in wing pattern. Male genitalia: Uncus pointed. Gnathos thin and blunt. Ventral valva margin medially constricted. Long thorns on a pedicle at the base of the costa of valva. Juxta deeply divided into two long and narrow arms, distally slightly bent inwards. Aedeagus cylindrical with numerous cornuti of about half length of aedeagus. Female genitalia: Antrum slightly v-shaped, well sclerotized. Corpus bursae pipe-shaped, with strongly sclerotized, spinose longitudinal folding, and with lateral appendix bursae (dome-like, ruffled structure bearing a set of stout and large teeth). Distribution and abundance: Euro-Caucasian. Widely distributed in Europe, from southernmost Iberian peninsula to the Baltic States, and from Fennoscandia to the Mediterranean (Fajčík & Slamka 1996; Viidalepp 1996; Mironov et al. 1998; Nestorova 1998; Huemer et al. 2009; Leraut 2009; Redondo et al. 2009; Malkiewicz 2012; Flamigni et al. 2016). In the south to Italy including Sardinia and Sicily and Greece including Crete. – Local in many parts of the distribution area, sometimes common. – Outside Europe known from Turkey and the Caucasus (Georgia, Armenia and Azerbaijan) (Viidalepp 1996). Phenology: Univoltine. From July to September (Bartsch 2003; Erlacher 2019), in southern Europe slightly earlier, from June to September (Redondo et al. 2009). Overwintering as small larva (L2 or L3; M. Leipnitz pers. comm.). Caterpillars found from March to mid-May (Bartsch 2003; Erlacher 2019). Adults active at night, readily attracted to light. Adults rest on rocks and under stones during daytime, easily flushed up from dry logs lying on the ground (‘wood pile’) (J. Gelbrecht pers. comm.). Biology: Larva polyphagous. Found on Helianthemum nummularium, Hippocrepis comosa, Campanula rotundifolia, Teucrium montanum and Vicia (Bartsch 2003) as well as on Loni­ cera periclymenum (M. Leipnitz pers. comm.) and on Lotus corniculatus (S. ­Erlacher own observation). Recorded on Armeria, Artemisia, Bupleurum, Calluna, Crataegus, Dorycnium, 68

Erica, Euphorbia, Fragaria, Galeopsis, Geranium, Gypsophila, Helianthus, Lychnis, Origanum, Plantago, Potentilla, Rubus, Rumex, Sanguisorba, Saxifraga, Scabiosa, Sedum, Silene, Stellaria, Thymus and Vaccinium (Skou 1986; Bˇe lín 2003; Redondo et al. 2009; Skinner 2009; Flamigni et al. 2016). Reared on Lonicera tatarica (M. Leipnitz, pers. comm.). Larvae of subsp. bellieri found on Genista sardoa (M. Leipnitz pers. comm.). Pupation on the ground among moss and plant parts in a fine, light spinning consisting of just a few threads (rearing observation; M. Leipnitz pers. comm.). Habitat: On heaths, fringes, dry hills, slopes and rocks, in all kinds of dry scrub- and woodland as well as in many other places, even in urban regions, preferably on calcareous and sandy soil (Skou 1986; Bartsch 2003; Redondo et al. 2009). From sea-level in the north up to 2000 m in the south (Nestorova 1998; Redondo et al. 2009). In southern Europe preferring montane habitats, with rare and local occurrence in the lowlands. Subsp. bellieri from 400 m up to 1500 m (Flamigni et al. 2016). Similar species: Charissa (Kemtrognophos) zeitunaria (Staudinger, 1901) is often larger (wingspan 28–35 mm), transverse lines on all wings usually less strongly dentate. Male and female genitalia of both species completely different. Genetic data: BINs: BOLD:ACE8297, BOLD:ABZ5552, BOLD:AAB5231, BOLD:ACF2016 (n=42 from Finland, United Kingdom, Germany, Austria, Spain, Italy, Croatia, Macedonia, Greece, Turkey). Genetically very homogeneous in the main cluster, but both the Iberian (BOLD:AAB5231; n=3; distance 1.7%) and the Sicilian cluster (BOLD:ACF2016; n=5; distance 3.3%) strongly diverging, thus suggesting diversity at least at subspecies level. One singleton from southern Italy (BOLD:ABZ5552) possibly referring to a pseudogene. Subsp. bellieri: BIN: BOLD:ACF2015 (n=7 from Sardinia and Corsica). Genetically homogeneous on each island (variation 0.0%), diverging from each other by 1.1%, both diverging from all

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other populations of C. obscurata at a minimum distance of 1.5%. Nearest species: North African C. crosi (Thierry-Mieg, 1910) (4.1%) and C. idrisarius (Zerny, 1934) (4.6%). Remarks: C. obscurata tends to develop substrate-adapted forms that recur at distant ­locations. In all cases, however, descriptions of subspecies are based on colour-deviant individuals within more or less uniform populations. This applies to the following individual forms, which are therefore synonymized with the nominotypical subspecies here: Gnophos obscurata argillacearia Staudinger, 1871 (syn. n.); Gnophos obscurata calceata Staudinger, 1861 (syn. n.); Gnophos obscurata lafauryata Oberthür, 1913 (syn. n.); Geometra obscurata pullaria Hübner, 1799 (syn. n.).

Subgenus Costignophos Wehrli, 1951 Costignophos Wehrli, 1951: Lambillionea (51): 23. Type species: Geometra pullata Denis & Schiffermüller, 1775.

Diversity and distribution: The subgenus Costignophos contains three species which all ­occur exclusively in Europe. Male genitalia: Uncus and gnathos well developed, beak- or hook-shaped. Juxta deeply divided. Saccus relatively long and stout. Valva with well sclerotized ventral margin. Costa of valva sclerotized distal end exceeding tip of valva. Aedeagus tubular to conical with only one cornutus, 1/3–1/2 length of aedeagus. Female genitalia: Antrum short and flat. Corpus bursae pipe-shaped, with strongly sclerotized, spinose, longitudinal folding, laterally with appendix bursae (dome-like structure with ring-shaped folds, and a set of stout and large teeth inside). Remarks: A review of this subgenus was given by Rezbanyai-Reser (1986).

12. Charissa crenulata (Staudinger, 1871) Gnophos crenulata Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 167 (Spain, Andalusia). Holotype ♀ (ZSM, examined). Gnophos pullata pyrenaica Oberthür, 1913: Études Lépid. comp. 7: 316, pl. 183, figs 1793, 1794 (France, East Pyrenees: Vernet-les-Bains). Lectotype ♀ (pl. 3, figs 12k, 12l, hereby designated) (ZFMK, examined, label data: Vernet [-les-Bains]). Validated at subspecific rank by Scoble (1999) and by Redondo et al. (2009), subspecies rank here tentatively accepted, requiring further study including DNA barcoding. Gnophos crenulatus aragonensis Zerny, 1927: EOS, Rev. Esp. Ent. 3 (4): 426, pl. 10, fig. 32 (Spain, Aragon: Albarracín, Valdovecar). Syntypes ♂♀ (NHMW, not examined). Validated at subspecific rank Scoble (1999), retained as synonym in Redondo et al. (2009), here tentatively accepted at subspecies rank, requiring further study (see Genetic data). Gnophos avilarius Reisser, 1936: Ent. Rdsch. 53 (9): 136, pl. 4, figs A1, A2 (Spain, Sierra de Gredos: Garganta de las Pozas). Syntypes ♂♀ (NHMW, not examined). Previously, validated at species rank (Scoble 1999; Leraut 2009; Redondo et al. 2009), herewith downgraded to subspecific rank of C. crenulata (stat. n.). Gnophos crenulatus [sic!] tormoi Expósito Hermosa, 1996: Saturnia 7: 14 (Spain, Alicante: ­Umbría de la Aitana). Holotype ♀ (Expósito, examined). Not listed in Scoble (1999), retained as synonym in Redondo et al. (2009) without implementing this nomenclaturally (syn. n.).

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External characters and abdomen: Wingspan 28–36 mm. Ground colour of wings usually ash-grey, but ranging from brownish to yellowish white. Fringes concolorous or slightly lighter. Transverse lines on all wings usually clearly visible, dark and dentate, discal spots usually unfilled. Underside of wings with darker grey terminal area, with a conspicuous, indented white postmedial line running across fore- and hindwings. Terminal area much darker and provided with light spots (on forewings at apex and in centre), creating strong contrast on wing underside. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: Ground colour very variable depending on colours of local substrates. See subspecies. Subsp. aragonensis Zerny, 1927 (Albarracín): Upperside of wings lighter grey, slightly cream-coloured, with well contrasted transverse lines. Terminal area bluish grey, with distinct terminal dots. Underside of wings darker, with blackish subterminal line. Subsp. avilarius Reisser, 1936 (Sierra de Gredos): Ground colour composed as an assemblage of grey, olive-tinged brown, ochreous and white. Ante- and postmedial lines clearly serrated. Discal spots large and clearly unfilled. In the middle of the terminal area a light spot. Fringes speckled. Underside much paler than in nominotypical subspecies. Subsp. pyrenaica Oberthür, 1913 (Pyrenees): Ground colour ochreous-grey. Medial area light orange to light reddish brown. Ante- and postmedial lines, fringes and underside largely as in the nominotypical subspecies. Male genitalia: Uncus arched conically, distally rounded. Gnathos from base to distal tip slim and slightly curved. Juxta divided, v-shaped. Juxta arms slender, slightly rounded, pointed at distal end. Costa of valva well sclerotized, distal end slightly curved, ventral margin well sclerotized. Saccus relatively long and stout. Aedeagus conically tapered, with a stout, narrow cornutus. Female genitalia: Antrum well sclerotized, quadrangular, margins slightly rounded, much wider than high. Distribution and abundance: West Mediterranean. Endemic to the Iberian peninsula (­Redondo et al. 2009) and in the Canigou in France (leg. L. Taurand). Phenology: Univoltine. From mid-July to late-August, occasionally until September (Zerny 1927a; Redondo et al. 2009). Most likely overwintering as larva (S. Erlacher, observations under rearing conditions). Adults nocturnal, attracted to light. Biology: Larva recorded on Sedum (Redondo et al. 2009) and other low-growing plants (Leraut 2009). Larva of subsp. avilarius reared on Rumex acetosella (Redondo et al. 2009), Lonicerea tatarica, Taraxacum and Clematis vitalba (M. Leipnitz pers. comm.). Pupation on the ground among moss and plant parts in a fine, light spinning (rearing observation; M. Leipnitz pers. comm.). Habitat: Xerothermophilous, montane. Mainly in mountain terrains and woodland areas (Leraut 2009; Redondo 2009). In central Portugal in stand of dense young pines with abundant undergrowth, found individually sitting on trunks (M. Leipnitz pers. comm.). From 700 m up to 2500 m, in the Sistema Central the subsp. avilarius from 1300 m up to 2200 m (Redondo et al. 2009). Subsp. pyrenaica in the Canigou from 1000 m up to 1600 m (leg. L. Taurand). 71

Similar species: C. italohelveticus and C. pullata, both allopatric with C. crenulata according to current knowledge (sympatry of C. crenulata and C. pullata, however, cannot be completely ruled out). C. crenulata differs from C. pullata in some phenotypic characters: The transverse lines on all wings are usually clearly visible and not broken into single dots as in C. pullata. On the underside of the wings the white transverse line is usually wider and medially more serrated. The dark terminal area contains light spots giving a contrast-rich ­appearance to the underside. While these two species are often distinguishable extern­ ally, discrimination from C. italohelveticus is hardly possible based on external characters and ­often requires dissection of the genitalia. In male genitalia of C. crenulata, the juxta is v-shaped (as it is also in C. italohelveticus), whereas it is u-shaped in C. pullata. The costa of the valva is usually slightly curved distally whereas it is more or less straight in C. italo­ helveticus (and clearly bent in C. pullata). In addition, the relative length of the single, almost straight, pointed cornutus is also different: The largest aedeagus of C. italohelveticus bears the shortest cornutus (length ratio of cornutus versus aedeagus 1:3.4 in C. italohelveticus; 1:2.55 in C. crenulata). In female genitalia of C. crenulata, the antrum is quadrangular with slightly rounded margins and is clearly wider than high, in contrast to the antrum of C. italohelveticus showing almost the same measures for height and width. Genetic data: BIN: BOLD:ACE7619 (n=12 from Spain). Genetically heterogeneous, populations from Andalusia (nominotypical subspecies) diverging from those of eastern Spain (n=2; subsp. aragonensis) at a distance of 1.2%. Populations from Sierra de Gredos (subsp. avilarius) BIN-sharing but diverging by only 0.8% from both subsp. aragonensis and nominotypical subspecies. Thus, genetic data support downgrading the taxon from species rank to subspecies of C. crenulata (see Remarks). Nearest species: C. italohelveticus (2.2%), C. pullata (1.7%). Remarks: Redondo et al. (2009) already recognized only pyrenaica as reasonably distinct subspecies of C. crenulata. Furthermore, they still treated avilarius as separate species as 72

also considered by Rezbanyai-Reser (1986) and Leraut (2009). However, current DNA barcode analyses show that avilarius is genetically much more similar to the nominotypical subspecies of crenulata than to the other two species of this subgenus. Based on the genetic evidence, on its allopatric distribution and on the almost complete lack of differential features in male and female genitalia, we herewith downgrade the taxon to subspecific rank of C. crenulata: Charissa crenulata avilarius (Reisser, 1936) (stat. n.). The taxon aragonensis Zerny, 1927 is based on specimens collected near Albarracín. Subspecific status was accepted by Scoble (1999) but not by Redondo et al. (2009), its status requires further clarification. The taxon tormoi Expósito Hermosa, 1996 was described based on four specimens from Umbría de la Aitana (Alicante). According to Expósito Hermosa (1996) it is more similar to subsp. pyrenaica than to aragonensis. However, no further differential characters were given in his description. This may just be a substrate-adapted individual form which is here synonymized with the nominotypical subspecies (syn. n.) in accordance with a similar suggestion in Redondo et al. (2009).

13. Charissa italohelveticus (Rezbanyai-Reser, 1986) Gnophos italohelveticus Rezbanyai-Reser, 1986: Nota lepid. 9 (1–2): 102, figs 6 (3), 8 (3), 10 (6), 11 (4), 12 (4) (Italy, Lugano TI: Mount Brè, Vetta, 920 m). Holotype ♂ (NML, not examined).

External characters and abdomen: Wingspan 29–38 mm. Ground colour of wings usually pale to darker ash-grey, sometimes with yellowish scales. Fringes concolorous or slightly lighter, slightly chequered. Transverse lines on all wings distinct and strongly jagged, on veins sometimes as strokes, discal spots large, mostly unfilled. Underside of wings with darker grey terminal area, with a conspicuous, indented white postmedial line running across fore- and hindwings. Hindwing underside with basal and medial area powdered with scales of monotonous grey, lighter than those of forewings, terminal area much darker and provided with light spots (on forewings only at apex and in centre), creating strong contrast on wing underside. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: Ground colour of the adults is very variable depending on the colour of the local substrate. Male genitalia: Uncus helmet-shaped, anteriorly and posteriorly bulging. Gnathos long and slim. Juxta divided, v-shaped, juxta arms relatively thick and stout, anterior processes clubshaped at tip. Costa of valva well sclerotized, distally more or less straight. Aedeagus long and slender, with a stout, narrow cornutus. Female genitalia: Antrum well sclerotized, margins slightly rounded, height almost the same as width. Distribution and abundance: South European. Distributed in easternmost France (Haute-­ Savoie), southern Switzerland (around Geneva, Valais, Ticino, Val Mesolcina and Valle di Poschiavo), northern Italy, Austria (Styria) (Rezbanyai-Reser 1986; SwissLepTeam 2010; ­Huemer 2013; Flamigni et al. 2016). Also known from north-western Greece (colls E. ­Friedrich, J. Viehmann) and from Peloponnese (coll. J. Gelbrecht).

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Phenology: Univoltine. From July to late September, mainly from late July to late August (Rezbanyai-Reser 1986). Overwintering as larva. Adults nocturnal, rarely flushed up from vegetation during daytime, but found sitting on trunks. Biology: Foodplants unknown from nature. Reared on Lotus, Vicia, Trifolium (Rezbanyai-­ Reser 1986), Thymus (Flamigni et al. 2016) and Lonicera tatarica, during hibernation also on Sanguisorba minor and Taraxacum (M. Leipnitz pers. comm.), successfully reared on potted plants (mixture of grasses, Plantago, partly withered). In autumn, the young caterpillars prefer withered parts of plants. After overwintering, they especially like to feed on Poa, but also on slightly withered Taraxacum and Plantago (Ziegler 2019). Pupation in a loose cocoon on the ground among moss, plant parts and small stones in a fine, light spinning (rearing observation; M. Leipnitz pers. comm.). Adults barely attracted to light, found at night when feeding on flowers, e.g. on Origanum (Ziegler 2019). Habitat: Xerothermophilous, montane. Inhabiting open habitats, bushy, warm, sunny and rocky slopes as well as forest edges (Rezbanyai-Reser 1986) and in warm, shady and bushy stands of Quercus pubescens, Juniperus and Pinus (M. Leipnitz pers. comm.), also in slightly cooler places within xerothermic habitats (Ziegler 2019). From 90 m up to 1000 m in the Alps (Rezbanyai-Reser 1986; Flamigni et al. 2016), up to 1050 m in Greece (J. Viehmann pers. comm.). Sometimes sympatric with C. pullata (Ziegler 2019). Similar species: Charissa crenulata and C. pullata (see species no. 12 and 14). Genetic data: BIN: BOLD:ABZ3077 (n=6 from France, Switzerland, Austria, Italy, Greece). Genetically slightly heterogeneous, maximum variation 0.6%. Nearest species: C. pullata (1.8%), C. crenulata (2.2%). Remarks: DNA barcode analyses support the status of C. italohelveticus as a separate species. 74

14. Charissa pullata (Denis & Schiffermüller, 1775) Geometra pullata Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 108 (Austria: Vienna district). Syntype(s) (lost). Gnophos impectinata Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 309 (France méridionale: Basse-Alpes). Syntypes ♂♀ (ZFMK, not examined). Herewith downgraded from subspecific rank of C. pullata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos albarinata Millière, 1859: Iconogr. Descr. Chenilles Lépid. inédits 1 (2): 398, pl. 1, figs 12–15 (France: Ain, Grande-Chartreuse, near Fourvoirie). Syntype(s). Herewith downgraded from subspecific rank of C. pullata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos pullata [“v. (ab.)”] confertata Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 167 ([Europe]). Syntype(s) (MNHU, not examined). Herewith downgraded from subspecific rank of C. pullata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos pullata [“var.”] nubilata Fuchs, 1872: Stettin. ent. Ztg. 33 (10–12): 429 (Germany: Geroldstein, valleys and side valleys of river Wisper). Syntype(s) (NSMW, not examined). Herewith downgraded from subspecific rank of C. pullata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos pullata kovacsi Vojnits, 1967: Annales historico-naturales Musei Nationales Hungarici 59: 380 (Hungary: Mount Börzsöny, Pogányvár, 450–550 m). Holotype ♂ (TMB, not examined). Herewith downgraded from subspecific rank of C. pullata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.).

External characters and abdomen: Wingspan 29–39 mm. Wings broad, ground colour of upperside greyish white, ash-grey, light beige grey, lead-grey or blackish grey, sometimes with suffusion of bright or dark scales on all wings. Transverse lines often broken into dots, thickened and darker at costa. Small black terminal dots present. Discal spots variable, usually unfilled. Fringes hardly chequered. Hindwing termen undulating. Underside of wings with conspicuous, proximally indented, white postmedial line on fore- and hindwing. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: Ground colour of the adults is very variable depending on the colour of the local substrate. It ranges from bright white (on limestone) to blackish grey (on primary rock). Even populations at small geographic distance can differ greatly in colour (e.g. Petry 1923; Bergmann 1955). Male genitalia: Uncus pointed, but stocky and slightly concave. Gnathos fairly thick and stout from base to tip, bent at right angles in the middle. Juxta divided almost down to the base, u-shaped. Costa of valva well sclerotized, distally clearly bent. Aedeagus relatively stout, with strong, narrow cornutus. Female genitalia: Antrum well sclerotized, margins clearly triangular. Distribution and abundance: European, submontane. In the north from Belgium through central Germany and southern Poland, eastwards to Ukraine, in the south-east to Romania and Bulgaria, and southwards to Greece. In the west, reaching French Pyrenees, but absent from Iberian peninsula (Leraut 2009; Redondo et al. 2009; Flamigni et al. 2016).

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Phenology: Univoltine. From June to early August. Main flight period in submontane regions in mid-June, in the Alps usually later, mid-July. Overwintering as larva, last instars found in May (Bartsch 2003; King & González-Estébanez 2015; S. Erlacher own observation). Adults active at dusk and night, attracted to light, resting in shady places during daytime. Biology: Larva polyphagous. Found on Gypsophila repens, Sedum album, Sedum rupestre, Teucrium montanum, Thymus, Origanum, Rubus, Cytisus scoparius, Lembotropis nigricans, Vaccinium, Valeriana (Bergmann 1955), on Asplenium ruta-muraria, Coronilla coronata, Genista sagittalis (Bartsch 2013), and under cushions of Sanguisorba minor and Helianthemum nummularium (M. Leipnitz pers. comm.). Recorded on Artemisia, ­Galium, Geum, Gnaphalium, Lactuca, Plantago (Flamigni et al. 2016), and on Sedum maximum (S. ­Erlacher own observation). Successfully reared on Lotus corniculatus, Rumex and Taraxacum (Flamigni et al. 2016), on Hippocrepis comosa, Saxifraga aizoides (Ziegler 2019), as well as under mats of Rumex scutatus (King & González-Estébanez 2015). Reared on Loni­cera tatarica, during hibernation on Sanguisorba minor (M. Leipnitz pers. comm.). Cater­pillars active at night, resting on ground or in rock crevices during daytime. Immediately before pupation also active in daytime, apparently when searching for pupation places (S. Erlacher own observation). Pupation on the ground among moss and plant parts in a fine, light spinning consisting of just a few threads (rearing observation; M. Leipnitz pers. comm.). Habitat: Xerothermophilous, montane. Mountains and hills with xerothermic rock and scree habitats, in bushy slopes, in partially shaded, rocky hollow paths. Sometimes also in smallscale biotopes (Bartsch 2013). From 200 m up to 1500 m, in the Aosta Valley up to 2100 m (Flamigni et al. 2016). Similar species: C. pullata differs from C. crenulata and C. italohelveticus in some phenotypic characters: The transverse lines on all wings are usually less visible and more broken into single dots in C. pullata. On the underside of the wings the white transverse line is usually 76

narrower and less serrated than in the other two species. The dark terminal area usually does not contain light spots as in C. crenulata and C. italohelveticus, giving a much less contrasting appearance to the underside. In certain cases, dissection of the genitalia is required for secure identification: In male genitalia of C. pullata, the juxta is u-shaped and the distal end of the costa is clearly bent. In female genitalia, the antrum is triangular in C. pullata (quadrangular in C. crenulata; with slightly rounded margins and height=width in C. italohelveticus). Genetic data: BIN: BOLD:AAC7656 (n=24 from Germany, Austria, Italy, Croatia, Romania, Macedonia). Genetically homogeneous. Nearest species: C. crenulata (1.7%), C. italohelveticus (1.8%). Remarks: As some other Charissa species, C. pullata tends to develop substrate-adapted individual forms that recur at distant locations. This applies to the following forms, which are therefore synonymized here with the nominotypical subspecies: Gnophos albarinata Millière, 1859 (syn. n.), Gnophos pullata confertata Staudinger, 1871 (syn. n.), Gnophos impectinata Guenée, 1858 (syn. n.), Gnophos pullata kovacsi Vojnits, 1967 (syn. n.), and Gnophos pullata nubilata Fuchs, 1872 (syn. n.).

Subgenus Dysgnophos Wehrli, 1951 Dysgnophos Wehrli, 1951: Lambillionea (51): 23. Type species: Gnophos difficilis Alphéraky, 1883.

Diversity and distribution: The subgenus Dysgnophos contains five species in the Palaearctic region, one species (C. turfosaria) also occurs in the Nearctic. They are mainly distributed in the mountains of Central Asia. Male genitalia: Uncus hook-shaped. Gnathos well developed. Valva slim to moderately wide, usually with thorn on costa. Juxta deeply split, with long, slender, and distally pointed juxta arms. Aedeagus at tip with fin-like extensions and a semi-annular cornutus. Female genitalia: Very similar to those of subgenus Pterygnophos Wehrli, 1951 (see E ­ rlacher et al. 2017). Corpus bursae pipe-shaped, with strongly sclerotized, spinose, longitudinal folding, laterally with appendix bursae (dome-like structure with ring-like folds, and a set of stout and large teeth inside). Antrum well sclerotized. Remarks: Viidalepp (1996) considered the taxon Dysgnophos as a separate genus, while B ­ eljaev (2016) treated it as a subgenus of Charissa. Based on morphological correspondence of male and female genitalia with the autapomorphic character states of genus ­Charissa (Erlacher et al. 2017), the taxon Dysgnophos is treated here as a subgenus of the latter. DNA barcode analyses strengthen this classification.

15. Charissa difficilis (Alphéraky, 1883) Gnophos difficilis Alphéraky, 1883: Horae Soc. Ent. Ross. 17 (3/4): 178, pl. 9, fig. 86 (Kyrgyzstan or Kazakhstan: foothills of Tian Shan, 3500 ft. [1067 m]). Syntypes ♂♀ (ZRAS, examined). Gnophos difficilis var. uniformis Staudinger, 1892: Dt. ent. Z. Iris 5: 188 (central Asia: Lepta). Holotype ♂ (MNHU, examined). Validated at subspecies rank by Scoble (1999), herewith downgraded to synonymy with C. difficilis (syn. n.), based on the absence of constant and significant differential features. Gnophos difficilis var. korlata Fuchs, 1903: Societas ent. 18 (6): 43 (north-western China, Xinjiang: Korla). Syntypes 2♂ (MNHU, examined). Validated at subspecies rank by Scoble (1999), at species rank by Viidalepp (1996), here again treated as subspecies of C. difficilis.

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Gnophos difficilis urmensis Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (suppl.): 593, fig 48h (Russia, Dagestan: Urma, Levashi, Kurush). Syntypes 3♂ (ZRAS and ZFMK, examined) (stat. rev.). Gnophos difficilis teberdensis Müller, 1978: Entomol. Berichte 1978 (1): 33, pl. 1, figs 11, 12; pl. 2, figs 11, 12; pl. 5, fig. 1; pl. 6, fig. 4 (Russia, north-western Caucasus: Teberda). Holotype ♂ (coll. B. Müller, examined).

External characters and abdomen: Wingspan 28–36 mm. Wings comparatively narrow, costa of forewing slightly convex. Ground colour of wings varying from reddish or yellowish brown, grey to yellowish and white. Transverse lines usually distinct, medial area usually prominent. Discal spots distinct, filled or unfilled. Fringes only slightly spotted. Underside of wings with uniform basal and medial area, terminal area darker, medially with brighter postmedial line. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: See subspecies. Very variable. Specimens from the European part of Russia (southern Ural) are brighter than all other known subspecies, wings are bright white to pale yellow and the wing pattern is only slightly visible, underside of the wings uniform, with contrasting coloured apical area of forewings and terminal area of hindwings. Possibly a well-defined, unnamed subspecies. Subsp. teberdensis (Müller, 1978) (north-western Caucasus, Teberda): Ground colour ochreous, rich in contrast. Wing pattern blackish and very clear. Transverse lines with blackish jags on veins. Discal spots distinct, blackish grey, with slight, bright filling. Fringes concolorous with ground colour, strongly chequered ochreous and grey. Underside of wings yellowish, with sharp pattern, more contrasted than in the other subspecies. Unknown from Europe. Subsp. urmensis (Wehrli), 1953 stat. rev. (eastern Caucasus, Dagestan): Smaller than nomino­typical subspecies (28–31 mm), ground colour lighter, whitish grey. Transverse lines on wing upperside fine but distinct, with blackish grey dots on veins. Discal spots blackish grey, distinct, only slightly filled with white scales. Wing underside homogeneous. Unknown from Europe. Male genitalia: Uncus wide, bicuspid, with a short blunt tip in the middle and two rounded tips laterally (“mouse head-like” sensu Wehrli 1922c: 11). Gnathos short and blunt. Valva narrow, costa more sclerotized, at distal end of sclerotized part with a simple thorn. Ventral margin of valva slightly s-shaped. Juxta deeply divided, juxta arms thin, evenly slender. Aedeagus tubular with a cornutus shaped as plate with sclerotized swelling. Female genitalia: Corpus bursae pipe-shaped, with strongly sclerotized, spinose, longitudinal folding, laterally with appendix bursae (dome-like structure with ring-like folds, and a set of stout and large teeth inside). Antrum well sclerotized, relatively long, with funnel-shaped ostium. Distribution and abundance: Central Asian and Caucasian, montane. In Europe only known from southern Ural (Schibendy valley, located in the southern edge of Orenburg district, coll. K. Nupponen). – Outside Europe four subspecies are described: the nominotypical subspecies from Tian Shan mountains, south-eastern and eastern Kazakhstan, Kyrgyzstan, Mongolia (Altai and Govy Altai, coll. P. Skou), subsp. urmensis Wehrli, 1953 from eastern Caucasus (North Ossetia in Russia, Dagestan, Armenia) (Wehrli 1953; Rjabov & ­Vardikjan 1964; Müller 1978), subsp. teberdensis Müller, 1978 from north-western Caucasus (­Teberda), and subsp. korlata Fuchs, 1903 from China (Korla). 78

Phenology: Univoltine. From early June to late August (Rjabov & Vardikjan 1964; Müller 1978), in Central Asia slightly earlier, from mid-May to mid-July. In southern Ural in early June (K. Nupponen pers. comm.). Adults nocturnal, attracted to light. Biology: Unknown. Successfully reared on withered Taraxacum (Müller 1978), larvae growing very slowly and overwintering with a length of 7 mm. Habitat: Xeromontane. Limestone steppe in southern Ural (K. Nupponen pers. comm.). In the Caucasus region in mountain steppes and subalpine mountain meadows, steppe grass and open pine forest (Müller 1978). From 1000 m in south-eastern Kazakhstan (coll. M. Sommerer) up to 3500 m in Kyrgyzstan (coll. ZSM). Similar species (see Text-figs 29–30): C. difficilis differs from C. turfosaria in some pheno­ typic characters: On the underside of the wings the terminal area and the postmedial line are more clearly visible in C. difficilis, giving a more contrasting appearance to the underside. However, in certain cases, dissection of the genitalia is required for secure identification. Genetic data: BIN: BOLD:ABA3449 (n=27 from Russia, Urals, Kazakhstan, Kyrgyzstan, Mongolia). Genetically homogeneous. Nearest species: Charissa orphninaria (Hampson, 1902), comb. n. (transferred from Gnophos, see Scoble 1999) from Pakistan (3.7%); C. turfosaria (4.6%). Remarks: Viidalepp (1996) considered urmensis as bona species. However, comparison of the type specimens and their genitalia clearly shows the conspecifity with C. difficilis: Charissa (Dysgnophos) difficilis urmensis (Wehrli, 1953) (stat. rev.). Gnophos finitimaria Fuchs, 1899 (Jahrb. Nass. Ver. Naturk. 52: 149; Russia: Krasnojarsk; Lectotype ♂, MNHU, ­examined) was listed as synonym of ‘Gnophos’ difficilis in Scoble (1999), but recognized as synonym of Gnophos ochrofasciata Staudinger, 1895 in Erlacher et al. (2017).

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Text-figs 29–30. Diagnostic characters (indicated) of European Charissa species, subgenus Dysgnophos. Underside of wings. Occasionally, external characters do not allow reliable identification, in which case the genitalia need to be examined. Text-fig. 29. C. difficilis (Alphéraky, 1883), male. Kazakhstan: prov. Almaty, Bokaydin-tau, Zailisky Alatau, 13.–15.vi.1994 (coll. Sommerer). Text-fig. 30. C. turfosaria (Wehrli, 1922), male. Russia: Republic Altai, Aktasch, 16.vi.2012 (coll. Schacht).

16. Charissa turfosaria (Wehrli, 1922) Gnophos turfosaria Wehrli, 1922: Dt. ent. Z. Iris 36: 13, pl. 1, fig. 7; pl. 2, figs 29, 45 (Russia: Sajan mountains). Syntypes 2♂ (ZFMK and MNHU, examined). Gnophos glaciata Wehrli, 1922: Dt. ent. Z. Iris 36: 14, pl. 1, fig. 8; pl. 2, figs 30, 46 (Russia: Sajan mountains, Tannuola). Syntypes 2♂ (ZFMK and MNHU, examined). Synonym of C. turfosaria (Scoble 1999). Gnophos exilis Wehrli, 1922: Dt. ent. Z. Iris 36: 15, pl. 1, fig. 10; pl. 2, figs 38, 54 (Russia: Sajan mountains, Munko Sardyk). Lectotype ♀, hereby designated (ZFMK, examined, label data: Sajan, genit. prep. Wehrli–606). In Scoble (1999) synonymized with Gnophos subsplendidaria Wehrli, 1922, here retained synonym of C. turfosaria (syn. n.), based on examination of the lectotype and its genitalia. Gnophos benepunctaria Wehrli, 1922: Dt. ent. Z. Iris 36: 16, pl. 1, fig. 24; pl. 2, figs 37, 53 (Russia: Sajan mountains). Syntype ♀ (ZFMK, examined). Synonym of C. turfosaria (Scoble 1999). Gnophos macguffini Smiles, 1979: J. Nat. Hist. 13: 106, figs 3, 4, 14, 18, 20 (Canada, North West Territories: Victoria Is.). Holotype ♂ (CNC, examined). Validated at species rank in Scoble (1999), herewith downgraded to synonymy of C. turfosaria, based on clinal character transitions, and DNA barcode-sharing (syn. n.). Unavailable names (incorrect subsequent spelling): mcguffini: Viidalepp (1996).

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External characters and abdomen: Wingspan 26–37 mm. Wings usually broad, sometimes narrow, costa of forewing straight or slightly concave. Ground colour of wings varying from yellowish brown, grey, bluish grey to reddish grey, and from roughly scaled to shiny. Transverse lines usually clear, medial area not contrasted. Discal spots distinct, filled or unfilled. Fringes not dotted. Underside of wings with uniform basal and medial areas, terminal area concolorous or slightly lighter. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: Extremely variable, even within comparatively small areas of occurrence. Specimens from central Ural mountains rich in contrast, coarsely scaled; reminiscent of what Wehrli (1922c) described as “benepunctaria” from Sajan mountains (synonym of C. turfosaria). Male genitalia: Highly variable. Uncus bent and pointed, beak-shaped. Gnathos stout, more or less obtuse, from slightly pointed to broad, sometimes even slightly heart-shaped. Valva usually slender, costa more strongly sclerotized towards apex, last third with one or several weak or stout thorns. Ventral margin of valva usually parallel to costa. Juxta deeply divided into two arms, juxta arms of same length, but variable in thickness and shape, slightly bent or parallel-sided. Aedeagus tubular with a cornutus shaped as plate with sclerotized swelling. Female genitalia: Corpus bursae pipe-shaped, with strongly sclerotized, spinose, longitudinal folding, laterally with appendix bursae (dome-like structure with ring-like folds, and a set of stout and large teeth inside). Antrum well sclerotized, relatively short, narrowing anteriorly. Ostium with sub-triangular postvaginal plate. Distribution and abundance: Holarctic. In Europe, C. turfosaria only occurs in Polar Urals (Viidalepp 1996, as “mcguffini” [sic!]). – Outside Europe from eastern slope of central Ural mountains (Denezhkin Stone mountain, Sverdlovsk district, coll. K. Nupponen; Beljaev 2016, as “C. macguffini”) across Altai mountains in Russia (colls J. Gelbrecht, U. ­Ratzel,

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B.  Schacht, ZSM), Kazakhstan (coll. P. Skou), Govi-Altai mountains in Mongolia (colls P. Skou, M. Sommerer), Buryatia (coll. R. Müller) and Tynda in southern Russia (coll. P. Skou) to Magadan area in Russia Far East (coll. P. Skou). Also recorded from Alaska and Canada (McGuffin 1977 as “Gnophos sp.”; Smiles 1979; Pohl et al. 2018 as “Gnophos macguffini”). Phenology: Univoltine. From mid-May to mid-July. Rearing from eggs of a female collected in mid-June resulted in adults in October of the same year (J. Gelbrecht pers. comm.). Adults nocturnal, attracted to light. Resting on rocks and easily disturbed during daytime (K. Nupponen pers. comm.). Biology: Larval foodplant unknown. Habitat: Inhabiting mountain tundra (K. Nupponen pers. comm.), Siberian taiga, light, herbrich coniferous forest with larch, spruce, pine, and single rocks (J. Gelbrecht pers. comm.). From 200 m up to 2300 m in Russia Altai (colls B. Schacht; ZSM) and from 700 m in Magadan area in Russia Far East (coll. P. Skou) up to 2500 m in Govi Altai mountains (colls P. Skou; M. Sommerer). Similar species: C. difficilis (see no. 15 and Text-figs 29–30). Genetic data: BINs: BOLD:AAD4767 (n=42 from European Russia, central and eastern Russia, Mongolia and north-western Canada). One BIN-sharing DNA barcode from Turkmenistan (outside the so far known distribution) requiring further study. Genetically slightly heterogeneous, but maximum variation only 0.8%, with two slightly more diverging (but BIN-sharing) specimens from Mongolia. Three additional BINs (BOLD:AAO7048; BOLD:ACY4875; BOLD:ABA9746; n=9 from Russia and Mongolia) at distances of 2.9–3.4%, probably not conspecific, requiring further study. Nearest species: Charissa orphninaria (Hampson, 1902), comb. n. (transferred from Gnophos, see Scoble 1999) from Pakistan (4.1%); C. difficilis (4.6%). Remarks: Even Wehrli (1922c) was deceived by the extreme variability in the external appearance and the shape of the genitalia resulting in the description of not less than five species from the Sajan mountains. Later, Smiles (1979) added another description based on specimens from Canada. However, detailed comparison of all type specimens, the study of numerous genitalia slides (n=60) as well as DNA barcode analyses clearly showed the conspecifity with C. turfosaria (= Gnophos macguffini Smiles, 1979 syn. n.).

Subgenus Cnestrognophos Wehrli, 1951 Cnestrognophos Wehrli, 1951: Lambillionea (51): 26. Type species: Gnophos praeacutaria Wehrli, 1922.

Diversity and distribution: Subgenus Cnestrognophos with about 20 species in the Palae­ arctic region, most of them occurring in Central Asia. Only three species known from Europe. Male genitalia: Juxta deeply divided down to the base. Costa well sclerotized and curved, medially with short process bearing one or more thorns. Centre of valva with a sclerotized spinose crest, in some species modified to a sclerotized clasp. Female genitalia: Corpus bursae with a folded posterior sclerotization and a hyaline anterior part with a half-rounded, brain-like, sclerotized protrusion.

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17. Charissa mutilata (Staudinger, 1879) Gnophos mutilata Staudinger, 1879: Horae Societatis Entomologicae Rossicae 14: 457 (Turkey: Amasia). Holotype ♂ (MNHU, examined).

External characters and abdomen: Wingspan 22–30 mm. Ground colour of wings usually yellowish grey, but ranging from whitish to dark ochreous. Fringes usually lighter. Transverse lines on all wings orange, discal spots orange-brown, filled or unfilled. Hindwing termen rounded. Underside of wings less marked, lighter, ground colour uniform. Frons, collar, thorax and abdomen concolorous with wings. Male antennae serrated, with long cilia, female antennae filiform. Variation: Considerably varying in size and coloration. Specimens of the first generation are much larger than those of the second. The ground colour of the wings depends on the substrate and varies from almost white to dark ochreous with bluish grey parts. Male genitalia: Uncus beak-shaped, abruptly narrowing towards distal end, blunt. Gnathos slender, distally rounded. Juxta deeply divided down to the base, juxta arms slender and pointed, skewer-shaped. Costa of valva well sclerotized and curved, distal third covered with long setae, medially with short and stout thumb-shaped process bearing a single, stout thorn; ventral margin of valva well sclerotized, with short and stout thorn-shaped process. Harpe developed as a sclerotized spinose crest with pointed, protruding proximal and distal extensions. Saccus short and small, bent backwards. Aedeagus tubular, without cornutus. Female genitalia: Antrum inconspicuous, small. Corpus bursae with posterior, longitudinally folded sclerotization of 0.83–1.16 mm length and a hyaline anterior part with half-rounded brain-like appendix bursae. Distribution and abundance: (South-east) European – West-Asiatic. So far, in Europe, only known from Bulgaria, here erroneously identified as C. pentheri (Beshkov 2017). Records

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from Greece (Rebel 1902; Hacker & Lukasch 1984) most probably referring to C. peloponnesiaria. – In Europe local and rare. – Outside Europe widespread throughout Turkey and Russian Armenia (Wehrli 1953). Phenology: Probably bivoltine, from Europe only known in the first generation. Specimens from Bulgaria collected from late April to early May (Beshkov 2017). In Turkey bivoltine, from mid-April to mid-June and again from mid-July to mid-September. Adults active at night, attracted to light. Biology: Unknown from nature. Under rearing conditions polyphagous on low plants (J. Gelbrecht pers. comm.). Habitat: Xerothermophilous, montane. Open, rocky slopes on limestone. In Bulgaria from 700 m up to 975 m (Beshkov 2017). In Turkey from 700 m up to 2400 m, in Russian Armenia from 2300 m up to 3300 m (Wehrli 1953). Similar species: Adults of C. mutilata closely resemble those of C. peloponnesiaria and C. pentheri, but are often darker and more strongly contrasted. In male genitalia, both protrusions of the harpe longer and more cuspidate than in C. peloponnesiaria. For full diagnosis see C. peloponnesiaria. Genetic data: BINs: BOLD:AAL9633 (n=4 from Bulgaria), AAD5323 (n=8 from Turkey). European and Turkish BINs diverging by 3.0%. Nearest species: C. peloponnesiaria (3.9%).

18. Charissa peloponnesiaria Erlacher & Erlacher, 2017 Charissa peloponnesiaria Erlacher & Erlacher, 2017: Zootaxa 4341 (1): 89, Pl. 1, figs 2a, 2b (Greece, Peloponnese: Parnon Mountains, NE Kosmas, 850 m). Holotype ♂ (coll. Erlacher).

External characters and abdomen: Wingspan ♂ 21–30 mm, ♀ 27 mm. Ground colour of wings whitish or ochreous, scattered with darker scales. Wing pattern with transverse lines and discal spot sometimes distinct and edged orange. Underside pale and dull. Frons, collar, thorax and abdomen concolorous with wings. Variation: The ground colour of the adults often varies from whitish to ochreous. Male genitalia: Uncus beak-shaped, abruptly narrowing towards distal end, blunt. Gnathos slender, distally rounded. Juxta deeply divided down to the base, juxta arms slender and pointed, skewer-shaped. Costa of valva well sclerotized and curved, distal third covered with long setae, medially with short and stout thumb-like process bearing a single strong thorn. Ventral margin of valva well sclerotized, with a short and stout thorn-like process. Harpe developed as a sclerotized, spinose crest with rounded proximal and cusp-like distal protrusion. Saccus short and small, bent backwards. Aedeagus tubular, without cornuti. Female genitalia: Antrum inconspicuous, small. Corpus bursae with posterior, ­longitudinally folded sclerotization. Anterior part hyaline with a half-rounded brain-shaped appendix bursae. Distribution and abundance: Endemic to the southern Balkan peninusla. Only known from the north and south-east of Peloponnese peninsula and from the island of Kephalonia in the Ionian Islands (Erlacher & Erlacher 2017b). Phenology: Bivoltine. Scarce data from late April to mid-May (leg. S. Beshkov, leg. J. ­Viehmann) and from late August to late September. Adults nocturnal (Erlacher & Erlacher 2017b). 84

Biology: Unknown. Habitat: Xerothermophilous, montane. Found in open, dry hills and calcareous rock habitats with sparse vegetation. From 500 m up to 2200 m (Erlacher & Erlacher 2017b; J. Viehmann pers. comm.). Similar species: Adults of C. peloponnesiaria closely resemble those of C. mutilata, but are slightly paler and less deeply coloured. Adults of C. pentheri are larger (wingspan: 31 mm) and their wings always lack the orange shading occasionally occurring in the two other species. In the genitalia, the species is particularly recognizable by the shape of the harpe: unlike in the other European species of this subgenus, the harpe of C. peloponnesiaria is developed as a spinose crest with a rounded proximal and a short cuspidate distal protrusion, while in C. mutilata both protrusions of the harpe are longer and cuspidate (pl. 6, figs 17c, 18c). In C. pentheri the harpe is more evenly shaped without any protrusions, but this species is prima facie also distinguishable from C. peloponnesiaria by the thin costal thorns of the valva. Similarly, in female genitalia, the discrimination between C. pentheri and C. peloponnesiaria is more obvious than between C. mutilata and C. peloponnesiaria: C. pentheri has clasp-like lateral sclerites in contrast to C. mutilata and C. peloponnesiaria where the lateral sclerites are flatter, the antrum is larger and rounded and furthermore the appendix bursae is situated at the left side of the corpus bursae (ventral view). The female genitalia of C. mutilata and C. peloponnesiaria are much more similar: in both species the antrum is very small and the appendix bursae is situated at the right side of the corpus bursae. However, in C. peloponnesiaria the folded longitudinal sclerotization of the corpus bursae is shorter (length of the folded longitudinal sclerotization in C. peloponnesiaria: 0.77 mm; length of the folded longitudinal sclerotization in C. mutilata: 0.83–1.16 mm) (­Erlacher & Erlacher 2017b).

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Genetic data: BINs: BOLD:AAD5324 (n=5 from Greece). Genetically heterogeneous due to a diverging specimen from Kefalonia island, intraspecific variation 2.5%. Nearest species: C. mutilata (3.9%).

19. Charissa pentheri (Rebel, 1901) Gnophos pentheri Rebel, 1901: Verh. zool.-bot. Ges. Wien 51: 801 (Bosnia-Herzegovina: Mount Plasa, 1300 m). Syntypes 1♂1♀ (NHMW, not examined). Gnophos pentheri petrina Thurner, 1936: Zeits. Österr. Ent.-Ver. 21: 14 (Macedonia: Petrina near Ochrid, 1600 m). Syntypes (NHMW, not examined, one specimen labelled as ‘allotype’ in ZSM, examined and DNA barcoded). Validated at subspecific rank (Scoble 1999), herewith downgraded to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.).

External characters and abdomen: Wingspan 29–32 mm. Ground colour of wings usually uniform whitish grey, without other coloration. Wing pattern with transverse lines and discal spot somewhat faded. Fringes slightly yellowish. Underside of wings pale and dull. Frons, collar, thorax and abdomen concolorous with wings. Male antennae serrated. Variation: Very little. Wings are more or less scattered with dark or yellowish scales. Male genitalia: Uncus beak-shaped, distally rounded. Gnathos slender. Juxta deeply divided down to the base, juxta arms slender and pointed, skewer-shaped. Costa of valva well sclerotized and curved, distal third covered with long setae, medially with short and stout thumb-like process bearing a single more or less thin thorn, which can be absent. Ventral margin of valva well sclerotized, without process. Harpe developed as an evenly shaped, sclerotized, spinose field without protrusions. Saccus short and small, bent backwards. ­Aedeagus tubular, without cornuti. Female genitalia: Antrum with clasp-like sclerotized lateral sclerites. Corpus bursae with a posterior, longitudinally folded sclerotization and a hyaline anterior part with a half-rounded brain-shaped appendix bursae situated at the left side of the corpus bursae (ventral view). Distribution and abundance: (South-east) European. Endemic to the Balkan peninsula. Known only from Bosnia-Herzegovina, Montenegro, and Macedonia. – One of the rarest European species of Charissa. – Unknown from outside Europe. Phenology: Very few records from late May to early July (Rebel 1901; Thurner 1936). Adults nocturnal (Daniel et al. 1951). Biology: Unknown. Habitat: Xerothermophilous, montane. Found in calcareous rock habitats. From 1300 m up to 1600 m (Rebel 1901; Thurner 1936). Similar species: Adults of C. pentheri closely resemble those of C. mutilata and C. peloponnesiaria, but are larger (wingspan: 31 mm) and their wings always lack the orange shading occasionally occurring in the two other species. In male genitalia of C. pentheri, the harpe is more evenly shaped, without any protrusions. In female genitalia, it has clasp-shaped lateral sclerites in contrast to C. mutilata and C. peloponnesiaria where the lateral sclerites are flatter, the antrum is larger and rounded and furthermore the appendix bursae is situated at the left side of the corpus bursae (Erlacher & Erlacher 2017b). It could also be confused

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with bright specimens of C. ambiguata, however, the male antennae of the latter are filiform while those of C. pentheri are distinctly serrated. Genetic data: So far, only sequenced to short sequence (133bp) for the allotype of synonym petrina. Nearest species: C. mutilata (3.7%), genetic distances revealed from such short sequences requiring confirmation. Remarks: Species of the genus Charissa tend to develop substrate-adapted individual forms. However, there are only colour-deviant individuals within more or less uniform populations. This also applies to specimens of C. pentheri from Ochrid which were described as subspecies petrina by Thurner (1936). It is therefore synonymized here with the nominotypical subspecies (syn. n.).

Subgenus Euchrognophos Wehrli, 1951 Euchrognophos Wehrli, 1951: Lambillionea (51): 25. Type species: Gnophos variegata D ­ uponchel, 1830.

Diversity and distribution: The subgenus Euchrognophos contains nine species in the Palaearctic and Oriental regions. These are small, often quite colourful moths. The male antennae are often diagnostic and can be either dentate, serrated or bipectinate (see Text-figs 31–44). Female antennae always filiform. Underside of wings usually well contrasted with conspicuous black and white pattern. Females much larger than males, wings more rounded, underside of wings much darker, with more pronounced dark spots in the terminal area. Variation: Extremely variable, even within the same populations, but also between different populations depending on the substrate. 87

Male genitalia: Uncus triangular, blunt at tip. Gnathos well developed, of about the same length as uncus. Valva bipartite, divided into a dorsal, more hyaline and a ventral, more sclerotized part; the latter distally knife-shaped and usually with a smaller dorsal extension of varying thickness and length; costa usually with one, sometimes with two thorns. Juxta split, with slender juxta arms. Aedeagus with two stout cornuti. Female genitalia: Corpus bursae without protuberances, strongly sclerotized, with distinctly spined longitudinal folding; at the anterior end with two hyaline diverticula. Ductus bursae absent. Antrum well sclerotized, sometimes with noticeable folding; lamella postvaginalis cylindrical, sometimes extending antrum. Remarks: Viidalepp (1996), Leraut (2009) and Redondo et al. (2009; 2010) validated the taxon Euchrognophos at genus rank, in contrast to Scoble (1999), Huemer et al. (2009), Corley (2015), Flamigni et al. (2016) and many others. Based on morphological similarity of male and female genitalia with the autapomorphic character states of the genus Charissa (Erlacher et al. 2017), we treat the taxon Euchrognophos as a subgenus of the latter here. DNA barcode analyses strengthen this classification.

20. Charissa assoi (Redondo & Gastón, 1996) Gnophos (Euchrognophos) assoi Redondo & Gastón, 1996: Zapateri revista aragonesa de entomología 6: 156, figs 1, 9 (Spain, Zaragoza: Torralba de los Frailes, 1050 m). Holotype ♂ (private collection of Victor Redondo, Zaragoza, not examined).

External characters and abdomen: Wingspan 17–25 mm, subsequent generation(s) smaller than the first. Wings with ground colour varying from ochreous to grey; black scales arranged in alternating roof tile-like rows from costa to inner margin of fore- and hindwings. Transverse lines usually clear, ochreous to orange. Discal spots sometimes indistinct, filled or unfilled. Fringes brighter than ground colour, not chequered. Male antennae distinctly dentate (Text-figs 31, 38). Variation: Somewhat variable individually, in colour and size. Male genitalia: Uncus blunt at tip, forming an equilateral triangle. Gnathos stout, narrower towards tip, blunt at tip. Ventral part of bipartite valva with clearly sclerotized edge, almost rectangular; the knife-shaped distal process clearly exceeding edge of valva; dorsal extension sometimes present as a small spike; costa with one thorn, which can be absent. Juxta split slightly more than halfway, juxta arms distally slightly curved. Aedeagus proximally with a shorter and distally with a longer cornutus (average length ratio of short versus long cornutus 0.5:1). Female genitalia: Corpus bursae slender, boomerang-shaped; at the anterior end, before the transition to the hyaline diverticulum, not or hardly narrower; twice as long as the antrum. Antrum rectangular, more strongly sclerotized slightly arched ledges at transition towards corpus bursae. Lamella postvaginalis funnel-shaped, exceeding antrum. Distribution and abundance: West Mediterranean, Iberian endemic. Previously known only from montane habitats in central Spain (Redondo et al. 2009: Segovia, Guadalajara, Teruel, Zaragoza, Tarragona), later found also in coastal and arid places to the south, i.e. Murcia (D. Stadie pers. comm.), Almeria (Garre et al. 2013) and Malaga province (P. Hale pers. comm.). – Local and in very limited numbers (P. Skou own observation).

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Phenology: Bivoltine. From mid-April to mid-June and again from mid-September to early November (Redondo et al. 2009; Garre et al. 2013; P. Skou own observation). In mountainous areas obviously only one generation from early July to late August (P. Skou own observation). Biology: Larva unknown. Habitat: Probably xerothermophilous. On rocky landscapes with many trees and bushes (P. Skou own observation), on rocks and in gorges (Redondo et al. 2009; Garre et al. 2013). From 450 m up to 1100 m in central Spain (Redondo et al. 2009; Garre et al. 2013), up to 2000 m in the Sierra Nevada (coll. ZSM) and down to sea-level in Almeria. Similar species: C. assoi is externally similar to allopatric C. variegata. The latter, however, with only slightly dentate male antennae (more distinctly dentate in C. assoi). It is also possible to confuse C. assoi with C. mucidaria which shares the same habitat but has distinctly bipectinate male antennae. Finally, C. assoi can easily be distinguished from all other species of the subgenus Euchrognophos by the examination of male and female genitalia. Genetic data: BIN: BOLD:AAE8722 (n=9 from Spain). Genetically homogeneous. Nearest species: Iranian C. dubitaria (5.0%), European C. variegata (5.9%). Remarks: The female genitalia, which were used for the original description, are broken in the middle of the corpus bursae (see Redondo & Gastón 1996). Due to that artifact they appear much stouter than they actually are.

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Text-figs 31–44. Male antennae of European Charissa species, subgenus Euchrognophos. Upper row lateral view, lower row ventral view. Text-fig. 31–32. C. assoi (Redondo & Gastón, 1996), distinctly dentate. Spain: Sierra Nevada, Veleta, 2000 m, 8.xii.1971, leg. J. Klimesch, (ZSM). Text-fig. 33–34. C. corsica (Oberthür, 1913), serrated. Italy/Sardinia: Olbia-Tempio, San Pantaleo, 190 m, ♂: 4.iv.2007, e.o., leg. B. Müller (coll. Erlacher). Text-fig. 35–36. C. corsica (Oberthür, 1913), bipectinate. France: Corsica, v.1923 (ZSM). Text-fig. 37–38. C. staudingeri (Wnukowsky, 1929), shortly bipectinate. Greece: Naoussa, 2005, e.o. (coll. Erlacher). Text-fig. 39–40. C. subtaurica (Wehrli, 1934), dentate. Turkey: Adana, Kozan, 18.iii.2001, leg. F. Doğanlar (coll. Erlacher). Text-fig. 41. C. variegata (Duponchel, 1830), slightly dentate. Italy/Malta: St. Julians, 11.–12.v.2000, lux, leg. F. Stöckl (coll. Pöll). Text-fig. 42. C. variegata (Duponchel, 1830), slightly dentate. Greece: Epirus, n/w Loannina/Zagoria, Monodendri, 1200 m, 24.v.2002, lux (coll. Friedrich). Text-fig. 43–44. C. mucidaria (Hübner, 1799), distinctly bipectinate. Italy: Liguria, Noli (Savona), 5.–30.ix.1951, leg. J. Klimesch (ZSM).

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21. Charissa corsica (Oberthür, 1913) Gnophos variegata [var.] corsica Oberthür, 1913: Études Lépid. comp. 7: 301, pl. 179, fig. 1735 (Corsica). Syntypes 1♂2♀ (ZFMK, examined).

External characters and abdomen: Wingspan 20–28 mm, second generation smaller than the first. Females often twice the size of males. Wings with ground colour varying from pale beige to bluish grey; pattern uniform to well contrasting. Transverse lines often interrupted, dark grey to orange. Discal spots usually distinct and filled. Underside of wings strongly contrasting, terminal area darkened, each wing with whitish spot at centre, forewing with another whitish spot on apex. Fringes brighter than ground colour, usually chequered. Male antennae serrated, exceptionally shortly bipectinate (Text-figs 32, 33, 39, 40). Variation: Very variable in colour, pattern and size. Wing pattern ranges from uniformly coloured specimens to well contrasted specimens. Male genitalia: Uncus blunt at tip, forming an almost equilateral triangle. Gnathos stout, narrower and spoon-shaped towards tip. Ventral part of bipartite valva with knife-shaped distal process clearly exceeding edge of valva; dorsal extension only hinted; costa with one thin thorn. Juxta deeply split, clearly more than halfway, juxta arms distally straight, slightly bent at tip. Aedeagus proximally with a short and distally with a longer cornutus (average length ratio of short versus long cornutus 0.75:1). Female genitalia: Corpus bursae bean-shaped, slightly narrowing at anterior end, before transition to hyaline diverticulum, somewhat less than twice length of antrum. Antrum stretched, with sclerotized and slightly arched ledges at the transition towards corpus bursae, posteriorly narrower. Lamella postvaginalis cylindrical, with sclerotized indentation, exceeding antrum. Distribution and abundance: Endemic in the Tyrrhenian region, occurring in Corsica, Sar­ dinia, Tuscan Archipelago and coast of southern Tuscany (Flamigni et al. 2016). – Common, sometimes found in numbers (P. Skou own observation). Phenology: Bivoltine, exceptionally trivoltine. From early March to mid-November in two or three generations depending on altitude (Flamigni et al. 2016; own data). Under rearing conditions, the pupae make a diapause lasting several weeks during the summer months (M. Leipnitz pers. comm.). Adults active at night, attracted to light. Biology: Larva unknown from nature, probably polyphagous. In northern Sardinia, females observed in the early evening laying eggs on the underside of flowers and ripening fruits of Daucus (M. Leipnitz pers. comm.). Reared, amongst others, on Daucus, Taraxacum, Vicia and Lonicera (Flamigni et al. 2016). Pupation in rock crevices, under rearing conditions in a relatively firm, white-grey to amber-coloured, multi-walled cocoon (M. Leipnitz pers. comm.). Habitat: Xerothermophilous, calciphile. Preferably in open, rocky habitats with bushes. In Sardinia in bushy and rocky terrain, in open bushy wastelands, rocky valleys, oak and gorse heaths, pastures (M. Leipnitz pers. comm.); from sea-level up to 1200 m, in Corsica up to 1500 m (Flamigni et al. 2016). Similar species: C. corsica can be confused with C. variegata, but is distinguished from the latter by its clearly serrated antennae of the males (see Raineri 1985b). Furthermore, C. corsica has larger dark spots in the terminal area, in particular on the underside of the wings. 91

Doubtful cases require examination of the genitalia for a reliable identification. Genitalia have different shapes of valva (male), and corpus bursae and ostium (female). In addition, the costal process of the valva is clearly developed in C. corsica whereas it is absent in C. variegata (exception: specimens from Malta with thin process). The two species are allopatric at a distance of about 70 km between the nearest collecting sites in Tuscany (Flamigni et al. 2016). Genetic data: BIN: BOLD:AAF2918 (n=5 from Sardinia). Genetically heterogeneous, one singleton from southernmost Sardinia (BOLD:ADA4171) diverging by 2.3%. Nearest species: C. dubitaria from Iran (3.8%), C. staudingeri (4.9%).

22. Charissa staudingeri (Wnukowsky, 1929) Gnophos dubitaria [“var.”] graecaria Staudinger, 1892: Dt. ent. Z. Iris 5: 192 (Greece). Syntypes 2♂ (MNHU, examined). Junior primary (and secondary) homonym of Gnophos ambiguata graecaria Staudinger, 1871. Gnophos dubitaria staudingeri Wnukowsky, 1929: Zool. Anzeiger 83 (9/10): 223. Replacement name for graecaria Staudinger, 1892. Herewith raised from subspecies of C. dubitaria (­Scoble 1999) to species rank (stat. n.), based on significant and constant differential features and on the deep COI divergence (see Genetic data).

External characters and abdomen: Wingspan 20–29 mm, second generation smaller than the first. Colour of wings varying from beige to bluish grey. Transverse lines usually clear, serrated, dark to light orange. Discal spots usually filled. Fringes brighter than ground colour, chequered. Male antennae shortly bipectinate (Text-figs 34, 41). Variation: Very variable in size and colour.

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Male genitalia: Uncus blunt at tip, forming an equilateral triangle. Gnathos stout, forming an isosceles triangle, blunt at tip. Ventral part of bipartite valva with knife-shaped distal process clearly exceeding edge of valva; dorsal extension clearly developed; costa with one, sometimes two thorns. Juxta split halfway, juxta arms slim, like tines of a fork. Aedeagus proximally with short and distally with very long cornutus (average length ratio of short versus long cornutus 0.46:1). Female genitalia: Corpus bursae long and slender, bean-shaped; not or hardly narrowing at anterior end, before transition to hyaline diverticulum; three times as long as antrum. Antrum rectangular, without sclerotized ledges at transition towards corpus bursae. Lamella postvaginalis cylindrical, exceeding antrum. Distribution and abundance: East Mediterranean (Circum Aegean). Occurring in mainland Greece, Crete, and in some Aegean islands (e.g. Samos: Fritsch et al. 2014). – In Samos common in both generations (D. Fritsch pers. comm.). – Outside Europe on the west coast of Turkey.

Phenology: Bivoltine. From late March to mid-June and from late September to mid-­ November in at least two overlapping generations (M. Leipnitz pers. comm.). Caterpillars found in early May (Wagner 2019). In Crete probably three generations from late April to mid-October (P. Skou own observation). Under rearing conditions, pupae with diapause for several weeks during summer months (M. Leipnitz pers. comm.). Hibernation as larva in different developmental stages. Adults active at night, attracted to light. Biology: Larva polyphagous on herbs and dwarf shrubs on rocks and dry grassland. Found on Euphorbia acanthothamnos (Wagner 2019), on Antirrhinum majus, Sedum sediforme (Samos: Fritsch et al. 2014) and in flowers and old fruit stalks of wild Foeniculum vulgare (Crete: M. Leipnitz pers. comm.). Reared on various small herbs (e.g. Taraxacum, Poly­ gonum aviculare, Genista) (S. Erlacher own data), and on flowers, fruits and leaves of 93

­Lonicera tatarica, Daucus carota, Bupleurum falcatum (M. Leipnitz pers. comm.). Under rearing conditions, pupation in a relatively firm, white-grey to amber-coloured, multi-walled cocoon (M. Leipnitz, pers. comm.). Habitat: Xerothermophilous, montane. Inhabiting rocky habitats, low, soft-leaved scrubland and stony, dry grassland. In Crete mainly in bushy terrain interspersed with rocks and perennials, open bushy wastelands, overgrown rocky valleys and cattle-grazed lands (M. Leipnitz pers. comm.). From sea-level up to 800 m in the Mount Parnassus, in Crete from sea-level up to 1500 m (P. Skou own observation), mainly from 200 m up to 800 m (M. Leipnitz pers. comm.), in Samos usually up to 500 m, exceptionally up to 1000 m (D. Fritsch pers. comm.). Adults flushed up from island-like growing stocks of Poterium spinosum during daytime (Crete: M. Leipnitz pers. comm.). Similar species: C. staudingeri can externally very easily be confused with C. subtaurica and C. variegata. However, the males of C. staudingeri have shortly bipectinate antennae while those of the other species are filiform. Identification of females requires examination of the genitalia: C. staudingeri has a slender, slightly curved corpus bursae, while it is more curved in the other species. In C. staudingeri, the antrum is relatively thin and not kinked as in the other species. C. staudingeri can be externally distinguished from C. dubitaria by its less ochreous ground colour and a more obscured terminal area of the wing underside. Also the genitalia differ strongly: The aedeagus of C. staudingeri has a much longer cornutus than that of C. dubitaria. In female genitalia of C. staudingeri, the corpus bursae is much longer than the equivalent in C. dubitaria. Genetic data: BIN: BOLD:AAD2249 (n=12 from Greece and Mediterranean south-western Turkey). Genetically slightly heterogeneous, maximum variation 1.2%. Nearest species: Iranian C. dubitaria (4.4%; n=8), C. subtaurica (4.8%), C. corsica (4.9%). Remarks: C. staudingeri was originally described as “variation” (subspecies) of Gnophos dubitaria (now Charissa) by Staudinger (1892a) and was considered as such for a long time. Our studies, including DNA barcoding, have shown that this taxon is not a subspecies of C. dubitaria but a bona species.

23. Charissa subtaurica (Wehrli, 1934) Gnophos variegata subtaurica Wehrli, 1934: Mitt. münch. ent. Ges. (24/2): 54, pl. 1, fig. 25/26 (Turkey: Maras, Bertiz Jaila; Syria: Akbès; Lebanon: Beirut). Syntypes ♂♀ (ZFMK, examined). ‡ variegata [“var.”] subtaurica Wehrli, 1932: Mitt. münch. ent. Ges. (22/1): 9. Non binominal, infrasubspecific, not available (see Hausmann 1994a).

External characters and abdomen: Wingspan 19–30 mm, second generation much smaller than the first. Wings with ground colour varying from grey or bluish grey to matt orange. Transverse lines usually clear, ochreous to orange. Discal spots usually well recognizable, usually unfilled. Fringes yellowish beige, chequered. Male antennae dentate (Text-figs 35, 42). Variation: Extremely variable in size and colour. Male genitalia: Uncus blunt at tip, forming an equilateral triangle. Gnathos stout, forming an isosceles triangle, blunt at tip. Ventral part of bipartite valva with knife-shaped distal process clearly exceeding edge of valva; dorsal extension clearly developed; costa with one thin thorn. Juxta split slightly more than halfway (average ratio of split versus juxta length 94

0.55:1), juxta arms slim, horn-shaped bent at tip. Aedeagus proximally with short and distally with long cornutus (average length ratio of short versus long cornutus 0.55:1). Female genitalia: Corpus bursae long, more or less bent, not or hardly narrower at anterior end, before transition to hyaline diverticulum, twice as long as antrum. Antrum slightly funnel-shaped, on left side (in ventral view) with sclerotized ledge at transition towards corpus bursae. Lamella postvaginalis cylindrical to slightly conical, exceeding antrum. Distribution and abundance: East Mediterranean. In Europe only in Samos (Fritsch et al. 2014), possibly occurring also elsewhere in the East Aegean. – Not rare, but with varying abundance, often collected in light traps. In Samos very common in both generations (D.  Fritsch pers. comm.). – Outside Europe from western, central and southern Turkey, across Cyprus and Lebanon to Israel and Jordan (Hausmann 1994a; 1995).

Phenology: Bivoltine. In Samos from early April to mid-June and again from mid-­September to mid-November (Fritsch et al. 2014; D. Fritsch pers. comm.). Outside Europe (Turkey to Israel) from mid-March to late June, at higher elevation until early August, with second generation from late September to late November. Caterpillars in Samos found in late March, May, September and December (Fritsch et al. 2014; D. Fritsch pers. comm.). Under rearing conditions, caterpillars of the first generation make a diapause of some weeks. Hibernation as small larva (M. Leipnitz pers. comm.). Adults active at night, attracted to light. Biology: Larva probably polyphagous. Found on Euphorbia acanthothamnos, Antirrhinum majus (Fritsch et al. 2014) and Teucrium divaricatum (D. Fritsch pers. comm.) and on flowers and seeds of Bupleurum rotundifolium (M. Leipnitz pers. comm.). Reared on various other plants, e.g. Taraxacum, Polygonum aviculare, Genista (S. Erlacher own observation) and Lonicera tatarica (M. Leipnitz pers. comm.). Like all species of this subgenus, caterpillars ­jerkily move back and forth when being disturbed. Pupation in rock crevices, under rearing conditions in a relatively solid, white to yellowish, multi-walled cocoon (M. Leipnitz pers. comm.). 95

Habitat: Xerothermophilous, montane. Inhabiting dry and hot oak and pine forests, rocky habitats and stony, dry grassland. In Samos from sea-level up to 1000 m, but usually below 500 m (Fritsch et al. 2014; D. Fritsch pers. comm.). Outside Europe usually from 600 m up to 1600 m (M. Leipnitz pers. comm.), sometimes down to sea-level (western Turkey: J. Gelbrecht pers. comm.), exceptionally up to 1800 m (South Anatolia), in Israel on Mt. Hermon up to 2000 m (A. Hausmann own data). Similar species: Externally, C. variegata cannot be distinguished from C. subtaurica. Hausmann (1994a; 1995) mentioned the following differences, which can be confirmed here with slight modifications: Male genitalia of C. subtaurica with slightly wider basal processes of juxta, an almost always present thorn in the middle of valva costa, and a slightly narrower and longer distal cornutus in the aedeagus. The juxta arms are shorter than those of C. variegata and not as deeply split as in the latter. In female genitalia, the lateral sclerotization from the lamella postvaginalis towards the corpus bursae is less pronounced. In addition, the corpus bursae of C. subtaurica is usually slightly longer than that of C. variegata. Genetic data: BIN: BOLD:AAC4341 (n=19 from Greece, Cyprus, Turkey, Lebanon, Israel, Jordan). Genetically homogeneous, maximum variation 0.7%, but one singleton from central Israel at a minimum distance of 1.1%. Nearest species: C. variegata (1.4%).

24. Charissa variegata (Duponchel, 1830) Gnoph[os] variegata Duponchel, 1830: in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 8 (1): 216, pl. 184, fig. 4; pl. 185, fig. 8 (France: Grenoble). Syntypes (MNHN, not examined). Gnophos variegata sicula Wehrli, 1931: Mitt. münch. ent. Ges. 21: 45 (Italy, Sicily: Catania). Syntypes 4♂ (ZFMK, examined). Downgraded from subspecific rank of Charissa variegata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies by Flamigni et al. (2016). Gnophos variegata cava Vojnits, 1968: Annls hist.-nat. Mus. natn. hung. 60: 222, figs 2, 3a (Hungary: Odvashegy, Budaörs). Holotype ♂ (TMB, not examined). Herewith downgraded from subspecific rank of Charissa variegata (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.).

External characters and abdomen: Wingspan 20–32 mm, second generation much smaller than the first. Wings with ground colour varying from whitish orange through grey or greyblue to matt orange. Transverse lines mostly clear, ochreous to orange. Discal spots usually well recognizable and unfilled. Fringes yellowish beige, more or less chequered. Male antennae slightly dentate (Text-figs 36, 43). Variation: Extremely variable in size and colour. Specimens from Sicily, originally described as subspecies sicula by Wehrli (1931), sometimes darker, ground colour dark grey with dark violet tinge, discal spots usually filled on forewing, usually unfilled on hindwing. According to Flamigni et al. (2016) it is only an individual form. Specimens from Malta are smaller than the nominotypical subspecies and usually pale beige or whitish orange. Male genitalia: Uncus blunt at tip, forming an equilateral triangle. Gnathos stout, forming an isosceles triangle, blunt at tip. Ventral part of bipartite valva with knife-shaped distal process clearly exceeding edge of valva; dorsal extension clearly developed; costa without thorn (exception: specimens from Malta). Juxta deeply split over clearly more than halfway (average ratio of split versus juxta length 0.6:1), juxta arms slim and relatively long, horn-shaped bent

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at tips. Aedeagus proximally with short and distally with very long cornutus (average length ratio of short versus long cornutus: 0.48:1). Female genitalia: Corpus bursae long and more or less bent, not or hardly narrower at anterior end, before transition to hyaline diverticulum, twice length of antrum. Antrum slightly funnel-shaped, on left side (in ventral view) with sclerotized ledge at transition towards corpus bursae. Lamella postvaginalis cylindrical to slightly conical, exceeding antrum. Distribution and abundance: South European. Distributed in the central and eastern part of southern Europe, from western and southern France (Lhomme 1923–1935) to Bulgaria and Crimea, in the north to eastern France (Eure, Haute-Saône; Sauvagère 1991; Jugan 2001), Switzerland (up to Basel; Altermatt et al. 2006), Austria (Tyrol and Carinthia), Hungary and Romania and southwards to Sicily and Greece. – In large parts of the distribution area common. – Not found outside Europe. Records from Morocco (Rungs 1981) refer to Charissa rothschildi (Prout, 1929), which is systematically much closer to C. mucidaria than to C. variegata. Records from Turkey, Cyprus, Israel, Lebanon and Jordan refer to C. subtaurica.

Phenology: Bivoltine, exceptionally trivoltine. From late April to late July and from mid-­August to mid-October. Under good conditions in southern Europe emerging from February with records in all months until December, likely in three generations (Flamigni et al. 2016). At altitudes above 1000 m, the flight period is shorter, from mid-May to early September, at 1700 m there may be only one generation (Flamigni et al. 2016). Larva found from August to November and, again, after overwintering, from early March to mid-May. Hibernation in different developmental stages (L2–L4) while feeding from time to time on Sanguisorba minor (M. Leipnitz pers. comm.). Larvae of summer generation from May to July (Flamigni et al. 2016). Adults active at night, attracted to light, also found during daytime in shady spots of sunlit rocks. 97

Biology: Larva polyphagous. Found on Rumex, Petrorhagia, Sedum, Sempervivum, Clinopodium vulgare, Verbascum, Antirrhinum, Linaria, Cymbalaria muralis, Plantago, Achillea, Centaurea, Asplenium ceterach, A. trichomanes, A. ruta-muraria, Clematis vitalba, and C. flammula (Flamigni et al. 2016), on Scrophularia (Wagner 2019), on Centranthus, Chamaecytisus, and Thymus (Ziegler 2019). Found in upholstery and flowers of Thymus (Corfu: M. Leipnitz pers. comm.). Reared on Lonicera xylosteum (Flamigni et al. 2016, after J. Gelbrecht), L. tatarica (M. Leipnitz pers. comm.). Pupation in a whitish cocoon at the base of the food-plant (Flamigni et al. 2016), under rearing conditions pupation in a relatively firm, white-grey, multi-walled cocoon (M. Leipnitz pers. comm.). Habitat: Xerothermophilous. Inhabiting coastal areas, hills and mountainous areas, hot and dry places, like Mediterranean macchia, sunny hills partially covered with steppe-like grassland and rocks, rocky or stony slopes, embankments and old quarries. In Corfu, on slopes with rich vegetation, open wastelands and cattle pastures interspersed with rocks, bushes and shrubs (M. Leipnitz pers. comm.). From sea-level usually up to 1000 m (Flamigni et al. 2016; A. Hausmann own observation), up to 1900 m in the Alps. Similar species: Externally, C. subtaurica cannot be distinguished from C. variegata. Hausmann (1994a; 1995) mentioned the following differences, which can be confirmed here with slight modifications: Male genitalia of C. variegata with slightly narrower basal processes of juxta and without thorn in the middle of valva costa (exception: specimens from Malta). Furthermore, C. variegata has a slightly wider and shorter distal cornutus in the aedeagus. The juxta arms are longer than those of C. subtaurica and more deeply split than in the latter. In female genitalia, the lateral sclerotization from lamella postvaginalis towards corpus bursae is more pronounced. In addition, the corpus bursae of C. variegata is usually slightly shorter than that of C. variegata. Genetic data: BINs: BOLD:ACF2465, BOLD:ACF2468, BOLD:ACE9838, BOLD:ACE9839, BOLD:AAC1039, BOLD:ACF1036, (n=27 from Austria, Hungary, Romania, Italy, Malta, Croatia, Greece). Genetically very heterogeneous, maximum variation 5.2%, with many different haplotypes characteristic for certain regions. One specimen from Malta belonging to the widespread Italian BIN BOLD:AAC1039, at a distance of only 0.6%. All six BINs clustering together, being well separated from the nearest species C. subtaurica at a minimum distance of 1.4%. Remarks: Specimens from Malta (examined from mid-May and late July) are usually smaller and brighter, and in the male genitalia the costa bears a thin thorn, which is absent in the nominotypical form. Interestingly, no significant genetic differences were detected in DNA barcoding (see under Genetic data). Leraut (2009: 231) mentioned “subvariegata Staudinger, 1898” as a subspecies of C. variegata, which, however, is a species of the subgenus Cnestrognophos Wehrli, 1951 and does not occur in Europe. The Moroccan taxon Gnophos variegata rothschildi Prout, 1929 (Novitates Zoologicae 35: 145; types in NHMUK, examined) was still treated as subspecies of C. variegata in Scoble (1999). After detailed analysis we raise its status to species rank: Charissa rothschildi (Prout, 1929) (stat. n., comb. n.), based on constant and significant differential features and on the deep divergence in the COI gene. In male genitalia, the aedeagus of C. rothschildi has proximally a flat and very short cornutus-like sclerite and distally a dagger-shaped, very long cornutus, thus more similar to C. mucidaria than to C. variegata. In female genitalia, 98

the bursa copulatrix is very long, separated into a tubular ductus bursae and a pipe-shaped corpus bursae, similar to that of C. mucidaria. However, C. rothschildi clearly differs from the latter by its dentate male antennae which are distinctly bipectinate in C. mucidaria.

25. Charissa mucidaria (Hübner, 1799) Geometra mucidaria Hübner, 1799: Samml. Eur. Schmett. 5: pl. 28, fig 148 (Europe). Syntype(s). Gnophos mucidaria [“var.”] subsignaria Staudinger, 1892: Dt. ent. Z. Iris 5: 190 (Italy, Sicily: Kalchberg). Syntypes 1♂1♀ (MNHU, examined). Downgraded from species rank (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos mucidaria [“var.”] ochracearia Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (3rd Edition): 346 (Spain: Murcia; ?Ronda; Mauretania). Syntypes 6♂ (MNHU, examined). Herewith downgraded from subspecific rank of C. mucidaria (Scoble 1999, Leraut 2009) to synonymy, based on clinal character transitions towards nomino­ typical subspecies (syn. n.). Gnophos mucidaria [“var.”] lusitanica Mendes, 1903: Brotéria 2: 67 (Portugal, S. Fiel: Beira Baixa). Syntype(s) (not examined). Herewith downgraded from subspecific rank of C. muci­ daria (Scoble 1999; Leraut 2009) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos mucidaria herrichii Oberthür, 1913: Suite de la Revision des Phalénites 7: 301, pl. 178, fig. 1735 (France, Eastern Pyrenees). Holotype ♀ (not examined). Synonym of C. muci­ daria according to Scoble (1999). Gnophos mucidaria [“ab. (var.)”] grisearia Prout, 1915: in Seitz, Gross-Schmett. Erde 3: 391 (France [south], Spain: Andalusia). Syntypes (MNHU, 1♀ syntype from Andalusia (Ronda) examined). The original description by Staudinger (1871: 167), later cited by Prout (1915: 391), referred to a single female from Ronda, showing individual variation (Staudinger 1892a: 191), whilst Prout based his description on material from both France and southern Spain. Herewith downgraded from subspecific rank of C. mucidaria (Scoble 1999) to synonymy, based on clinal character transitions towards nominotypical subspecies (syn. n.). Gnophos nubilarius Reisser, 1936: Ent. Rdsch. 53: 145, pl. 3, figs C4–6, C8, D4–6, D8 (Spain: Sierra de Gredos, Garganta de las Pozas, 1900 m; Laguna de Gredos, 2000 m) Syntype(s) (NHMW, examined). Synonym of C. mucidaria according to Scoble (1999).

External characters and abdomen: Wingspan 19–31 mm, second generation much smaller than the first. Wings with ground colour varying from whitish through beige or ochreous to dark grey, with numerous dark scales arranged into longitudinal stripes, therefore speckled. Transverse lines usually clear, often blurred, brown to ochreous or orange. Discal spots of forewings usually relatively large and conspicuous, usually unfilled, on hindwings present but usually smaller, usually filled. Fringes of ground colour, more or less chequered. Male antennae clearly bipectinate (Text-figs 37, 44). Variation: Extremely variable in wing shape, colour and pattern: Coloration varying from almost whitish to heavily darkened with all conceivable transitions, depending on the substrate. Specimens occurring on granite substrates are often heavily speckled, while specimens on reddish sandstone soils are camouflaged by a reddish tint. The shape of the forewings can vary from slightly rounded to narrow with tapered apex. Male genitalia: Uncus slightly pointed, forming an equilateral triangle. Gnathos stout, forming an isosceles triangle, blunt at tip. Ventral part of bipartite valva with knife-shaped distal process clearly exceeding edge of valva, dorsal extension clearly developed, almost parallel 99

to distal process, but usually shorter than the latter, costa with a strong thorn. Juxta deeply split, clearly more than halfway (average ratio of split versus juxta length 0.63:1), juxta arms slim and straight, slightly pointed at tips. Saccus conspicuous, long. Aedeagus proximally with a flat and very short cornutus-like sclerite and distally with a dagger-like and very long cornutus (average ratio of short versus long cornutus 0.43:1). Female genitalia: Bursa copulatrix very long, separated into a tubular duct (= ductus bursae) and a pipe-shaped corpus bursae; the former three to four times length of antrum and with strongly sclerotized, longitudinal folding, the latter with a hyaline diverticulum and a dome-like appendix bursae bearing a set of large teeth. Antrum slightly funnel-shaped or rectangular. Lamella postvaginalis cylindrical, exceeding antrum. Distribution and abundance: West Mediterranean, with extension towards North-east Africa. In Europe from Iberian peninsula, France (up to Eure and Seine-et-Marne: Sauvagère 1991; Purser 2007; Thibaudeau et al. 2013), western Switzerland (only from Geneva: S ­ wissLepTeam 2010) to north-westernmost Italy. The species also occurs on several Mediterranean islands, including the Balearic Islands, Corsica, Sardinia (1♂, Aritzo, 10.v.1935, in coll. ZSM), Sicily, Malta, and Lampedusa (Flamigni et al. 2016). Also reported from Crete (Müller 1996) but not yet confirmed and therefore probably erroneous (Ruckdeschel 2007). Reports from the Balkan peninsula (Slovenia, Croatia, Bosnia) most likely refer to C. variegata (­Rebel 1904; Flamigni et al. 2016). – In large parts of the distribution area common or rather common and usually found in numbers. – Outside Europe from Morocco, Algeria, Tunisia, Libya to Egypt (examined). Reputedly in Turkey (Riemis 1994; Koçak & Kemal 2009), probably misidentified.

Phenology: Bi- or trivoltine. From February to mid-November in two or three generations. In southernmost Spain in all months of the year (P. Hale pers. comm.). In Italy from ­early March to mid-June and from mid-September to late October, at higher altitudes (e.g. 100

1100 m, Madonie, Sicily; 1600–2000 m, Sierra de Gredos, Spain) in only one generation from mid-June to late August (Flamigni et al. 2016; own data). In North Africa from late January to late October in two or three generations (own data). Overwintering as larva (Meyer 1919). Caterpillars found from March to October (Flamigni et al. 2016). Adults active at night, attracted to light. Biology: Larva polyphagous. Found on Rumex, Anagallis arvensis, Polygonum avicu­ lare, Sedum album, Trifolium (Leraut 2009; Redondo et al. 2009; Flamigni et al. 2016), and ­under Antirrhinum majus (G. King, pers. comm.). Reared on Taraxacum (S. Erlacher own observation). Recorded on Umbilicus rupestris and Sarcocapnos crassifolia (Morocco: Rungs 1981). Habitat: Xerothermophilous. Inhabiting hills and mountains, on hot and dry rocky places, such as steppe-like grassland, abandoned quarries, groves of Quercus pubescens and pine forest (Flamigni et al. 2016). From sea-level up to 1200 m, exceptionally up to 1780 m in Val di Susa (Italy) (Flamigni et al. 2016) or 2250 m in Sierra Nevada (Spain: Redondo et al. 2009; P. Skou own data). In North Africa from 300 m around Tataouine (Tunisia) up to 2400 m near Oukaïmeden (Morocco). Similar species: Males of C. mucidaria can be easily distinguished from similar species by their antennae, which are slightly dentate in C. variegata and C. subtaurica, more distinctly dentate in C. assoi, serrated in C. corsica, shortly bipectinate in C. staudingeri, while distinctly bipectinate in C. mucidaria. For correct identification of females sometimes dissection of genitalia is required, those of C. mucidaria differing from all other species of this subgenus in having a very long and slender ductus bursae and a dome-like appendix bursae with a set of large teeth. Genetic data: BIN: BOLD:AAB5230 (n=46 from France, Spain, Portugal, Morocco, Tunisia). Genetically homogeneous in the main cluster (maximum variation 0.7%), specimens from southern France (with almost identical sequences in Tunisia) diverging by 0.6%. In Tunisia with a second cluster, similarly diverging from the main cluster of C. mucidaria. Nearest species: C. rothschildi from Morocco (4.1%). Remarks: The name “catenulata” was introduced by Rambur (1866: pl. 19, fig. 5) figuring a specimen from Andalusia without accompanying text and without assignment to genus (uninominal, thus nomenclaturally unavailable). It has long been considered to be a synonym of C. onustaria. Wehrli (1926) treated it as a “form” of Charissa mucidaria (formerly Gnophos) from Granada and the Sierra Nevada. Examination of types, however, suggests synonymy with C. onustaria (see there), based on likely mislabelled type material from Corsica. C. mucidaria tends to develop substrate-adapted forms that recur at distant locations. In all cases, however, descriptions of subspecies are based on colour-deviant individuals within more or less uniform populations. This applies to the following individual forms, which are therefore synonymized here with the nominotypical subspecies: Gnophos subsignaria Staudinger, 1892 (syn. n.), Gnophos mucidaria ochracearia Staudinger, 1901 (syn. n.), Gnophos mucidaria lusitanica Mendes, 1903 (syn. n.), and Gnophos mucidaria grisearia Prout, 1915 (syn. n.).

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Subgenus Kemtrognophos Wehrli, 1951 Kemtrognophos Wehrli, 1951, Lambillionea 51: 11. Type species: Gnophos onustaria Herrich-­ Schäffer, 1852. Validated at genus rank by Viidalepp (1996) and Redondo et al. (2009), at infrageneric (subgenus) rank by Scoble (1999) and Leraut (2009).

Diversity and distribution: Subgenus Kemtrognophos with ten currently validated species, distributed from southern Spain (Andalusia) across southern and central Europe to eastern Europe. Outside Europe from Turkey and Levant across Transcaucasus, Iran to central Asian mountains, Mongolia and Korea in the east. One species in Morocco: Charissa idrisarius (Zerny, 1934), comb. n., requiring transfer from Gnophos (Scoble 1999) to Charissa (Kemtrognophos) after analysis of genitalia. External characters and abdomen: Medium-sized, wingspan 24–37 mm. Ground colour of wings brown to grey, often speckled. Discal spots usually ring-shaped on forewing upperside. Hindwing termen concavely arched between veins. Antennae of both sexes filiform, male antennae slightly thicker (exception: North African C. idrisarius with bipectinate male antennae). Further characters see description under genus Charissa. Proboscis present. Hindtibia of both sexes with two proximal and two distal spurs. Male genitalia: Uncus rounded at tip. Gnathos well developed. Posterior juxta arms broad, moderately long or long, anterior part of juxta small. Valva narrow and tapering towards tip. Costa of valva at centre with short, digitiform process (ampulla) bearing one or two terminal thorn(s), in C. zeitunaria appressed to costa. Non-European C. argillata (Brandt, 1938) and C. mardinaria (Staudinger, 1901) without costal process. Shape of cornutus diagnostic, either long and straight, or compact and strongly bent. Non-European C. ciscaucasica (­Rjabov, 1964), however, without cornutus. Female genitalia: Shape of corpus bursae diagnostic, usually long and slender over most of its length, strongly sclerotized, longitudinally furrowed, anteriorly with hyaline sac. Non-­ European C. ciscaucasica, however, without anterior hyaline sac. C. ambiguata and C. predotae with less compact and less sclerotized corpus bursae. The group of C. zeitunaria, C. argillata and C. mardinaria with long apophyses and sclerotized papillae anales with stiff setae. Differential diagnosis: Subgenus Sacrognophos (type species C. sacraria (Staudinger, 1895) from the Levant) differs from Kemtrognophos species in bipectinate male antennae (filiform in Kemtrognophos), in male genitalia cornutus developed as triangular sclerites (usually rod-shaped in Kemtrognophos). Biology, phenology, habitat: Larval hostplants only known for three of the five Euro­pean species, oligophagous on Polygonaceae (C. ambiguata, C. predotae) or polyphagous (C. onustaria). Adults of subgenus Kemtrognophos inhabiting montane habitats. Phenology uni- or bivoltine. Active at night, readily attracted to light. Immature stages: In C. ambiguata with shallow, dorso-lateral humps on 3–5 mid-segments, in C. onustaria and C. predotae densely covered with warts, without dorsal humps (­Euroleps 2018; Lepiforum 2018; own data). Pupa with two apical and two subapical hooked bristles on the cremaster (Euroleps 2018).

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26. Charissa ambiguata (Duponchel, 1830) Gnophos ambiguata Duponchel, 1830: in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 8 (1): 223–224, pl. CLXXXVI, fig. 2 (France: Basses Alpes). Syntype(s) (MNHN; not examined). Tephrosia ambiguaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 198. Unjustified emendation (cf. GME Vol. 3, p. 12). Gnophos meyeraria de la Harpe, 1853: Fauna Suisse Lépid. 4: 65, fig. 2 (Switzerland: Jura, near Bienne and Arbourg). Syntypes 2♂1♀. Gnophos ophthalmicata Lederer, 1853: Verh. zool-bot. Ges Wien, 3: 379, pl. V, fig. 5 (Siberia). Holotype ♂ (ZMB; not examined). Described at species rank, retained as synonym in Scoble (1999), here recognized at subspecies rank, based on slight differences in habitus (stat. n.). Gnophos pullularia Herrich-Schäffer, 1852: Syst. Bearb. Schmett. Eur. 6 (55): 72; ibidem (1851), 3; pl. LXXXI, fig. 498, 499 (Europe). Syntype(s). Retained as an infrasubspecific form in Leraut (2009), validated at subspecies rank in Scoble (1999), the latter accepted here. Gnophos vepretaria Speyer, 1867: Stettin. ent. Ztg. 28 (7–9): 352 (Poland: ‘Prussia’, Danzig). Syntype(s). Synonymized with subsp. pullularia in Scoble (1999). ‡ Gnophos obfuscaria: sensu Vorbrodt (1913: 176, 181) nec (Denis & Schiffermüller, 1775), misidentification. Vorbrodt explicitly refers to Hübner as original author and lists ­ambiguata as synonym. Vice versa, in Osthelder (1931: 520) obfuscaria (under the authorship of H ­ übner) is listed as synonym of ‘Gnophos ambiguata‘. Gnophos sheljuzhkoi Schawerda, 1924: Zeits. Österr. Ent.-Ver., 9 (10): 87–89 (Russia: distr. Belebej, station Belebej-Aksakon, Aksenovo). Lectotype ♂ (ZMKU, examined), designated by I. Kostjuk in Anikin et al. (2017), paralectotype figured under fig. 26e. Validated at species rank in Scoble (1999) and Leraut (2009, without examination), herewith downgraded to subspecies rank (stat. n.) of C. ambiguata based on diverging coloration without showing significant differential features in genitalia (see Remarks). Gnophos ambiguata tatrensis Vojnits, 1969: Opusc. Zool. Budapest, 9 (2): 375 (Hungary: High Tatra, Tátraszéplak). Holotype ♂ (TMB; not examined). Validated at subspecies rank in ­Scoble (1999), herewith downgraded to synonymy of subsp. pullularia based on the absence of significant and constant differential features (syn. n.). Kemtrognophos ambiguatus [sic!] viidaleppi Vojnits, 1975: Annales historico-naturales Musei nationalis hungarici, 67: 191–192, fig. 5 (Mongolia, Chövsgöl Aimak, near Somon Cecerleg and Somon Bajan-ul, 65 km west of Cecerleg, 1700 m). Holotype ♂ (TMB; not examined). Validated at subspecies rank in Scoble (1999), herewith downgraded to synonymy (syn. n.) of subsp. ophthalmicata based on the absence of significant and constant differential features. ‡ Charissa graecaria: sensu Scoble (1999) nec Staudinger, 1871. Erroneously validated as subspecies of C. ambiguata in Scoble (1999). See under subgenus Trilobignophos and see remarks to this species. Charissa herbuloti Leraut, 2009: Moths of Europe Vol. 2: 224–226, figs 117, 118, pl. LXI, figs 13–15 (France: Pyrénées-Orientales). Holotype ♂ (MNHN; examined; furthermore 1♀ paratype examined). Herewith downgraded from species rank to subspecies rank under C. ambi­ guata, based on constant differential features in habitus, but considering exact COI barcode sharing with the nominotypical subspecies (stat. n.). More research is needed to examine potential clinous transitions towards the nominotypical subspecies. See also Remarks. Unavailable names (infrasubspecific): nigrescens: Hannemann (1912).

External characters and abdomen: Wingspan males 26–29 mm, females 25–33 mm. Hindwing termen arched between veins. Ground colour of wings light to medium grey-brown, slightly shining. Discal spots ring-shaped, medium to dark brown. Transverse lines ­medium to dark brown. Antemedial line of forewing weak, with three inwardly directed jags. Post­ medial line of forewing dentate, with tips of jags darker than brown colour of line, near origin

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at costa strongly projecting towards termen, here bent at right angle. Subterminal line pale. Termen with dark brown or black dots. Fringes slightly chequered. Hindwing without antemedial line. Underside of forewing with light brown to light grey ground colour, discal spot weak, dark brown to dark grey, postmedial line dark brown, darker towards apex. Under­ side of hindwing with light brown to light grey ground colour, paler than forewing, discal spot dark brown to dark grey but very weak, postmedial line indistinct. Frons dark brown, vertex white. Variation: Variable. Ground colour of nominotypical subspecies uniform brown with pale grey areas, transverse lines and discal spots dark brown, subterminal line pale. Central and eastern European subsp. pullularia with uniform dark brown ground colour, transverse lines and discal spot slightly darker than ground colour. Subsp. herbuloti from the Pyrenees with comparatively dark ground colour (due to suffusion with brown scales), postmedial lines and antemedial line of forewing strongly marked, at inner termen approaching; delimitation from nominotypical subspecies awaiting further study. Subsp. sheljuzhkoi from southern European Russia with snow white ground colour, with very slight grey or brown suffusion, transverse lines and discal spots dark grey. Postmedial line often broken, with just the tips of the jags being marked. Subsp. ophthalmicata from central and eastern Asia with pale brown to grey ground colour, partially mottled, transverse lines and discal spot brown to dark grey. Specimens from North Korea with brown to grey ground colour, transverse lines and discal spots dark brown. Male genitalia: Uncus blunt at tip, elongate triangular. Gnathos well developed. Saccus long and wide. Juxta long, with inwards curved, narrow posterior juxta arms, their length three times length of anterior juxta arms. Valva wider at base, narrowing towards pointed apex, apex strongly setose, costal margin with a long, pointed thorn. Aedeagus straight, asymmetric at tip, cornutus stout, bent and pointed at tip, dilated at base, longitudinally furrowed. Length of cornutus (0.6–0.7 mm) half length of aedeagus (1.2–1.4 mm). No significant differences between subspecies, as also shown for subsp. herbuloti in the genitalia figures of the original description (Leraut 2009), though the latter shows slightly shorter aedeagus and cornutus, thus being transitional to the following species in male genitalia. Female genitalia: Antrum posteriorly broad, hyaline, with wide invagination. Ductus bursae narrowing anteriorly, constricted. Corpus bursae anteriorly dilated to a hyaline sac, at centre strongly sclerotized and longitudinally furrowed, sclerotization extended over 2/3 of length of corpus bursae, sometimes more. Length of bursa copulatrix 1.2–1.6 mm. Distribution and abundance: Euro-Siberian, montane. Nominotypical subspecies from southern France (incl. Massif Central) across Switzerland to northern Italian, Bavarian and Austrian Alps, and in the French, Swiss and south-west German Jura mountains. In the Pyrenees replaced by subsp. herbuloti Leraut, 2009 (delimitation from nominotypical subspecies requiring further study). Subsp. pullularia from central Germany to the Baltic States, and across Czech Republic and Slovakia to the Ukraine, and across European Russia to central Urals (Viidalepp 1996). Populations of the Balkan peninsula tentatively attributed to subsp. pullularia (see Genetic data). Scattered distribution in Poland, northern Germany, in Slovenia and very local in westernmost Hungary (‘Szakonyfalu’: Vojnits 1967). Extinct (without records after 1980) from northern Austria except for one single site near the 104

­Slovakian border (Huemer et al. 2009). Listed as present in Romania (Müller 1996; Hausmann et al. 2004; 2011a), but no records given in Szekely (2011). Putative records from the European part of Turkey (Okyar & Mironov 2008) based on confusion with C. onustaria (Okyar pers. comm.). Exact delimitation of subsp. pullularia from nominotypical subspecies sometimes difficult and requiring further study. Subsp. sheljuzhkoi distributed in southern European Russia, on the coast of the Black Sea and the Caucasian foothills, in southern Urals and in north-westernmost Kazakhstan (Viidalepp 1996), delimitation from subsp. pullularia and nominotypical subspecies awaiting further study. Absent from Iberian peninsula south of Pyrenees (Redondo et al. 2009) and from many northern regions of Europe, i.e. United Kingdom, Fennoscandia, northern and western France, ‘Benelux’ countries, Estonia) (Vojnits 1967). – In Europe local, in the Alps more widespread, not rarely in numbers at its localities. – Outside Europe represented by subsp. ophthalmicata in southern Siberia, Altai and Sayan mountains, Transbaikalia, in the east to Mongolia (Uliastai) and Russian republics Magadan and Sakha (former Yakutia) (Viidalepp 1996; Mironov et al. 2008).

Phenology: Univoltine, rarely with incomplete second generation. Usually from early June to mid-July, sometimes earlier or later, in south-western Germany with late flying specimens until mid-August (Bartsch 2003). Subsp. ophthalmicata until late July. Exceptionally, under good conditions, emerging from late April and with partial second generation until late September, e.g. in the southern Alps (Wehrli 1953; Rezbanyai-Reser & Expósito-Hermosa 1988; Flamigni et al. 2016). Hibernation in the ground as half-grown larva (Wehrli 1953; Bartsch 2003; Euroleps 2018). Active at night, both sexes readily attracted to light. Biology: Larva possibly polyphagous. Recorded on various low-growing plants like Plan­ tago, Campanula, Taraxacum, Lactuca, Artemisia, Chrysanthemum, Vaccinium uliginosum, Rubus idaeus, Ranunculus, and Sedum, also on Betula, Abies. (Koch 1988; Bartsch 2003; Redondo et al. 2009; Leraut 2009; Flamigni et al. 2016), probably most records referring 105

to rearings. Egg deposition observed on Sedum rupestre (Bergmann 1955; Bartsch 2003). Pupation between dry leaves (Flamigni et al. 2016). Habitat: Montane. Sun-exposed rocky slopes, screes, low-nutrient grassland on rocky soils, also in scattered coniferous forests (Bartsch 2003). In Hungary found in Luzulo-Fagetum and Dicrano-Pinetum callunetosum (Speyer 1868). In the north in the lowlands and on hills, e.g. in central Germany from 200 m up to 700 m (Erlacher 2019), in south-western Germany from 400 m up to 1200 m (Bartsch 2003), in the south of the distribution usually from 600 m up to 2000 m, but also here exceptionally down to 100 m or up to 2600 m (Flamigni et al. 2016). In central Asia up to 2700 m. During daytime resting on rocks, fenceposts and stems of pines (Speyer 1868 Bartsch 2003). Similar species (see Text-figs 45–48): Secure identification of C. ambiguata sometimes difficult due to large variation in wing pattern and coloration. C. predotae differing in much shorter aedeagus, cornutus more strongly bent and relatively short (1/3 length of aedeagus; half length in C.  ambiguata), cf. Rezbanyai-Reser & Expósito-Hermosa (1988). In female genitalia of C. predotae the sclerotized part of corpus bursae is much shorter and restricted to the posterior half or less (2/3 or more in C. ambiguata). Genetic data: Two BINs: BOLD:ACE4202, BOLD:ACF4447 (n=46 from Germany, Austria, France, Italy, Bulgaria, Russia, Kazakhstan, Mongolia, Korea). Genetically heterogeneous, shallowly split into two clusters at a distance of 1.1% from each other, the first BIN (BOLD:ACF4447) with specimens from southern France and northern Italy, across western Austria to southern Germany (thus including the area of the nominotypical subspecies), the other (BOLD:ACE4202) from south-easternmost Austria and Bulgaria across the Urals to Mongolia and Korea (several subspecies). Two DNA barcodes from the distribution area of subsp. herbuloti (Pyrenees) barcode-sharing with the nominotypical subspecies. Nearest species: C. predotae (1.7%) and C. onustaria (2.7%). Remarks: Gnophos sheljuzhkoi was described as species of Gnophos by Schawerda in 1924 and combined at species rank with subgenus Kemtrognophos by Wehrli (1953). Although Viidalepp (1996) recognized that taxon as a subspecies of C. ambiguata, this new status was not accepted by Müller (1996), Scoble (1999), Anikin et al. (2000) and Leraut (2009), based on assumption without any explanation. Extensive morphological examinations showed conspecifity of sheljuzhkoi with C.  ambiguata; we herewith propose subspecific status. Leraut (2009) described the ‘species’ C. herbuloti based on four males and two females from French Pyrénées Orientales, moreover mentioning a distribution area in central Spain, but without listing Spanish material. Because of the COI barcode sharing and very similar genital morphology of herbuloti and C. ambiguata we downgrade the former to a subspecies of the latter. This corresponds to the results of Rezbanyai-Reser & Expósito-­Hermosa (1988) and Redondo et al. (2009) who attributed the populations of the Pyrenees to C. ambiguata. The taxonomic status of herbuloti was examined based on two COI barcode sequences from the eastern Pyrenees (locus typicus) and on photos of the type specimens (Leraut 2009) and genital dissections of the male holotype and one female paratype.

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Text-figs 45–48. Diagnostic characters (indicated) on upperside of wings of selected Euro­pean Charissa species, subgenus Kemtrognophos. Text-fig. 45. C. ambiguata (Duponchel, 1830), male. Austria: North Tyrol, North Umhausen, 23.vi.1994 (coll. Erlacher). Text-fig. 46. C. predotae (Schawerda, 1929), male. Spain: prov. Teruel, Bronchales, 29.vi.1962 (coll. ZSM). Textfig. 47. C. onustaria (Herrich-Schäffer, 1852), male. Bulgaria: Pirin Mountains, Liljanovo, e.o. 15.ix.1982 (coll. B. Müller). Text-fig. 48. C. remmi (Viidalepp, 1988), male. Russia: Burjatia, Sajan Mountains, Irkuta, 16.vi.–13.vii.2013 (coll. R. Müller).

27. Charissa predotae (Schawerda, 1929) Gnophos predotae Schawerda, 1929: Verh. zool-bot. Ges Wien, 79 (1): 46–47 (Spain: Albarracin). Holotype ♂ (NHMW; not examined). ‡ Charissa onustaria: sensu Leraut (2009, partim: Moths of Europe Vol. 2: pl. LXII, fig. 6) and Wehrli (1953, partim: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 585) nec Herrich-Schäffer, 1852. Misidentification. Leraut’s figure shows a female from San Ildefonso, central Spain which is far outside the area of distribution of C. onustaria. Similarly, specimens of ‘C. onusta­ ria’ are mentioned for San Ildefonso, central Spain, and San Fiel, Portugal in Wehrli (1953).

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External characters and abdomen: Wingspan 24–31 mm. Hindwing termen arched between veins. Ground colour of wings dull pale brown, medium brown mottled. Discal spots ring-shaped, dark brown, sometimes elongate. Transverse lines dark brown to dark grey. Antemedial ‘line’ for forewing usually with very dark, partially black jags. Postmedial line dentate, with tips of jags almost black, proximally paler. Subterminal line brown to grey, near costa darker, fading towards inner termen. Termen with black dots. Fringes slightly chequered. Antemedial line absent from hindwing, subterminal line more or less broken into darker shades. Underside whitish grey, with slight brownish tinge. Discal spots dark brown or greyish. Postmedial line inconspicuous. Apical area of forewing dark brown. Variation: Variable in the intensity of the wing pattern and in the degree of dark suffusion. populations from Andalusia much darker grey or grey-brown. Male genitalia: Uncus comparatively broad, blunt at tip. Gnathos well developed. Saccus moderately long. Juxta moderately long, posterior juxta arms thick and distant from each other, straight or slightly curved inwards, inner side slightly wavy. Length of posterior arms 2–3 times length of anterior arms. Valva wider at base, narrowing towards pointed apex, apex moderately and shortly setose, ampulla on costal margin with a long, pointed thorn. Aedeagus narrow at centre, blunt at tip. Cornutus stout, bent and pointed at tip, dilated at base, longitudinally furrowed. Length of cornutus (0.4–0.45 mm) 1/3 length of aedeagus (0.95–1.1 mm). Female genitalia: Antrum posteriorly broad, alate, with wide shallow posterior invagination. Ductus bursae short, comparatively broad, slightly sclerotized. Corpus bursae anteriorly dilated to a globular hyaline sac, in the posterior half (or less) strongly sclerotized and longitudinally furrowed. Length of bursa copulatrix 1.1–1.6 mm. Distribution and abundance: (South-west) European. Distributed in the centre of the I­berian peninsula from prov. Teruel (Aragon) in the east to São Fiel (Portugal) in the west. ­Isolated occurrences in Andalusia and towards the north, e.g. prov. León, but allopatric with C. ambi-­

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guata herbuloti from the Pyrenees (Redondo et al. 2009). One single and very isolated record from Sicily (Hausmann 1993a; ZSM, dissected) not confirmed by further specimens, possibly mislabelled. – Local, usually in limited numbers. Phenology: Bivoltine. Flight period, depending on altitude, from early April to mid-July and from mid-August to late September (Redondo et al. 2009; own data), exceptionally emerging from March. Hibernation as a half grown larva. Active at night, both sexes readily attracted to light. Biology: Larva oligophagous on Polygonaceae. Recorded on Polygonum aviculare and Rumex acetosa (Mendes 1913; Redondo et al. 2009). Habitat: Xeromontane. On rocky slopes of hills and mountains. Vertical distribution from 500 m up to 2000 m (Redondo et al. 2009). Similar species (see Text-figs 45–48): On average smaller and with paler ground colour than its closest related species, C. ambiguata. External appearance of C. predotae much more reminiscent of C. ciscaucasica (Rjabov, 1964) and C. onustaria. Transverse lines of C. ciscaucasica more brownish and forewings wider. C. onustaria with more brownish tinge of ground colour and transverse lines. Male genitalia of C. predotae most similar to those of C. ambiguata, the latter with broader and longer aedeagus, longer cornutus and longer and narrower posterior juxta arms. Female genitalia of C.  ambiguata differing from those of C. predotae in the much larger sclerotization covering 2/3–3/4 of corpus bursae (covering posterior half in C. predotae). Genetic data: Two BINs: BOLD:AAD7406 (n=4 from central, eastern and southern Spain), BOLD:ACE4201 (n=3 from southern Spain). Genetically heterogeneous, in two BINs at a distance of 1.6%. Nearest species: C. ambiguata (1.7%). Remarks: C. predotae was described by Schawerda based on a single male from Albarracin. Wehrli (1953) assumed the species to be a variation of C. ambiguata. Wing coloration and pattern of C. predotae are reminiscent of those of C. onustaria, but morphology of genitalia revealed sister species relationship with C. ambiguata (Rezbanyai-Reser & Expósito-Hermosa 1988).

28. Charissa onustaria (Herrich-Schäffer, 1852) Gnophos onustaria Herrich-Schäffer, 1852: Syst. Bearb. Schmett. Eur. 6 (55): 73, ibidem (1851), pl. LXXXI, figs 496, 491 (non binominal on figures) (Turkey: Amasia). Syntype(s). Gnophos oneraria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 310. Unnecessary replacement name for onustaria Herrich-Schäffer, 1852, cf. Scoble (1999). Gnophos serraria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 304 (France: Corsica). Syntype(s) (ZFMK; examined). Validated at subsp. rank by Scoble (1999), here accepted. In the original description the locus typicus was stated as “Corsica or Andalusia”, Wehrli (1953), after examination of ‘the type’, retained the former as likely. Andalusia is not included in the distribution area of this taxon. ‡ catenulata Rambur, 1866: Cat. Syst. Lepid. Andalusie: pl. XIX, fig. 5 (France, Corsica, see below). Non binominal, not available. At the ZSM two specimens from the Rambur (ex coll. ­Mabille) collection, one labelled as ‘catenulata, Corse’, the other as ‘catenularia Type ­Rambur’, the latter without indication of locality. The specimens revealed to belong to C. onustaria after dissection. Hence it seems that Rambur described a Corsican taxon in his fauna of Andalusia. There is no need to designate a lectotype, as the name is nomenclaturally unavailable. See Remarks.

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Gnophos onustaria var. monotona Schawerda, 1932: Z. öst. Ent.Ver. 17 (2/3): 13, pl. 3, figs 9, 10 (Central Italy; Herzegovina: Mostar; Greece, Peloponnese: Morea). Syntypes (NHMW, not examined). Here classified as junior synonym (of subsp. serraria or of the nominotypical subspecies, lectotype designation required for clarification). Gnophos (Kemtrognophos) onustaria var. eugonia Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 585, pl. XLVIII, fig. f. (Azerbaijan: Talysh mountains, Ljakjar). Syntypes (all at ZFMK, examined), including the male and female ‘type’ of Wehrli are still at the ZFMK. Wehrli’s (1953) statement ‘types and paratypes in ZISP [but still at ZFMK], paratypes in my collection [now ZFMK]’ does not constitute a correct holotype designation, hence all types are syntypes. Since Wehrli, with his term ‘types’, intended a male holotype and a female allotype it is the preferable to designate the male ‘type’ as lectotype, which is herewith done: ‘Talysh mont., fl. Ljakjar, 11viii.1932, M. Rjabov’ [red label] ‘eugonia, Wehrli, Gnophos onustaria, HS, ♂ Type’, ZMFK. Paralectotype 1♀: ‘Daratshitshag, Armen[ia], 7000’, Rjabov, 15.vii.1934’ [red label] ‘eugonia Wehrli, Gnophos, onustaria HS, ♀ Type’. Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy of the nominotypical subspecies, based on the absence of constant and significant differential features correlated with geographical distribution (syn. n.). Unavailable names (infrasubspecific): autumnalis: Dannehl (1933); illineata: Schawerda (1930); kollmorgeni: Schawerda (1932); tangens: Schawerda (1930).

External characters and abdomen: Wingspan males 23–26.5 mm, females 28–32 mm, in the second generation sometimes only 20 mm. Hindwing termen arched between veins. Ground colour of wings dull beige or pale grey. Discal spots ring-shaped, brown or grey. Transverse lines brown or grey, jags of postmedial line darker, antemedial line of forewing often weak or absent, always absent from hindwing. Termen with black dots. Fringes distinctly chequered. Underside whitish grey. Discal spots conspicuous, dark brown to dark grey, usually filled, rarely ring-shaped. Postmedial line inconspicuous. Apical area of forewing conspicuous, dark brown. Terminal area of hindwing darker with dark brown shades or spots. Variation: Variable in contrast and intensity of wing pattern. Second generation usually smaller and paler. Subsp. serraria from Corsica and Apennine peninsula with more contrasting pattern, coloration more brown than grey. Specimens collected in the mountains darker and less strongly marked, therefore reminiscent of continental populations (Wehrli 1953). According to Flamigni et al. (2016) all populations from the Apennine peninsular belong to this subspecies, characterized by more whitish ground colour and well contrasting pattern. Sicilian populations and high mountain specimens attributed to this subspecies, too, though being darker (Flamigni et al. 2016). Male genitalia: Uncus slender, moderately long, blunt at tip. Gnathos well developed. Saccus digitiform projecting, usually narrow. Juxta with triangular external margins, posterior juxta arms thick and close to each other, thick and usually straight. Length of posterior arms about twice length of anterior arms. Valva wider at base, narrowing towards pointed apex, apex moderately and shortly setose, ampulla on costal margin with stout bristle. Aedeagus straight, long (2.0–2.2 mm) and narrow, obliquely truncate at tip, dilated at base. Cornutus very long (1.7–1.9 mm) and thin. Subsp. serraria differing in the shape of the juxta according to Leraut (2009), not confirmed by Flamigni et al. (2016). Female genitalia: Antrum very short, weakly sclerotized, posteriorly truncate. Ductus bursae straight, sclerotized, towards corpus bursae hyaline and narrower. Corpus bursae slen-

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der, long, sclerotized with longitudinal furrows, sclerotization anteriorly narrowing to a long tubu­lar sclerite in a membranous, elongate sac. Length of bursa copulatrix 3.5–4.2 mm. Distribution and abundance: (South) European – West Asiatic. Nominotypical subspecies with disjunct distribution areas in southern Europe in northernmost Spain (Vallhonrat 1989; Dantart 2000; Redondo et al. 2009; requiring confirmation), south-eastern France and north-western Italy, and in the Balkan peninsula. Isolated records in the French Massif Central (Saugues, Ht. Loire, coll. Herbulot/ZSM) and Peloponnese peninsula (coll. B.  Müller and ZSM). On Corsica, Sardinia, Sicily and Apennine peninsula replaced by subsp. serraria Guenée, 1858. – Often local, in Corsica and Apennine peninsula common. – Outside Europe from Turkey and the Levant (Lebanon, Israel, Jordan, cf. Hausmann 1991) to Caucasus, Transcaucasus, northern Iran (Wehrli 1953) and Kopet Dagh mountains in Turkmenistan (Viidalepp 1996).

Phenology: Bivoltine. Flight period depending on altitude. First generation from late March to early June, at higher elevation late April to late June. Second generation starting from mid-July, late July or early August to late September, sometimes until October (Wehrli 1953; Redondo et al. 2009; Flamigni et al. 2016; own data). In Israel and Jordan with first generation from mid-March to early May. Hibernation as larva. Active at night, both sexes readily attracted to light. Biology: Larva oligophagous on Polygonaceae and probably other plants. Recorded on Rumex and Polygonum (Lhomme 1935; Redondo et al. 2009; Flamigni et al. 2016), reared on Rumex acetosa, Polygonum aviculare (Mendes 1913) and Genista (Fritsch et al. 2014). Habitat: Montane. Dry grassland, shrubland, steppes and rocky slopes. Most common from 700 m up to 1600 m in Italy, up to 2700 m in the east of the distribution, in suitable rocky sites sometimes down to 200 m. 111

Similar species (see Text-figs 45–48): External appearance of C.  onustaria reminiscent of C. ciscaucasica (Rjabov, 1964) and C. predotae. C. ciscaucasica differing by slightly darker ground colour and less contrasted transverse lines. C. predotae differing by more greyish ground colour and transverse lines. Male genitalia of C. onustaria most similar to equivalents in C. ciscaucasica and C. remmi but cornutus lacking in C. ciscaucasica whilst being thicker and shorter in C. remmi. Female genitalia of C. onustaria most similar to equivalents in C. ciscaucasica but hyaline appendix bursae absent from the latter. C. graecaria (see no. 34) with pale postmedial fascia of wing underside proximally bordered by a dark, dentate line, terminal area of hindwing underside paler. Genetic data: BIN: BOLD:AAB6741 (n=33 from Italy, Greece, Bulgaria, Russia, Turkey, Israel, Jordan, Lebanon, Iran). Genetically homogeneous, but BIN BOLD:AAB6741 from northern Iran and Turkmenistan (currently identified as C. onustaria) diverging by 4.8%, possibly not conspecific (but without significant differences in genitalia). Nearest species: C. ciscaucasica (barcode-sharing) and C. ambiguata (2.7%). Remarks: Examination of two specimens of ‘catenulata’ from the Rambur collection (­Herbulot in ZSM) revealed details in genitalia similar to equivalents in C. onustaria subsp. serraria. Since one of these catenulata specimens was labelled with ‘Corse’ we retain this to be the correct locus typicus and the name to be synonym of Charissa onustaria subsp. serraria.

29. Charissa remmi (Viidalepp, 1988) Kemtrognophos remmi Viidalepp, 1988: Svyazi entomofaun Severnoi Evropy i Sibiri: 19, figs 1, 2 (Russia, Yakut[skaya] ASSR, Prizhim). Holotype ♂ (IZBE; not examined, paratype ♂ in coll. Skou examined and dissected). Validated at species rank in genus Charissa in Scoble (1999).

External characters and abdomen: Wingspan males 27 mm, females 28–30 mm. Hindwing termen almost round, very shallowly arched between veins. Ground colour of wings slightly shining light brown to grey, somewhat mottled. Discal spots brown to dark grey, oval, usually filled, rarely ring-shaped. Transverse lines brown to dark grey, postmedial line conspicuous, slightly dentate, antemedial line slightly curved, usually present on forewing. Terminal line dark grey, usually disrupted to streaks. Fringes light brown to grey, not chequered. Underside whitish grey with light brown tinge, towards base of forewing often darker grey-brown. Discal spots brown to dark grey, oval, usually filled, rarely ring-shaped. Postmedial line dark brown to grey, weakly developed, on hindwing almost lacking. Terminal streaks present. Variation: Variable in contrast and intensity of wing pattern. Male genitalia: Uncus broad, posteriorly rounded. Gnathos well developed. Saccus triangular at base, digitiform projecting anteriorly. Juxta with long, straight and thick posterior juxta arms, anterior arms absent. Valva wider at base, narrowing towards pointed apex, apex comparatively strongly setose, ampulla on costal margin long, with one or two spines. Aedeagus slightly curved, long (2.2 mm), truncate at tip, dilated at base. Cornutus stout, straight, longitudinally furrowed, long (1.5 mm). Female genitalia: Antrum small, weakly sclerotized. Ductus bursae long, narrowing at centre. Corpus bursae slender, long, sclerotized with longitudinal furrows, near ductus bursae less furrowed, sclerotization anteriorly narrowing and pointed, in a very small membranous sac. Length of bursa copulatrix approx. 4.2 mm. 112

Distribution and abundance: Siberian. In Europe restricted to Polar Urals, eastern prov. Komi (Mironov et al. 2008). – In Europe local, in limited numbers. – Outside Europe across central Russian Siberia and eastern Sayan mountains to republic of Sacha (former Yakutia) (cf. Viidalepp 1996).

Phenology: Univoltine. Flight period from late June to mid-July. Active at night, both sexes readily attracted to light. Biology: Larva unknown. Habitat: Montane. Scarce data from altitudes of 800 m up to 2000 m. Similar species (see Text-figs 45–48): External appearance of C. remmi reminiscent of C. ambi­ guata ophthalmicata. The latter, however, with slightly paler ground colour, medial area wider and forewings narrower. Male genitalia of C. remmi similar to those of C. zeitunaria and C. ciscaucasica (Rjabov, 1964) but C. zeitunaria with shorter juxta arms, narrower and more pointed uncus, C. ciscaucasica without cornutus. Female genitalia of C. remmi reminiscent of those of C. onustaria and C. ciscaucasica. C. onustaria with shorter ductus bursae and larger appendix bursae. Posterior fifth of corpus bursae strongly sclerotized and furrowed in C. ciscaucasica whilst being hyaline in C. remmi. Genetic data: BIN: BOLD:ABU7245 (n=5 from Russia). Genetically homogeneous. Nearest species: C. predotae (5.0%) and C. ambiguata (5.2%).

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30. Charissa zeitunaria (Staudinger, 1901) Gnophos obscuraria (sic!) var. zeitunaria Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. 1: 344 (Turkey: southern Taurus mountains). Lectotype ♂ (MNHU, examined), herewith designated: [yellow label] ‘Zeitun | [18]97 Haradj.’, [red label] ‘Origin.’, ‘v. | Zeitunaria | Stgr.’, ‘1/6’, ‘ex. coll. | STAUDINGER’ ‘‘SE–520 ♂ | genit.prep. S. ­Erlacher, 2011’, MNHU. Paralectotype 1♀ (herewith designated): [red label] ‘Origin.’, ‘3/6.’, ‘ex. coll. | STAUDINGER’ ‘SE–521 ♀ | genit.prep. S. Erlacher, 2011’, MNHU.

External characters and abdomen: Wingspan males 27.5–32 mm, females 28–33.5 mm. Hindwing termen strongly arched between veins. Ground colour of wings slightly shining brown, with orange and yellow tinge. Discal spots brown, oval, ring-shaped. Transverse lines inconspicuous, almost of ground colour, jags of postmedial line darker. Terminal line disrupted to dark grey streaks. Fringes slightly chequered. Underside whitish with light brown tinge, in terminal area darker brown. Postmedial line replaced by a pale ochreous fascia. Discal spots inconspicuous, small or absent. Variation: Variable in intensity of brown suffusion and distinctness of pattern.

Male genitalia: Uncus at base moderately wide, posteriorly with narrow, digitiform projection, curved ventrad. Gnathos developed. Saccus short, broad, anteriorly rounded. Juxta with short and thick posterior juxta arms, narrowing towards tip, anterior arms very small or absent. Valva very broad at base, narrowing towards pointed apex, apex weakly setose. Ampulla appressed to or fused with costal margin of valva, with one short spine. Aedeagus straight, very long (2.4–2.5 mm), slightly dilated at base. Cornutus stout, straight, longitudinally furrowed, very long (2.1–2.4 mm). Female genitalia: Papillae anales sclerotized, sub-rectangular, with stout setae. Apophyses anteriores and posteriores very long, 1.6–1.8 mm and 2.4–2.7 mm respectively. Antrum calyx-shaped, lateral edges sclerotized, posteriorly curved outside, at centre a sclerotized pocket. Ductus bursae broad, straight, sclerotized. Corpus bursae narrow, long, sclerotized with longitudinal furrows, anteriorly narrowing and pointed, here with numerous spinules. At anterior end a large membranous sac, with lateral, oval, membranous, sac-shaped appendix. Length of bursa copulatrix approx. 3.9–4.4 mm. Distribution and abundance: East Mediterranean – Anatolian. In Europe restricted to Crete (ZSM; Hausmann et al. 2004; 2011a). – In Europe local and rare, in Turkey sometimes collected in numbers. – Outside Europe from western to eastern Turkey, Armenia and Azerbaijan (Wehrli 1953), probably also in northernmost Iran. Phenology: Univoltine. Flight period from late July to mid-September, sometimes until mid-October (Wehrli 1953; own data). Active at night, both sexes readily attracted to light. Biology: Larva unknown. Habitat: Xeromontane. Scarce data from dry and rocky slopes. Vertical distribution from 1000 m up to 2500 m. Similar species: External appearance of C. zeitunaria reminiscent of the two Non-­European species C.  argillata (Brandt, 1938) from eastern Turkey and Iran and C.  mardinaria (Staudinger, 1901) from south-eastern Turkey and the Levant. C. argillata with paler ground colour and more brownish transverse lines, C.  mardinaria with darker brownish coloration. In male genitalia valva of both C. argillata und C. mardinaria bearing tapered ventral 114

projections at the tip of the sacculus (lacking in C. zeitunaria). Female genitalia of C. mardinaria with strongly curved corpus bursae and without the subapical, lateral membranous appendix bursae. Female genitalia of C. argillata with much broader corpus bursae lacking the sclerotization in the posterior half. C. graecaria (see no. 34) with pale postmedial fascia of wing underside proximally bordered by a dark, dentate line, terminal area of hindwing underside paler. Genetic data: BIN: BOLD:AAE2751 (n=5 from Turkey). Genetically heterogeneous, maximum variation 1.9%. Nearest species: C. argillata from Turkey (5.5%) and C. mardinaria (5.7%).

Subgenus Neognophina Wehrli, 1946 Neognophina Wehrli, 1946, Revue fr. Lépidopt. 10: 241. Type species: Gnophos intermedia Wehrli, 1917. Wehrli describes “Neognophina subg. n. (groupe intermedia)” for the second time, later on (Wehrli 1951: 24), which must be considered an objective synonym. Rhopalognophos Wehrli, 1951, Lambillionea 51: 24. Type species: Gnophos glaucinaria H ­ übner, 1799. Herewith downgraded to synonymy of subgenus Neognophina Wehrli, 1946, based on COI barcode analysis, suggesting closer relationships with the other two European Neo­ gnophina species syn.n.

Diversity and distribution: Three species in the western Palaearctic region. External characters: Underside of wings with diffuse but rather wide postmedial line, pale narrow fascia, terminal area brown and its inner margin rather diffuse. Male genitalia: Valva margins parallel or tapering, apex narrow. Base of valva with distinct and sclerotized, hook-shaped spine, ventrally with small sclerotized ridge (absent in C. glaucinaria). Posterior arm of juxta long, apical half a hollow tube. Aedeagus with ­serrate, 115

lateral extensions, without cornutus, or without serrate extensions and one very long cornutus (C. glaucinaria). Female genitalia: Lamella postvaginalis weakly sclerotized medially (sclerotization absent in C. glaucinaria). Ductus bursae wide (narrow in C. glaucinaria). Corpus bursae covered with parallel, spinose grooves. Biology: Larva of C. supinaria has been found on Asplenium ceterach (Flamigni et al. 2016). In captivity it has been reared on Lotus corniculatus, Clematis, Sedum and Campanula, among others (Flamigni et al. 2016). Immature stages: Larva of C. supinaria grey to olive-green, with whitish longitudinal stripe laterally, a pair of divergent lines dorsally (most prominent on segments 5–8) and short dorsal projections on 11th segment (Lepiforum 2018). Pupal cremaster of C. supinaria [illustrated as C. intermedia] with four pairs of curved hooks, apical pair most prominent (Patočka & Turčani 2005). Remarks: Genitalia morphology of C. glaucinaria, particularly female genitalia, is similar to species in subgenus Kemtrognophos. DNA barcodes do not support this relationship, and this is the reason why the following three species are combined in subgenus Neognophina.

31. Charissa glaucinaria (Hübner, 1799) Geometra glaucinaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 28, fig. 150 (Europe). Syntype(s). Gnophos falconaria Freyer, 1842: Neuere Beitr. Schmettkde 4 (63): 153, pl. 377, fig. 3 (Alps). Syntype(s). Validated at subspecific rank (Scoble 1999), but here retained as infrasubspecific form (Leraut 2009; Flamigni et al. 2016). [Gnophos] sartaria Herrich-Schäffer, 1848: Syst. Bearb. Schmett. Eur. 3 (13): 75; ibidem (1845), pl. 12, figs 66, 67 (Morea, Dalmatia, Turkey). Syntype(s). Non binominal. Junior synonym of Gnophos falconaria (Scoble 1999). Gnophos glaucinata Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 299. Emendation of glaucinaria Hübner. Gnophos panessacaria Trimoulet, 1858: Actes Soc. Linn. Bord. 22: 62 (France: St-Médard). Syntype(s). Junior synonym of Geometra glaucinaria. Gnophos glaucinaria ab. plumbearia Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb.: 167 (Europe). Syntype(s). Validated at subspecific rank (Leraut 2009, misspelled as plumbaria). Tentatively validated here at subspecies rank, requiring more research. Gnophos glaucinaria etruscaria Staudinger, 1892: Dt. ent. Z. Iris 5: 188 (Italy: Apennines, Vallombrosa). Holotype ♂. Validated at subspecific rank under Charissa crenulata (Scoble 1999), but validated at subspecific rank under Charissa glaucinaria by Flamigni et al. (2016). Here, tentatively retained as valid at subspecies rank, requiring more research. Gnophos operaria f. anastomosaria Schwingenschuss, 1918: Verh. zool.-bot. Ges. Wien 68 (6– 8): 154 (Austria: Beler Kalkalpen, Hochschwab). Syntypes 3♂. Junior synonym of Geometra glaucinaria. Gnophos glaucinaria f. intermediaria Turati, 1919: Atti Soc. ital. Sci. nat. 58 (2): 178 (Italy: Apennines, Modenese, 1000 m). Syntype(s). Validated at subspecific rank (Scoble 1999), considered a junior synonym of Gnophos glaucinaria etruscaria by Flamigni et al. (2016). Gnophos glaucinaria var. jurassica Osthelder, 1921: Mitt. münch. ent. Ges. 11 (6–8): 31 (Germany: Bavaria, Kelheim). Holotype ♂ (ZSM). Junior synonym of Geometra glaucinaria. Gnophos glaucinaria juravolans Wehrli, 1924: Mitt. Schweiz. ent. Ges. 13 (7): 344 (Switzerland: Jura). Syntype(s) (ZFMK). Validated at subspecific rank (Scoble 1999). Here, tentatively validated at subspecies rank, requiring more research.

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Gnophos glaucinaria f. perstrigata Wehrli, 1924: Dt. ent. Z. Iris 38: 85, pl. 1, fig. 3 (France: Alpes Maritimes). Syntype(s) (ZFMK). Junior synonym of Geometra glaucinaria. Gnophos fischeri Wehrli, 1934: Int. ent. Z. 27 (47): 535, figs 17, 18 (France: Vosges). Syntypes 18♂4♀ (ZFMK). Validated at subspecific rank by Scoble (1999), considered a junior synonym of Gnophos glaucinaria plumbearia by Leraut (2009). Gnophos reisseri Schawerda, 1938: Ent. Rdsch. 55 (47): 555 (Spain: Castilia, Sierra de Gredos, Pozas de Gredos, 1900 m). Holotype. Junior synonym of Geometra glaucinaria. Gnophos glaucinaria salvatorensis Schwingenschuss, 1942: Z. wien Ent. Ver. 27 (10): 247 (Italy: Sicily, Monte San Salvatore). Syntypes 2♂1♀. Validated at subspecific rank (Scoble 1999; Flamigni et al. 2016), here tentatively retained as valid at subspecies rank, requiring more research. Gnophos glaucinaria supinodes Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 599 (Yugoslavia: Zara, Dalmatia, Bosnia). Syntypes ♂♀ (ZFMK). Validated at subspecific rank (Scoble 1999), here tentatively retained as valid at subspecies rank, requiring more research. Gnophos glaucinaria turatii Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 599. Unnecessary replacement name for Gnophos glaucinaria f. intermediaria. Gnophos glaucinaria peruni Varga, 1975: Acta Univ. ebrecen. (2) (Biol.) 12: 84, fig. 12, text-fig. 4 (Bulgaria: Pirin Mountains, Vichren-Hütte, 2060 m). Holotype ♂. Validated at subspecific rank (Scoble 1999), here tentatively retained as valid at subspecies rank, requiring more research. Unavailable names (misspelling): plumbaria: Leraut (2009: 231).

External characters and abdomen: Wingspan 26–32 mm. Wings grey to yellowish, often with olive-green tinge, irrorated densely with dark scales, creating contrasting appearance. Forewing with denticulate, blackish transverse lines, widened at costa. Antemedial line often as row of blackish brown dots or almost absent, whereas postmedial line distinct, its outer margin bordered with paler area. Wavy line weakly present near forewing costa. Hindwing with well-developed, denticulate postmedial line. Terminal area of wings often darker. Discal spots large, pale-centred. Terminal line as row of blackish brown dots. Fringes chequered. Hindwing termen undulating. Underside of wings brown, darkest in terminal area. Post­ medial line narrow, bordered with pale and narrow fascia. Terminal area brown and its inner margin rather distinct, with pale blotch on each wing. Discal spots weak on underside. Overall impression of underside contrasting. Frons, labial palpi, collar, thorax and abdomen concolorous with wings. Male antennae ciliate-fasciculate, with short cilia. Female antennae filiform. Hindlegs in both sexes with 2+2 spurs, male hindleg swollen. Abdominal sternites unmodified. Variation: Wing pattern rather uniform, but wing colour varies from whitish grey to yellowish grey, to ash or lead grey. Area between antemedial and postmedial lines, and terminal area often slightly darker than rest of wing. Postmedial line often the most distinct line. Terminal area often irrorated with dark grey, silvery, and olive-green scales. Underside of wings often contrasting, particularly inner margin of terminal area well-defined. Fringes often white, whitish, or yellowish grey, but may be grey, strongly or weakly chequered. Numerous subspecies have been described based on external characters, apparently wing coloration of adults is adapted to local substrate. Validity of subspecies is debatable, detailed descriptions are available elsewhere (e.g Varga 1975a; Leraut 2009; Flamigni et al. 2016). Male genitalia: Uncus wide, triangular, weakly setose. Gnathos arms narrow, fused apically. Tegumen wide. Valva margins parallel, valva narrow, long, weakly setose, apex concave between acute ventral tip and round apex of costa. Costa wide, sclerotized, base upturned, apex round. Anterior arms of juxta long and fused basally, posterior arms of juxta short. 117

Saccus elongated. Aedeagus long, slightly curved, caecum short. Cornutus massive, almost as long as aedeagus, with two fused, parallel spines. Vesica opens at 135 degree angle, narrow simple sac, cornutus attached to its tip (vesica not illustrated here, but it is shown in Malkiewicz 2012). Sclerotized tip of valva may be absent (see Flamigni et al. 2016), apex of uncus varies from round to acute, apices of ventral arm of valva may be acute or rounded, and shape of saccus varies from weakly elongated to distinctly elongated (variation of saccus is illustrated in Flamigni et al. 2016). Female genitalia: Papillae anales large, setose. Apophyses posteriores slightly longer than apophyses anteriores. Lamella antevaginalis and lamella postvaginalis membranous. Ductus bursae wide, short, most sclerotized laterally. Corpus bursae covered with parallel sclerotized, weakly spinose grooves. Sclerotization expanded, tongue-shaped at anterior end. Anterior end partly membranous. Shape of tongue-shaped sclerotization varies from triangular to elongated to pointed (variation is illustrated in Flamigni et al. 2016). Distribution and abundance: European. In the Alps and through most of the Apennines in I­taly, in Central Massif in France and in the Pyrenees with isolated occurrences in north-­ western Spain, southern Italy, southern Bulgaria, Greece and through southern central ­Europe from western Ukraine and southern Poland to north-western France. A single record by Ribbe (1912) from Sierra Nevada in southern Spain, probably due to misidentification. We do not differentiate tentative subspecies in the distribution map. – Relatively common, mainly found in limited numbers, but now and then more abundant. – Outside Europe recorded from Turkey (Koçak & Kemal 2009), but we have not been able to trace specimens from Turkey.

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Phenology: Uni- or bivoltine. Recorded from late April to early May and from late May to mid-October. The long and uninterrupted flight period likely depends on altitude, possibly also on latitude. In central Germany from mid-July to early September (Erlacher 2019). In Slovenia in one long generation from mid-June to late September (S. Gomboc pers. comm.). Larva from May to early July and from late July, hibernating to early May (Ziegler 2019). Active at night, both sexes readily attracted to light. Also found while resting during day time, but hardly detected due to its camouflaged coloration. Adults often under stones or in damp niches (Bergmann 1955). Biology: Polyphagous. Recorded on Carex, Polygonum, Lychnis, Silene, Sedum, Hippocrepis, Thymus, Teucrium, Linaria, Plantago, Scabiosa, Campanula, Artemisia, Taraxacum, Genista and Vaccinium myrtillus (Flamigni et al. 2016). Habitat: Montane. In a rather large variety of open and semi-open habitat types. In coll. Skou from 360 m up to 2575 m. According to Flamigni et al. (2016) also found close to sea-level. According to Ziegler (2019) from 500 m up to 2500 m and according to S. ­Gomboc (pers. comm.) from 700 m up to 2100 m. In the limestone Carpathians in western and northern Slovakia from 500 m up to 1500 m (Z. Las˘tu˚ vka pers. comm.). Similar species: Externally, Charissa glaucinaria, C. pfeifferi and C. supinaria are very similar, see diagnostic characters in Text-figs 52–55. C. pfeifferi and C. supinaria subsp. intermedia with postmedial line more distinct on wing underside (postmedial line less developed in C. glaucinaria). Both male and female genitalia of these taxa allow reliable identification. Also C. crenulata, C. pullata and C. italohelveticus are similar, see Text-figs 49–51. Genetic data: BIN: BOLD:AAC1853 (n=41 from Germany, Austria, Spain, France, Italy). Genetically heterogeneous, maximum variation 2.9%, the westernmost specimens from western Germany, western Austria and northern Spain (n=4) BIN-sharing with the main cluster but diverging by a minimum pairwise distance of 0.9%. Nearest species: C. pfeifferi (4.6%) and C. intermedia (4.8%).

32. Charissa pfeifferi (Wehrli, 1926) Gnophos pfeifferi Wehrli, 1926, Mitt. münch. ent. Ges. 16 (8–12): 95 (Turkey: Anatolia, Egerdir, Akşhehir). Syntypes 5, including ♂ and ♀ (ZFMK).

External characters and abdomen: Wingspan 26–31 mm. General appearance as in C. glaucinaria, but with following diagnostic differences (see Text-figs 52–53): irrorations paler, antemedial and postmedial lines paler, therefore overall appearance more uniform and less contrasting. Underside of wings contrasting between dark brown and beige. Postmedial line well-developed. Pale fascia wide, weakly curved outwards on forewing tornus. Terminal area with three well-defined pale blotches: on forewing apex, middle of forewing and middle of hindwing. Variation: Wing colour varies from whitish grey to yellowish sand-colour to ash- or leadgrey. Often terminal area darkest, with ash or silvery scales, with weak paler blotches barely visible on upperside. Pale fascia between postmedial line and terminal area only slightly paler than rest of wing. Male genitalia: Uncus wide, triangular, weakly setose. Gnathos arms narrow, fused apically, weakly elongated. Tegumen wide. Base of valva wide, ventral margin rounded. Apical half of valva narrower, margins tapering, apex narrow. Costa of valva weakly serrate at base, 119

with distinct and sclerotized hook-shaped spine. Ventrad of spine small sclerotized ridge. Anterior arm of juxta long, apical half a hollow tube. Aedeagus stout, with short caecum, apex with serrate, triangular extensions laterally. Cornutus absent. Female genitalia: Papillae anales weakly elongated, setose. Apophyses posteriores about twice as long as apophyses anteriores. Lamella postvaginalis with small, round sclerotization medially. Lamella antevaginalis a narrow, weakly sclerotized ring. Ductus bursae short, wide. Corpus bursae rather narrow, central part covered with parallel, spinose grooves. Anterior end of corpus bursae membranous. Distribution and abundance: South-east European-Anatolian. In Europe only found in Bulgaria, in mainland Greece and in Samos. – Local, usually found in limited or very limited numbers. – Outside Europe in western and southern Turkey.

Text-figs 49–55. Diagnostic characters (indicated) of selected Charissa species. Occasionally, external characters do not allow reliable identification, in which case the genitalia and/or DNA barcodes need to be examined. Text-fig. 49. C. crenulata (Staudinger, 1871), female, Spain: Teruel, Moscardon, e. o. 13.viii.1998 (coll. Gelbrecht). Text-fig. 50. C. italohelveticus (Rezbanyai-Reser, 1986) female, Switzerland: Valais, Raron, e. o. 17.vii.1987 (ZSM). Text-fig. 51. C. pullata (Denis & Schiffermüller, 1775), male, Austria: North Tyrol, Fließ am Inn, 25.vi.2008 (coll. Skou). Textfig. 52. C. glaucinaria (Hübner, 1799), male, France: Hautes Alpes, La Grave, Villar-d’Arêne, 2000 m, 26.vii.2000 (coll. Skou). Text-fig. 53. C. pfeifferi (Wehrli, 1926), male, Turkey: Antalya, Bei Dağları, Saklikent, 21.v.1999, (coll. Gelbrecht). Text-fig. 54. C. supinaria (Mann, 1854), male. Croatia (ZFMK), syntype. Text-fig. 55. C. supinaria subsp. intermedia (Wehrli, 1917), male, Austria: North Tyrol, Zirl, 26.vi.1961 (ZSM). %

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Phenology: Bivoltine. In Samos from early April to late May and from early September to mid-October (D. Fritsch pers. comm.). In mainland Greece the relatively few records are from early May, early and late June, late July, early and mid-September, and early October. Adults active at night, readily attracted to light. Biology: Unknown in nature. Reared from eggs on Genista (Fritsch et al. 2014), and Genista tinctoria (D. Stadie pers. comm.). Habitat: Montane, xerothermophilous. In mainland Greece in open and semi-open habitats, uncultivated areas with rocks from 250 m up to 1700 m, in Greece up to 1900 m. In Samos in similar places, from 200 m up to 900 m (D. Fritsch, pers. comm.). In the Taurus mountains (Turkey) found in rocky gorges and steep escarpments on limestone (D. Stadie pers. comm.). Similar species: Externally, Charissa glaucinaria, C. pfeifferi and C. supinaria are very similar, see diagnostic characters in Text-figs 52–55. C. supinaria has narrow serrate extensions in aedeagus (C. pfeifferi has triangular serrate extensions in aedeagus). C. supinaria has lamella postvaginalis with narrow, ring-like medial sclerotization and corpus bursae is wide (C. pfeifferi has lamella postvaginalis with plate-like medial sclerotization and corpus bursae is narrow). Genetic data: BIN: BOLD:AAD1467 (n=14 from Greece, Turkey, Lebanon). Genetically homogeneous, maximum intraspecific divergence 0.6%. Nearest species: C. supinaria (2.1%).

33. Charissa supinaria (Mann, 1854) Boarmia supinaria Mann, 1854: Verh. zool.-bot. Ver. Wien 4 (Abh.): 566 (Croatia: Rijeka [­Fiume], Istria). Syntype(s) (ZFMK, examined). Gnophos glaucinata supinata Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insected (Spec. gén. Lépid.) 9: 299. Emendation of Boarmia supinaria Mann. Gnophos glaucinaria intermedia Wehrli, 1917: Verh. naturf. Ges. Basel 28: 253 (Switzerland: Jura). Holotype (ZFMK). Validated at species rank (as Charissa intermedia) (Scoble 1999), or at subspecific rank (as Charissa supinaria intermedia) (Flamigni et al. 2016). Here, tentatively validated at subspecies rank, requiring more research, see Distribution and Genetic data. Gnophos supinaria var. hypophana Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 596 (Macedonia: Ohrid?). Syntypes 4♂ (ZFMK). Junior synonym of Gnophos glaucinaria intermedia Wehrli, 1917. Gnophos intermedia benestriata Kovács, 1954: Annls hist.-nat. Mus. natn. hung. (N. S.) 5: 313 (Hungary: Branyiszko). Holotype ♂ (TMB). Validated at subspecific rank (Scoble 1999), downgraded here to synonymy with Boarmia supinaria Mann, 1854 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos intermedia budensis Kovács, 1954: Annls hist.-nat. Mus. natn. hung. (N. S.) 5: 314 (Hungary: near Buda, Pilisvörösvar). Holotype ♂ (TMB). Validated at subspecific rank (­Scoble 1999), downgraded here to synonymy with Boarmia supinaria Mann, 1854 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos intermedia dioszeghyi Kovács, 1954: Annls hist.-nat. Mus. natn. hung. (N. S.) 5: 313 (Hungary: Retyezát, 1300 m). Holotype ♂ (TMB). Validated at subspecific rank (Scoble 1999), downgraded here to synonymy with Boarmia supinaria Mann, 1854 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation. Gnophos intermedia f. francothuringiaca Bergmann, 1955: Gross-Schm. Mitteldeutschlands 5 (2): 923, 933, 934, 945, pl. 269, figs C1–6, pl. 270, figs A1, 2, B1, 2 (Germany: Thuringia, Frankenwald, Höllental near Hirschsprung, Kobersfels near Burgk, Muckenberg near Lobenstein). Syntypes ♂♀ (MNHU, examined). Validated at subspecific rank (Scoble 1999), herewith downgraded to synonymy based on the lack of significant differential features in genitalia and DNA barcodes (syn. n.).

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Gnophos interdemia gulsensis Wolfsberger, 1959: Z. wien. ent. Ges. 44 (3): 40, pl. 1, figs 1–4 (Austria: Steiermark, Gulsenberg/Murtal, 600–900 m). Holotype ♂. Validated at subspecific rank (Scoble 1999), downgraded here to synonymy with Boarmia supinaria Mann, 1854 (syn. n.), based on the absence of constant differential features, falling within intraspecific variation.

External characters and abdomen: Wingspan 26–29 mm. General appearance as in C. glau­ cinaria, but with following diagnostic differences (see Text-figs 52, 54–55): Wings rather uniform pale grey-brown, antemedial and postmedial lines weak, therefore overall appearance rather homogeneous. Underside of wings with diffused markings. Pale fascia on forewing tornus weakly curved inwards. Terminal area with small and diffuse pale blotches. Variation: Wing colour is variable, ranging from rather uniform pale grey-brown to pale yellowish. Irrorations may be scarce and grey, or numerous and dark brown to olive-green to silvery to blackish. Wing coloration of adults is adapted to different substrates, resulting in the description of numerous subspecies, and thus in redundant taxonomy. Validity of the isolated population in Thuringia (‘francothuringiaca’) controversial, here downgraded to synonymy despite the peculiar yellowish green tinge of the ground colour. Subsp. intermedia is more contrasting, particularly on underside) see Colour plates figs. 33c-e and Text-fig. 55): postmedial line is distinct, often weakly angled on forewing and wide fascia is pale. Genitalia are without diagnostic characters from the nominotypical subspecies, and characters given in the literature (Wehrli 1922a; Leraut 2009) are not constant (Flamigni et al. 2016). Not all specimens can be identified based on wing pattern and colour, transitional specimens are discussed and illustrated in Flamigni et al. (2016). Although the taxon intermedia is genetically diagnosable (see Genetic data), its status needs more research, particularly focusing on specimens from different altitudes from north-eastern Italy, Slovenia, Croatia and southernmost Austria. Male genitalia: Generally as in C. pfeifferi, but in C. supinaria base of valva narrower, ­ edeagus apex with lateral rod-shaped extensions, vesica opens at 90 degree angle, swollen a at base, without cornutus (vesica not illustrated here, but shown in Malkiewicz 2012). Female genitalia: Generally as in C. pfeifferi, but in C. supinaria lamella postvaginalis narrow, without central plate and with wide corpus bursae. Distribution and abundance: Central- and south-east European. Nominotypical subspecies found from Adriatic coast to Greece and Bulgaria, but with disjunct distribution; recently recorded as new for Albania (S. Beshkov pers. comm.). Subspecies intermedia with disjunct distribution in the Alps and with scattered occurrences in central and eastern Europe; recently recorded as new for Hungary (E. Friedrich pers. comm.). – Usually found in limited numbers. – Unknown from outside Europe. Phenology: Uni- or bivoltine. From late April to late May and from mid-August to mid-­ October. In south-western Germany in one generation from mid-May to early July (subsp. intermedia), however, total flight period not lasting more than four weeks in each year (Ebert et al. 2003). In central Germany from late April to late May (Erlacher 2019), under rearing conditions regularly a second generation in autumn. Two generations in the Czech and Slovak republics, the first in first half of May and the second in August (Z. Las˘tu˚ vka pers. comm.). In Slovenia and Croatia in two clearly separated generations from late April to early June and from early August to late September (subsp. supinaria) (S. Gomboc pers. comm.). Adults active at night, attracted to light. Resting in shady places of sun-exposed rocks during daytime (central Germany: S. Erlacher own observation). 123

Biology: Probably polyphagous. Found on Sedum rupestre (Bergmann 1955). Recorded on Asplenium ceterach and in Italy reared on Lotus corniculatus (Flamigni et al. 2016). In south-western Germany reared on Taraxacum (Ebert et al. 2003), in central Germany reared on Plantago, Lactuca, Taraxacum, Campanula, Lonicera, Clematis, and Sedum album (Bergmann 1955). Habitat: Dry slopes and rocky places (Ebert et al. 2003). In Croatia on limestone rocks near the coast. From sea-level (subsp. supinaria) up to 1300 m, in south-western Germany from 500 m up to 1000 m, in central Germany around 400 m (Erlacher 2019). In the Czech and Slovak republics on relatively dry and warm but also on more shaded and cooler rocky places from 300 m up to 600 m (Z. Las˘tu˚ vka pers. comm.). Similar species: Externally, Charissa glaucinaria, C. pfeifferi and C. supinaria are very similar, see diagnostic characters in Text-figs 52–55. C. supinaria has narrow serrate extensions in aedeagus (C. pfeifferi has triangular serrate extensions in aedeagus). C. supinaria has lamella postvaginalis with narrow, ring-like medial sclerotization and corpus bursae is wide (C. pfeifferi has lamella postvaginalis with plate-like medial sclerotization and corpus bursae is narrow). Genetic data: BIN: BOLD:ABY4044 (n=30). Diverging into two main clusters at a constant minimum distance of 0.6%: The first (n=9) from north-easternmost Italy, Croatia, Bulgaria referring to the nominotypical subspecies, the second (n=21) from Germany, Austria, Roma­ nia referring to subsp. intermedia. Thus, genetic data support downgrading of intermedia to subspecies rank under C. supinaria. Two barcoded specimens from north-easternmost Italy (Friuli) diverging by 0.8% from nominotypical subspecies. Two barcoded specimens from south-eastern Austria diverging by 1.4% from the main cluster of subsp. intermedia. The isolated population in Thuringia showing a constant divergence of 1bp (0.15%; n=4) from

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the other populations of subsp. intermedia. Nearest species: C. pfeifferi (2.1% from both nominotypical subspecies and subsp. intermedia). Remarks: More research is needed to clarify the distribution border between nominotypical subsp. supinaria and subsp. intermedia in north-eastern Italy, Slovenia and western Croatia, where these taxa occur very close to each other.

Subgenus Trilobignophos Wehrli, 1951 Trilobignophos Wehrli, 1951: Lambillionea (51): 23. Type species: Gnophos pollinaria Christoph, 1887.

Diversity and distribution: The subgenus Trilobignophos currently contains four species in the Palaearctic, mainly in the Middle East. The moths are medium-sized, with a more or less blackish white contrasted wing underside. Due to the external similarity, the identification of the adults is very difficult. A taxonomic revision of this subgenus is urgently needed. Male genitalia: Uncus elongated and pointed. Gnathos weakly developed. Juxta divided almost down to the base, juxta arms either forked and thin or v-shaped and slightly dilated. Valva relatively narrow. Costa of valva well sclerotized, approximately in the middle usually with one, sometimes with two spines on a relatively long process. Aedeagus with four to six cornuti. Female genitalia: Corpus bursae with strongly sclerotized, spirally spined, longitudinal folding of different extensions, anterior part sometimes bearing a round and spiny plate (lamina dentata). Antrum well sclerotized.

34. Charissa graecaria (Staudinger, 1871) Gnophos ambiguata [“v.”] graecaria Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 37: 167 (Greece). Syntypes 2♂ (MNHU, examined). In the original description erroneously assigned to Gnophos ambiguata, in Scoble (1999) still as subspecies of Charissa ambiguata, here recognized as senior synonym of C. certhiatus. Gnophos certhiatus Rebel & Zerny, 1931: Denkschr. Akad. Wiss. Wien, Math.-naturw. Klasse 103: 115, figs 1, 2, 5 (Albania: Pashtrik). Syntypes 2♂2♀ (NHMW, examined). Here downgraded to synonymy of C. graecaria after examination of both syntype series (syn. n.). Gnophos certhiatus minorasiaticus Wehrli, 1936: L’amateur papillons 8 (10): 154 (Turkey, Anatolia: Akşhehir, Sultan Dagh, 1700–2200 m.). Syntypes 1♂1♀ (ZFMK, examined). Here downgraded to synonymy of C. graecaria, after examination of both syntype series, and based on clinal character transitions (syn. n.).

External characters and abdomen: Wingspan 28–35 mm. Wings relatively broad. Ground colour of wing upperside creamy white, whitish grey or brown. Transverse lines clearly visible, serrated, sometimes broken into dots. Terminal line broken into small black dots at veins endings. Discal spots variable, usually unfilled. Fringes concolorous with forewings, not chequered. Hindwing margin undulate. Underside of forewing with inconspicuous, proximally indented, white postmedial line, being weaker on hindwing. Frons, collar, thorax and abdomen largely concolorous with wings. Antennae filiform in both sexes. Variation: Ground colour of adults very variable, depending on colour of the (limestone) substrates, ranging from white to light brown.

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Male genitalia: Uncus elongated and pointed. Gnathos weakly developed. Juxta divided almost down to base, v-shaped, juxta arms slim, distal half slightly broader than proximal half, at tip again tapered but rounded. Valva relatively narrow. Costa of valva well sclerotized, approximately in the middle usually with one, sometimes with two spines on a relatively long process. Aedeagus distally slightly tapering, with about four to five cornuti of about one-third length of aedeagus. Female genitalia: Antrum well sclerotized. Corpus bursae with strongly sclerotized, spinose, longitudinal folding in the posterior part down to about half of corpus bursae, anteriorly bordered straight (circular). Anterior half of corpus bursae with a round and spinose plate (lamina dentata). Distribution and abundance: (South-east) European – West Asiatic. In Europe widely distributed in the Balkan peninsula, from Slovenia and Croatia (Lasan 2000) across Bosnia and Herzegovina, Albania and Montenegro (Rebel & Zerny 1931), to Greece, including Peloponnese, and Bulgaria. – Outside Europe in central (Wehrli 1953) southern (Konya Province: Sultan Dagh) and northern Turkey (Kastamonu Province: Yaraligöz-Pass).

Phenology: Univoltine. From late June to mid-September. Hibernating as larva (rearing observation; J. Gelbrecht pers. comm.). Adults nocturnal, attracted to light, rarely flushed up from vegetation. Biology: Larval foodplants unknown. Under rearing conditions polyphagous on low plants (J. Gelbrecht pers. comm.). Habitat: Montane. In Greece in alpine meadows, mainly on limestone. From 550 m in Pirin mountains (Müller 1986) up to 1700 m on Mount Parnassus (coll. E. Friedrich).

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Similar species: C. graecaria might be confused with C. pullata, C. italohelveticus, and C. ambi­ guata. However, C. graecaria can be differentiated from C. pullata and C. ­italohelveticus by the underside of the wings which is much less contrasting, the underside of the hindwings is brighter than that of the forewings. In addition, the terminal spots on the wing upperside are more pronounced in C. graecaria than in the other two species. However, this also applies to C. ambiguata. In the latter, however, the postmedial line is curved inwards before reaching the costa. Identification of whitish specimens from calcareous substrates is difficult based on external characters and requires dissection of genitalia. For differences from C. zeitunaria see no. 30. Genetic data: BIN: BOLD:AAF2916 (n=7 from Macedonia and Greece). Genetically slightly heterogeneous (maximum variation 1.6%). Nearest species: C. zejae (2.5–3.0%) from Dagestan and Iran (Elburs). Another cluster at a distance of 4.9% from eastern Turkey was clearly identified as C. zejae after dissection of male and female genitalia, requiring further study. Remarks: C. graecaria was originally described as a variation of ‘Gnophos’ ambiguata. Sixty years later, Rebel & Zerny (1931) described Gnophos certhiatus as a new species. However, examination of the type material of certhiatus clearly showed its conspecifity with C. graecaria (= Gnophos certhiatus Rebel & Zerny, 1931, syn. n.). As most congeners, C. graecaria tends to develop substrate-adapted forms recurring at distant locations. This also applies to the taxon minorasiaticus, which is therefore synonymized with the nominotypical subspecies of C. graecaria, here: Gnophos certhiatus minorasiaticus Wehrli, 1936 (syn. n.).

Psodos Treitschke, 1825 Psodos Treitschke, 1825: in Ochsenheimer, Die Schmetterlinge von Europa, 5: 434. Type species: Phalaena equestrata Fabricius, 1777. Torula Boisduval, 1840: Genera et index methodicus Europæorum Lepidopterorum: 230. Type species: Phalaena equestrata Fabricius, 1777. Synonym (Scoble 1999). Glacies Millière, [1874] 1869: Iconogr. Descr. Chenilles Lépid. inédits 3: 429. Type species: Psodos alticolaria Mann, 1853. Validated at genus rank in Scoble (1999), Huemer et al. (2009), Malkiewicz (2012), Huemer (2013) and Flamigni et al. (2016), downgraded here to synonymy with Psodos Treitschke, 1825 (syn. n.) due to phylogenetic reasons (see remarks). Triglavia Povolny & Moucha, 1956: Sbornik Ent. Oddeleni Narodniho Mus. Praze 30: 164. Type species: Psodos spitzi Rebel, 1905. Listed as synonym of Glacies in Scoble (1999). Trepidina Povolny & Moucha, 1956: Sbornik Ent. Oddeleni Narodniho Mus. Praze 30: 165. Type species: Phalaena canaliculata Hochenwarth, 1785. Listed as synonym of Glacies in Scoble (1999). Alpina Povolny & Moucha, 1956: Sbornik Ent. Oddeleni Narodniho Mus. Praze, 30: 168. Type species: Phalaena coracina Esper, 1805. Listed as synonym of Glacies in Scoble (1999). Unavailable names: Psothus Agassiz, 1847; Psothos Spuler, 1910; Psolos Berthet, 1938 (see Fletcher 1979).

Diversity and distribution: 15 species, most of them restricted to European high mountains. Only three species, Psodos sajana Wehrli, 1919, Psodos coracina (Esper, 1805), and Psodos canaliculata (Hochenwarth, 1785), occurring in central and northern Asian high mountains and one subspecies (P. sajana daisetsuzana Matsumura, 1927) on the Japanese island Hokkaido. External characters and abdomen: Ground colour of body and wings dark brown to black. Wings with distinct dark markings; upperside with ante- and postmedial, subterminal and terminal lines. Antemedial line not continued on the hindwings; ante- and postmedial lines

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serrated or rounded; terminal line interrupted or almost continuous. Underside of wings weakly marked, with paler terminal area and white to silvery lines (except in P. quadrifaria with characteristic yellow-orange postmedial band). Discal spot visible on all wings from both sides. Within genus Psodos male and female similar in wing pattern but size of females often slightly smaller than males. The genus Psodos is characterized by the following characters (adults): ground colour of wings and body dark brown; strongly covered with hairy scales, especially around head; eyes small. Male genitalia: Uncus triangular, apically blunt or pointed. Gnathos triangular or tongueshaped, of various length between species. Valva wide, valvula membranous, sacculus sclerotized. Costa of valva strongly sclerotized, bearing multiple thick spines, connected to cucullus. The latter covered with hairy bristles. Juxta well developed and with two strongly sclerotized lobes, apically spined. Aedeagus thin, straight to s-shaped curved, distally ­tapered, with or without spines. Female genitalia: Papillae anales short, round, covered with tiny spines. Apophyses posteriores much longer (at least twice) than apophyses anteriores. Lamella postvaginalis strongly sclerotized, lateroventral part widely u-shaped, mediodorsal part usually developed as heart-shaped sclerite; posterior margin of sclerotized disc usually arched; inverted posterior part of sclerite, the ‘uncus gap’ (sensu Wehrli 1921) either widely opened or as a little notch. Lamella antevaginalis strongly sclerotized, mostly as a narrow, grooved band, clasping lamella postvaginalis ventrally. Antrum funnel- or tube-shaped, strongly sclerotized. Colliculum short, membranous. Ductus bursae sclerotized (with few exceptions), connected to a pear-shaped membranous corpus bursae. Signum strongly sclerotized, ovally inverted. Biology, phenology, habitat: Caterpillars probably hiding in sheltered places under rock fragments or in fissures, probably polyphagous, most instars feeding on low-growing plants. Adults univoltine, diurnal, heliophilous, flying only in sunlight. Flight period from mid-May to early September, with main activity in July. Adults mainly flying during morning hours in ­sunny sheltered places or sitting with folded-up wings usually on rocks. Some species also active in the afternoon after a break during midday (e.g. P. alticolaria, pers. observation). Vertical distribution from 200 m in Fennoscandia up to 4500 m in the Alps. Preferring habitats with vegetation-poor rocks or gravel surfaces in higher locations or in deeper alpine meadows. Immature stages: So far there is not much known about Psodos eggs, caterpillars and pupae. Successful rearing of these moths adapted to the high mountain climate is very difficult due to the lack of knowledge concerning their living conditions and food-plants. Remarks: The validation of the genus name Glacies was based on the assumption that the alpine species Sciadia tenebraria (auct.) (formerly Orphne tenebraria Esper, 1806) is more closely related to the species of ‘Glacies’ than to Psodos quadrifaria (Sulzer, 1776) (­Povolny & Moucha 1955). Therefore, Povolny & Moucha (1955) separated P. quadrifaria from Psodos by using the genus name Torula for that single species and the genus name Psodos for the rest of the group, overlooking that P. quadrifaria was actually the type species of the genus Psodos. Later, Varga (1975a; 1975b) corrected this nomenclatural mistake by using the replacement name Glacies for the remaining species. Remarkably, the species of Sciadia (cf. Huemer & Hausmann 2009) share the autapomorphic character states of Psodos as homoplasies due to their very similar ecobiotic mode of life. On the other hand, characters of the antennae, male and female genitalia and DNA analysis showed that the species of Sciadia are more closely related to Elophos, which was the reason to transfer some species of Elophos to Sciadia. This means, however, that there is no need to maintain Psodos as 128

a monotypic genus aside Glacies. Therefore, we consider Glacies as a junior synonym of Psodos Treitschke, 1825 (=Glacies Millière, [1874] syn. n.).

35. Psodos alpinata (Scopoli, 1763) Phalaena alpinata Scopoli, 1763: Ent. Carn.: 228, fig 571 (Slovenia, Carnia: Bohinj). Syntype(s), lost. Geometra horridaria Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener Gegend: 100 (Austria: Vienna district). Syntype(s) lost. Phalaena carbonata Schrank, 1802: Fauna boica 2 (2): 52 (Germany, Bavaria: Hohenschwangau Mts.). Syntype(s) not traced.

External characters and abdomen: Wingspan 21–27 mm. Upperside of wings brown, light brown or greyish brown, poorly contrasted. Distal margin in both wings slightly lighter. Postmedial and antemedial lines of forewing as well as medial line of hindwing dark and serrated; no silvery, shiny scales. Forewing with dark discal spot. Underside of wings almost uniform, except weakly lighter terminal area of wings. Antennae dorsally white-scaled. Variation: Little. Wing pattern more or less prominent. Medial areas sometimes darker than rest of wings. Male genitalia: Uncus broad, blunt. Gnathos widely tongue-shaped, reaching up to base of uncus. Costa of valva strongly sclerotized, distally curved, with numerous spines, distinctly separated from apex of valva. Juxta process characteristically slender, reaching up to base of costa of valva, terminally with strong spines pointing towards centre; inner margin of juxta processes slightly curved, connected to an x-shaped caulis (additional muscle-less sclerite between juxta and aedeagus). Saccus broadly rounded (in contrast to digitiform shape in most other Psodos species). Aedeagus straight, with two small, lateral spines at tip.

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Text-figs 56–58. Diagnostic characters (indicated) of selected European Psodos species. Upperside of wings. Text-fig. 56. P. alpinata (Scopoli, 1763), male. Austria: North Tyrol, Vennatal, 28.vi.1947 (ZSM). Text-fig. 57. P. canaliculata (Hochenwarth, 1785), male. Austria: North Tyrol, Hoher Burgstall, 16.vii.1941 (ZSM). Text-fig. 58. P. coracina (Esper, 1805), female. Italy: South Tyrol, Veneto, Dolomites, Valparolo Pass, 13.vii.1994 (coll. Erlacher).

Female genitalia: Antrum short, broad, tongue-shaped. Lamella postvaginalis: lateroventral part forming a straight line; sclerite distinctly heart-shaped; posterior margin of sclerite distinctly arched; ‘uncus gap’ widely opened with fissure extending anteriorly, posterior part of uncus gap strongly sclerotized. Lamella antevaginalis developed as a narrow, grooved band. Ductus bursae very short, partially sclerotized, with several folds. Corpus bursae membranous, signum slit-shaped. Distribution and abundance: European, alpine, with peri-alpine disjunction. Widely distributed in the Alps but with scattered distribution in the south-western Alps. Also recorded from several parts of Sudety mountains (Czech Republic) and Carpathians (Romania). Erroneously recorded from Pyrenees by Wehrli (1953) (Tautel 2011). – Very common, usually found in numbers. Phenology: Univoltine. From late May to late August, peaking in July. Larva from mid-July to mid-September, overwintering, and from May to June. 130

Text-figs 59–61. Diagnostic characters (indicated) of selected European Psodos species. Underside of wings. Text-fig. 59. P. alpinata (Scopoli, 1763), male. Austria: North Tyrol, Vennatal, 28.vi.1947 (ZSM). Text-fig. 60. P. canaliculata (Hochenwarth, 1785), female. Italy: Trentino-Alto Adige, Adamello, Rifugio Mandrone, late vii.1967 (coll. TLMF). Text-fig. 61. P. coracina (Esper, 1805), female. Italy: South Tyrol, Veneto, Dolomites, Valparolo Pass, 13.vii.1994 (coll. ­Erlacher).

Biology: Larva polyphagous. Recorded on low-growing plants (e.g. Leontodon, Rhododendron, Vaccinium), reared on Taraxacum (Flamigni et al. 2016). Larval stages: egg oval, in early stages yellowish, later brown with blood-red stains (Prout 1915; Forster & Wohlfahrt 1981). Habitat: Of all Psodos species (except P. quadrifaria) least specific in habitat choice, including alpine vegetation mats, both dry and humid, as well as on open rocky slopes, ridges and screes. Vertical distribution from 1400 up to 3100 m, usually between 1800 and 2500 m, exceptionally up to 3600 m (Adamello) and 4000 m (Margelkopf). Similar species: P. alpinata can easily be distinguished from the other species of the genus by the weak and uniform pattern of the wings with very weak markings especially on the underside. However, examination of genitalia recommended for secure identification. In contrast to all other Psodos species, male genitalia with broad and blunt uncus. Genetic data: BIN: BOLD:AAH7952 (n=23 from Germany, Austria, Switzerland, Czech Republic, Italy). Genetically slightly heterogeneous, maximum variation 1.2%. Nearest species: P. coracina (4.0%). 131

36. Psodos canaliculata (Hochenwarth, 1785) Phalaena canaliculata Hochenwarth, 1785: Schr. Berlin. Ges. Naturf. Freunde 6: 336, pl. 7, fig. 5 (Locus typicus and type deposition unknown). Geometra trepidaria Hübner, 1809: Samml. Eur. Schmett. 5, Geometrae. Pl. 66, fig. 343 (­Europe). Syntype(s) lost. Psodos trepidaria [forma] schwingenschussi Wehrli, 1919: Mitt. Ent.-Ver. Basel 11: 7, pl. 1, 2 (Carpathians: High Tatra. Lectotype ♂ (hereby designated) (ZFMK, examined, label data: Tatra, Blumengarten, 1800 m, 22.vii.1914, leg. Piesczek, prep. G0250). – Paralectotype 1♀ (hereby designated) (ZFMK, examined, label data: Tatra, Blumengarten, 1800 m, 21.vii.1914, leg. Piesczek, prep. G0253). Validated at species rank under Psodos by Scoble (1999), here downgraded to subspecies of P. canaliculata (stat. n.).

External characters and abdomen: Wingspan 17–24 mm. Ground colour of wing upperside varying from light to dark brown, with scattered creamy scales. Both ante- and postmedial lines serrated, with creamy edging. Antemedial line absent from hindwing. Medial band darker brown. Terminal line interrupted, with creamy edging. Underside light brown, with more uniform pattern; distal and costal areas lighter, sometime covered with more creamy scales. Antennae dorsally chequered black and white. Discal spots visible on all wings. Variation: Upperside of wings more covered with greenish, yellowish or silver metallic scales, resulting in duller appearance or darker and more uniform coloration. Underside with most conspicuous terminal area in the genus, often with traces of dark subterminal band, or nearly uniform. Subsp. schwingenschussi (High Tatra: wings light brown, with lighter terminal area and strongly contrasted wing pattern. Wing underside usually lighter, sometimes darker. Male genitalia: Uncus long and pointed. Gnathos tongue-shaped, reaching over the base of uncus. Costal process of valva well sclerotized. Costa of valva not separated, covered with a short row of sclerotized spines. Juxta processes asymmetrical, apically covered with a group of medially directed spines; the longer process reaching up to base of costa of valva; the shorter one three-quarters of the former. Aedeagus slightly arched, with numerous tiny apical spines. Female genitalia: Antrum slender, tubular, approximately as long as corpus bursae, ostium broad. Lamella postvaginalis with heart-shaped sclerite; uncus gap developed as a tiny notch. Lamella antevaginalis a narrow, grooved band. Colliculum short, membranous. Ductus bursae short, sclerotized. Corpus bursae pear-shaped. Signum mouth-shaped. Distribution and abundance: European, Alpine, with peri-alpine disjunction. Widely distributed in the Alps. Local occurrences in the Massif Central of France, in High Tatra and in several parts of the Carpathians (subsp. schwingenschussi). – Common but in limited numbers. – Specimens from Russia (Tuva) and Kazakhstan (leg. M. Česánek) seemingly conspecific, requiring further research (see Genetic data). Phenology: Univoltine. From late June to early September, mainly from mid-July to early August. Biology: Larva polyphagous on low-growing plants, especially on Orobanchaceae and Plantaginaceae (Flamigni et al. 2016): Recorded on Linaria alpina, Odontites lutea, Rhinanthus minor, Pedicularis comosa, P. rostratocapitata (= rostrata), Plantago alpina (Lhomme 1923–35). Reared to pupa by Wehrli & Imhoff (1922). Egg relatively large, oval, smooth, light green. Larva light brown, dorsal line continuous, alternating light and dark brown, 132

lateral line yellowish. Controversial descriptions of larva given by Prout (1915) and Forster & Wohlfahrt (1981). Pupa smooth, slender, shiny yellow brown, cremaster dark, at tip with four hooked cilia (Hoffmann 1911; Wehrli & Imhoff 1922; Forster & Wohlfahrt 1981). Habitat: Vertical distribution from 1000 up to 3700 m, mainly from 2000 up to 2800 m. Subsp. schwingenschussi from 1700 up to 2200 m. Preferring alpine grasslands, where vegetation is interrupted by stones, rocks and boulders, but also on rocky terrain and scree (Flamigni et al. 2016). Similar species: Confusion possible with P. bentelii, P. coracina and P. belzebuth. However, P. canaliculata differing in male genitalia by strongly asymmetrical juxta, with two ventral lobes of different length, in female genitalia by extremely long tubular antrum and wide sclerite of lamella postvaginalis. P. canaliculata with vague rest of pale line very close to termen of wing underside (in P. belzebuth the whole terminal area slightly paler). Genetic data: Four BINs: BOLD:AAE6736 (n=12 from Germany, Austria, Switzerland, northern Italy), BOLD:AAO1675 (n=2 from Russia: Tuva and Kazakhstan), BOLD:ACP8192 (n=1 from Italy: Friuli), BOLD:ABX0826 (n=1 from Italy: Aosta Valley). Genetically very heterogeneous, BINs diverging by 3.4–3.8%. Nearest species: P. alticolaria (5.7%), P. wehrlii (5.7%), P. noricana (5.8%). Remarks: The subsp. schwingenschussi was described as a separate species by Wehrli (1919). The original description does not exclude that schwingenschussi could also be a local form of P. canaliculata. Wehrli (1953) supplements that P. canaliculata is replaced by schwingenschussi in the Carpathians and is also characterized by asymmetrical juxta processes, differing in length and width from the alpine form. Based on the amount of ­examined material and on the P. canaliculata-related asymmetry in genitalia, schwingenschussi is treated as a subspecies of P. canaliculata (stat. n.). Psodos canaliculata shows a considerable amplitude of geographical variation and even subsp. schwingenschussi tends 133

to form isolated p ­ opulations in local restricted areas (Wehrli 1953; Povolny & Moucha 1955). In Scoble (1999), P. pyrenaica and P. werneri are listed as subspecies of P. canaliculata. Here, both taxa are treated as bona species (see remarks under species no. 37 and 42).

37. Psodos werneri Schawerda, 1916, bona species, stat. n. Psodos trepidaria werneri Schawerda, 1916: Verh. zool.-bot. Ges. Wien 66 (3–5): 245 (north-western Montenegro: Volujak mountains, 2000 m). Lectotype ♂ (hereby designated) (NHMW, examined, label data: [unreadable], Volujak, 2000 m, genit. prep. SE–716. – Paralectotype 1♂ (hereby designated) (NHMW, examined, label data: [unreadable], Volujak, 2000 m, genit. prep. SE–718). Validated as subspecies of ‘Glacies’ canaliculata in Scoble (1999), herewith raised to species rank, based on constant and significant differential features in genitalia.

External characters and abdomen: Wingspan 22–25 mm. Upperside of wing brown, pattern with poor contrast. Ante- and postmedial lines serrated, almost invisible. Antemedial line absent from hindwing. Underside brown, terminal area slightly lighter. Discal spot almost invisible from upperside. Variation: Unknown. Male genitalia: Uncus long, blunt. Gnathos tongue-shaped, reaching over base of uncus. Costal process of valva slightly sclerotized, not separated, covered with a short row of sclerotized spines. Juxta processes asymmetrical, apically covered with a group of medially directed spines. The longer process reaching up to base of costa of valva, the shorter one three-quarters of the length of the former. Hyaline membrane connecting juxta to vaseshaped caulis. Aedeagus slightly s-shaped, with unique large, stout sub-apical spine (the latter diagnostic within genus).

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Female genitalia: Unknown. Distribution and abundance: (South-east) European. Only known from Volujak mountains in north-western Montenegro, possibly more widespread in the Balkan peninsula. – Rare and very local. Phenology: Unknown. Biology: Unknown. Habitat: Adults collected at 2000 m. Similar species: For a secure identification examination of genitalia is necessary. In male genitalia differing from all other Psodos species by the unique large, stout sub-apical spine on aedeagus. Genetic data: Not yet DNA barcoded.

38. Psodos coracina (Esper, 1805) Phalaena coracina Esper, 1805: Die Schmetterlinge 4 (2), (53): 74, pl. 197, fig. 5 (’Alps of Jura mountains’ [possibly wrong according to Wehrli (1953)]. Psodos coracina tundrana Wehrli 1919: Mitt. Ent.-Ver. Basel 11: 8, pls 1, 2 (Russia, Sajan: Arasagun gol). Lectotype ♂ (hereby designated) (ZFMK, examined, label data: Russia, ­Sajan, Arasagun gol, #96). – Paralectotypes 7♂3♀ (hereby designated) (ZFMK, examined, label data: Russia/Mongolia, Sajan, Munku Sardyk, #95). Validated at subspecies rank by Scoble (1999). Psodos coracina transiens Wehrli, 1921: Mitt. Schweiz. ent. Ges. 13 (3/4): 170, figs 68, 69 (Austria, Tyrol: Arzelscharte). Lectotype ♂ (hereby designated) (ZFMK, examined, label data: Austria, Nordtirol, Karwendel, Innsbruck: Arzler Scharte, 24.vi.1920, leg. A. Deutsch). – Para­ lectotypes 10♂4♀ (hereby designated) (ZFMK, examined, label data: Nordtirol, Karwendel, Innsbruck: Arzler Scharte, 24.vi.1920, leg. A. Deutsch, #212). Validated at subspecies rank by Scoble (1999), here downgraded to synonymy (see Remarks). Psodos coracina lappona Wehrli, 1921: Mitt. Schweiz. ent. Ges. 13 (3/4): 170, figs 65, 66 (Norway, Finnmark: Alten). Lectotype ♂ (hereby designated) (ZFMK, examined, label data: Sweden, Norrbotten, Lule Lappmark, leg. F. Dannehl, prep. G0702). – Paralectotypes 4♂3♀ (hereby designated) (ZFMK, examined, label data: Lappland, vii.1908, leg. F. Dannehl, prep. G0703). Validated at subspecies rank by Scoble (1999), confirmed. Psodos coracina [forma] pseudonoricana Wehrli, 1921: Mitt. Schweiz. ent. Ges. 13 (3/4): 169, pl. 9, figs 63, 64 (Slovenia: Triglav). Lectotype ♂ (hereby designated) (ZFMK, examined, label data: Slovenia, Carniola, Julische Alpen: Triglav, 25.vi.1908, #817). – Paralectotype 1♀ (hereby designated) (ZFMK, examined, label data: Slovenia, Carniola, Julische Alpen: Triglav, 20.vii.1913, #691). Validated at subspecies rank by Scoble (1999) and Flamigni et al. (2016), but downgraded to synonymy by Leraut (2009), here retained as infrasubspecific form. Psodos coracina dioszeghyi Schmidt, 1930: Int. Ent. Zeits. 24 (3): 25, pl. A, fig. 1; pl. B, fig. 9; pl. C, figs 15–20 (Romania, Retyezat Mts.: Zanogasee, over 2000 m). Lectotype ♂ (hereby designated) (NHMW, examined, label data: Romania, Retyezat, Aragyes-teto, 18.–19.vii.1914). Validated at species rank under Psodos by Scoble (1999) and Leraut (2009), herewith downgraded to subspecies of P. coracina (stat. n.). Psodos coracina tundranoides Malicky, 1966: Mitt. Schweiz. ent. Ges 38 (3/4): 177 (Switzerland, Wallis: Mount Bonvin, 2995 m). Holotype ♂ (TLMF, examined). Paratypes: 1♂2♀ (TLMF, examined). Validated at subspecies rank by Scoble (1999), here downgraded to synonymy, syn. n. (see Remarks).

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Glacies coracina bureschi Varga, 1975b: Acta Biol. Debrecina 12: 77–90 fig. 5; text-fig. 2 (­Bulgaria, Pirin Mts.: NW side of Vichren, 2800 m). Holotype ♂ (Coll. Z. Varga. Not examined). – Paratype 1♂ (ZSM, examined). Validated at subspecies rank by Scoble (1999), confirmed.

External characters and abdomen: Wingspan 19–24 mm. Upperside of wings shiny, wing pattern well contrasted, ground colour light grey to dark brown, with slightly lighter terminal area. Forewing with dark brown basal area, ante- and postmedial lines dark grey with light edging, serrated; terminal line brown, weakly interrupted. Markings on hindwings less intense, antemedial line absent. Underside of wings slightly lighter, postmedial line present on both wings. Due to high variation of wing pattern several subspecies are described in Europe. Variation: See subspecies. Specimens from Slovenia, Triglav region and East Tyrol with less contrasted pattern on forewings, underside varying from less to distinctly contrasted. Subsp. bureschi (Pirin mountains): ground colour of wing upperside uniform, pattern little contrasting; dark graphite grey (Varga 1975b). Subsp. dioszeghyi (Retyezat mountains): wing pattern less intense, transverse lines partly very weak and less contrasted; terminal area of underside indistinctly contrasted from rest of wing colour. Subsp. lappona (Fennoscandia): very variable, colour and pattern varying from less contrasted, dark brown to lighter with distinctly contrasted transverse lines. Upperside of wings slightly metallic blue scaled. Underside with weakly contrasted terminal area. Male genitalia: Uncus relatively broad, distally pointed. Gnathos tongue-shaped, reaching up to base of uncus. Valva broad, basally with strong sclerotization. Costa of valva distinctly separated from apex of valva, distally curved, with numerous strongly sclerotized spines arranged in one row, getting longer towards apex. Juxta process short (shorter than in all other Psodos species), terminally with a row of spines, medially directed, with one to three large spines; juxta process not reaching up to base of costa of valva. A hyaline membrane connecting juxta with funnel-shaped caulis. Aedeagus relatively long compared with ­juxta processes, straight, with two small lateral spines at tip. Genitalia of Slovenian populations (‘pseudonoricana’) larger than those of the other subspecies, juxta processes broader, longer and almost reaching up to base of costa of valva, distal spines distinctly more strongly marked. In male genitalia of subsp. lappona dorsally directed spine of juxta processes sometimes absent. Female genitalia: Papillae anales small, broad. Antrum short, tongue-shaped, broader only towards ostium. Lateroventral part of lamella postvaginalis forming a straight line, sclerite distinctly heart-shaped, posterior margin of sclerite distinctly arched, ‘uncus gap’ developed as deep gap, posteriorly not sclerotized. Lamella antevaginalis developed as a narrow, grooved band. Ductus bursae very short, membranous. Corpus bursae membranous, signum absent. Distribution and abundance: Euro-Siberian, arctic-alpine. Widely distributed in the Alps and in the higher areas of the Aragonese and Catalan Pyrenees. Also present in Balkan pen­ insula in the Bulgarian Pirin Mountains (subsp. bureschi) and in the Retyezat Mountains of the Romanian south-western Carpathians (subsp. dioszeghyi). In contrast to all other species of Psodos, P. coracina is widely distributed in the Scandinavian mountains, from western Norway to Murmansk coast of the Russian peninsula Kola (Skou 1986; Viidalepp 1996). Occasionally distributed in Scotland (Gardner 1950; Skinner 1984), in the Tatra in Lower 136

Austria and in the southern Czech Republic (Malkiewicz 2012; Huemer 2013). – Common, usually in numbers. – Outside Europe replaced by subsp. tundrana in central Asia in the Sajan mountains (Skou 1986; Viidalepp 1996).

Phenology: Univoltine. From mid-June to early September with main flight period in July. Larva overwintering twice (Skinner 1984). Biology: Larva polyphagous on low-growing plants, e.g. possibly on Malva (Flamigni et al. 2016). In Scandinavia recorded on Empetrum nigrum and Betula nana (Skinner 1984; Skou 1986; Flamigni et al. 2016), in Scotland on Arctostaphylos uva-ursi (Skinner 1984), in Pyrenees on Dryas octopetala (Flamigni et al. 2016). Reared on Polygonum aviculare, Sonchus and Leontodon (Fleischmann 1900) and on Plantago lanceolata (W. Wolf pers. comm.). Adults active during daytime, resting on sun-heated ground, rarely on flowers. In Italy larva found from early August to mid-September and in June; hibernation in first winter as larva, in second winter as pupa (Flamigni et al. 2016). Egg oval, colour turning from yellow-green through coral-red, bronze-brown to black (Gardner 1950). Larva reminiscent of that of P. noricana, but light brown instead of grey-yellow (May & Fleischmann 1899; Prout 1915; Gardner 1950; Forster & Wohlfahrt 1981). Pupa light brown, turning black before hatching (Gardner 1950). Habitat: Vertical distribution from 1700 up to 3600 m, in Scandinavia also found at sea-level, here up to 1100 m (Skou 1986); on sunny meadows, scree and rock (Flamigni et al. 2016); in Scandinavia in marshes, mountain birch forests as well as in the conifer zone and on mountain moors (Skou 1986). Similar species: Psodos coracina in particular reminiscent of P. canaliculata and P. ­alpinata. However, wing upperside of P. canaliculata more or less sprinkled with metal greenish, 137

y­ ellowish or silver scales, resulting in more speckled appearance, ground colour of P. cora­ cina more uniform, bright dark grey with conspicuous transverse lines and discal spot (­Flamigni et al. 2016). Wing upperside of P. alpinata more uniform than that of P. coracina, wing pattern hardly recognizable. See under P. noricana for differences from f. pseudonoricana. For secure identification, examination of genitalia is required. In contrast to P. alpinata, ‘uncus gap’ in female genitalia of P. coracina dorsally not sclerotized and corpus bursae not or only slightly sclerotized. Genetic data: BIN: BOLD:AAD3142 (n=44 from Finland, Sweden, United Kingdom, Germany, Switzerland, Austria, Italy, Slovenia, Russia, Kazakhstan). Genetically slightly heterogeneous, mostly without clear-cut geographical patterns, but seven BIN-sharing DNA barcodes from Kazakhstan and Russia (Altai, Tuva: subsp. tundrana Wehrli, 1919) at a minimum distance of 0.6%. Nearest species: P. alpinata (4.0%), P. alticolaria (5.2%), P. sajana (4.9%). Remarks: Wehrli (1953) suggested the type locality was almost certainly in Graubünden (Switzerland). In that case, the type locality of P. coracina is overlapping with the distribution range of transiens. Also the distribution range of tundranoides cannot clearly be distinguished from the nominotypical subspecies. Therefore, these taxa are downgraded to synonyms of Psodos coracina coracina Esper, 1805 (= Psodos coracina transiens Wehrli, 1921 syn. n., = Psodos coracina tundranoides Malicky, 1966 syn. n.). Due to morphological conformity in the male and female genitalia, Psodos dioszeghyi is proposed to be a subspecies of P. coracina (Psodos coracina dioszeghyi Schmidt, 1930 stat. n.). Despite small differences in male genitalia between pseudonoricana Wehrli, 1921 and the nominotypical subspecies, we follow here the concept of Leraut (2009) treating this taxon as synonym to P. coracina coracina. Similar to P. noricana f. kusdasi and P. alticolaria f. chalybaeus, it seems to be just an ecological form adapted to the geological substrate. This is also supported by the lack of significant genetic differences with respect to the respective nominotypical subspecies. The Japanese taxon daisetsuzana Matsumura, 1927 (Insecta matsumura 1 (4): 186; Syntypes 10♂, examined) was considered subspecies of P. coracina by Scoble (1999), but treated as synonym of the Asian species P. sajana Wehrli, 1919 by Beljaev (2016), the latter view here confirmed.

39. Psodos bentelii Rätzer, 1890 Psodos bentelii Rätzer, 1890: Bull. Soc. ent. Suisse 8 (6): 224 (Switzerland, Valais: Gornergrat). Syntypes 3♂1♀ (NMBE, not examined). Psodos perlinii Turati, 1914: Atti Soc. ital. Sci. Nat. (Pavia) 53: 468–619 (Italy, Brescia, Valcamonica: Lago Salarno, 3000 m). Lectotype ♂ (designated in Erlacher & Junghans 2009) (ZFMK, examined, label data: Italy, Brescia, Valcamonica, Lago Salarno, 3000 m, prep. G1447). Validated at subspecies rank under P. coracina by Scoble (1999), subordinated as subspecies of P. bentelii in Erlacher & Junghans (2009). ‡ Psodos bentelii [forma] zermattensis Wehrli, 1919: Mitt. Ent.-Ver. Basel & Umgebung 11: 5, pl. 1, 2 (Switzerland, Valais: Zermatt). Syntypes (ZFMK, examined). Infrasubspecific. Validated at subspecies rank by Scoble (1999), treated as a synonym of P. bentelii by Flamigni et al. (2016). Psodos bentelii retyezatensis Bartha, 1933: Mitt. münch. ent. Ges. 23: 41, pl. 7, fig. 1 (Romania: Retyezat Mts.). Syntypes 2♂ (HNHM, not examined). Validated at subspecies rank by Scoble (1999), here accepted. Psodos telekii Bartha, 1933: Mitt. münch. ent. Ges. 23 (2): 39, pl. 7, fig. 2 (Romania: Retyezat Mts., 2300 m). Holotype ♂ (HNHM, not examined). Synonym of P. bentelii (Scoble 1999).

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External characters and abdomen: Wingspan 21–28 mm. Upperside of wings brown irrorated with white scales; ante- and postmedial lines serrated, dark brown; terminal line brown, interrupted. Antemedial line absent from hindwing. Underside dark, costal area and terminal area of forewing slightly lighter, apex dark brown; postmedial and terminal lines visible, with white edging. Discal spot present on all wings. Variation: See subspecies. Subsp. perlinii (Adamello mountains): forewings black, with greenish blue to metallic shiny scales; two distinct white lines in the terminal area of wing underside, proximal one broader than distal (for detailed description see Erlacher & Junghans 2009). Subsp. retyezatensis (south-western Carpathians): smaller, forewings less contrasted, markings weak, medial line weakly serrated. Upperside of hindwings and underside of wings almost without markings. Male genitalia: Uncus pointed. Gnathos long, slender, pointed, almost reaching up to tip of uncus. Valva broad, basally with strong sclerotization. Costa of valva strongly sclerotized, apically separated, with numerous strongly sclerotized spines, arranged in one row. Juxta process long, reaching up to base of costa of valva, basally conical and slender, distally broadened, slightly arched on the inner side, terminally with a group of strongly sclerotized and medially directed spines; underneath a larger triangular tooth and another, dorsally directed spine on the inner side of juxta. Caulis elongated, with hyaline membrane connected to juxta. Saccus pointed. Aedeagus thin, slightly s-shaped, vesica with numerous small spines. Female genitalia: Antrum widely funnel-shaped. Lamella postvaginalis with lateroventral margin forming a straight line; sclerite widely heart-shaped; posterior margin of sclerite not distinctly arched; uncus gap developed as small notch without anteriorly extending fissure 139

Text-figs 62–64. Diagnostic characters (indicated) of selected European Psodos species. Upper­ side of wings. Text-fig. 62. P. bentelii Rätzer, 1890, male. Italy: Aosta, Matterhorn, Breuil-­ Cervinia, 15.vi.1967 (coll. Gelbrecht). Text-fig. 63. P. noricana Wagner, 1898, male. Austria: North Tyrol, Ötztaler Alpen, Obergurgel, 17.vii.1951 (ZSM). Text-fig. 64. P. belzebuth Praviel, 1938, male. France: Alpes-de-Haute-Provence, Colmars, 29.vi.1924 (ZSM).

Lamella antevaginalis developed as a grooved band with medial gap and weakly sclerotized, dorsally directed, arched corners. Colliculum short, membranous. Ductus bursae sclerotized. Corpus bursae large, membranous. Signum strongly sclerotized, inverted mouth-shaped,. Distribution and abundance: European, West-Alpine. Widely distributed in the western Alps from French Alps to Dolomites. Replaced by subsp. perlinii in Adamello mountains, by subsp. retyezatensis in south-western Carpathians. – Usually in limited numbers. Phenology: Univoltine. From early July to late August, with main flight period from mid-July to late July. Adults active in the morning. Biology: Larva polyphagous, feeding on low-growing plants: Recorded e.g. on Silene acaulis (Flamigni et al. 2016). Reared on Sedum album and S. telephium, but also on Trifolium sp., Daucus carota, Plantago sp., Artemisia absinthium and Taraxacum sp. (Wehrli & Imhoff 1922; Thomann 1930; Forster & Wohlfahrt 1981; Flamigni et al. 2016). Hibernating twice (Flamigni et al. 2016).

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Text-figs 65–67. Diagnostic characters (indicated) of selected European Psodos species. Underside of wings. Text-fig. 65. P. bentelii Rätzer, 1890, male. Italy: Trentino-Alto Adige, Adamello, Rifugio Mandrone, 7.–10.vii.1964 (coll. TLMF). Text-fig. 66. P. noricana Wagner, 1898, male. Austria: North Tyrol, Innsbruck, Nordkette, 24.vii.1994 (coll. Erlacher). Text-fig. 67. P. belzebuth Praviel, 1938, male. France: Alpes-de-Haute-Provence, Colmars, 29.vi.1924 (ZSM).

Habitat: Vertical distribution from 1500 up to 3300 m, mainly from 2000 up to 3000 m, one specimen from 3800 m (Trifthorn). Preferring habitats on rocky walls and steep slopes with grass and lichen tufts (Flamigni et al. 2016). Similar species: In contrast to P. noricana without green tinge even in fresh specimens. P. canaliculata differing in size (on average slightly larger), and in more uniform dark slate colour. Examination of genitalia required for secure identification. P. bentelii clearly differing from all other Psodos species by distally club-like broadened juxta processes in male genitalia. Female genitalia of P. bentelii similar to those of P. noricana, the former often with screw-like rotation in anterior part of antrum, never occurring in P. noricana. In addition, medial gap of lamella antevaginalis of P. bentelii considerably narrower and more conspicuous than equivalent in P. noricana.

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Genetic data: Three BINs: BOLD:ACE3447 (n=4 from Italy: Lombardy, South Tyrol), BOLD:AAL3170 (n=4 from Switzerland: Valais; Italy: Aosta Valley), BOLD:ACV2332 (n=2 from north-eastern Italy: Veneto, Dolomites). Genetically heterogeneous, BINs at distances of 1.1% (BOLD:ACE3447 / BOLD:AAL3170), north-eastern Italian populations closer to P. noricana. Nearest species: P. noricana (1.8% from north-eastern Italian BOLD:ACV2332), P. belzebuth (4.6%). Remarks: P. bentelii zermattensis cannot be considered a geographically isolated population, in genitalia identical with those of P. bentelii, also occurring near Zermatt. Therefore, P. bentelii zermattensis was treated as a synonym of P. bentelii by Flamigni et al. (2016). Most probably, telekii Bartha, 1933 is only an aberration of subsp. retyezatensis (Bartha 1933; Wehrli 1953; Povolny & Moucha 1955). However, further material and integrative study is necessary to re-examine the status.

40. Psodos noricana Wagner, 1898 Psodos noricana Wagner, 1898: Verh. zool.-bot. Ges. Wien 48 (10): 715 (Austria, Styria, Hochschwab: Fölzalm, 1472 m). Psodos noricana var. carpathica Schwingenschuss, 1915: Verh. zool.-bot. Ges. Wien 65 (5/6): 83 (Slovakia, Tatra: Beler Kalkalpen, 1800 m). Holotype ♂ (ZFMK, examined). – Paratypes: 1♂1♀ (ZFMK, examined, label data: Tatra, Beler Kalkalpen, 1800 m, gen. preps. 209 and G0245). Validated at subspecies rank by Scoble (1999), here confirmed. Psodos noricana kusdasi Wehrli, 1945: Mitt. Schweiz. ent. Ges. 19 (9): 337 (Austria, Salzburg, Hohe Tauern: Gleiwitzer Hütte). Lectotype ♀ (hereby designated) (ZFMK, examined, label data: Austria, Salzburg, Glocknergruppe, Brandlscharte, Gleiwitzer Hütte, 2400 m, 07.–13. vii.1941, leg. K. Kusdas, prep. G0246). – Paralectotype 1♂ (hereby designated) (ZFMK, examined, label data: Austria, Salzburg, Glocknergruppe, Brandlscharte, Gleiwitzer Hütte, 2400 m, 07.–13.vii.1941, leg. K. Kusdas, G7508). Further Paralectotypes in NHMW. Validated at subspecies rank by Scoble (1999), downgraded here to synonymy of P. noricana Wagner, 1898 (syn. n.) based on the almost overlapping distribution with the nominotypical subspecies. Psodos burmanni Tarmann, 1984: Z. Arb.gem. Österr. Ent. 36 (1/2): 1, pl. 1, figs a–d; pl. 2, figs a–b (Austria, North Tyrol, Ötztal Alps, Obergurgl, Rotmoostal: Rotmoosferner, 2400 m. Holotype ♂ (TLMF, examined). – Paratypes 7♂3♀ (TLMF, examined). Validated at species rank by Scoble (1999), downgraded to synonymy by Leraut (2009).

External characters and abdomen: Wingspan 20–27 mm. Upperside of wings grey-brown to light brown, with very few scattered metallic blue scales, fresh moths with distinctly green, shiny scales. Terminal area of wings (on both sides) slightly paler. Ante- and postmedial lines serrated, terminal line interrupted; antemedial line absent from hindwing. Underside light brown. Discal spot present, hardly visible on underside. Antennae dorsally indistinctly chequered black and white. Variation: See subspecies. Populations on primary rock in eastern Alps (High Tauern, Deferegg Alps to Ötztal Alps) slightly smaller, wings light yellowish brown. Subsp. carpathica (High Tatra): larger, wings darker and with more contrasted pattern. Male genitalia: Genitalia very large. Uncus short, pointed. Gnathos long, slender and pointed, almost reaching up to tip of uncus. Costal process of valva well sclerotized, apically separated from costa of valva, with strongly sclerotized stout spines. Juxta process large, lateral margins distally extended, apically sickle-shaped, inner margins covered with medially directed, strongly sclerotized spines; the largest triangular, at the anterior end, diagnostic. 142

A hyaline membrane connecting juxta with caulis. Aedeagus slightly s-shaped, apically with numerous small spines. Female genitalia: Antrum widely funnel-shaped. Lamella postvaginalis with widely heartshaped sclerite; posterior margin of sclerite weakly arched; ‘uncus gap’ with large notch, without anteriorly extending fissure. Lamella antevaginalis developed as a narrow, grooved band, with medial, widely opened gap. Colliculum short, membranous. Ductus bursae widely sclerotized. Corpus bursae pear-shaped, membranous. Signum strongly sclerotized, inverted mouth-shaped. Distribution and abundance: European, east-alpine, with peri-alpine disjunction. Widely distributed in the eastern Alps, especially on limestone. Additionally present in Germany (Bavaria), France (Chartreuse Mts.), northern Italy (from Bergamasque Alps to Carnic and Julian Alps). Isolated occurrences in High Tatra (i.e. Belianske Tatra; subsp. carpathica) and in the Retyezat Mountains. – Common, but usually in limited numbers.

Phenology: Univoltine. From mid-May to early September. Biology: Larva reared on Polygonum aviculare, Sonchus spp. and Leontodon (Fleischmann 1900). Egg oval, colour turning from yellow-green to bronze-brown. Larva grey-yellow, in early stages more brightly coloured with dark dorsal line and blue-white lateral line (May & Fleischmann 1899; Prout 1915; Forster & Wohlfahrt 1981). Pupa slender, shiny yellow-brown, dorsal line dark brown, shafts for legs and antennae greenish brown, cremaster at tip tapering to a single projection (May & Fleischmann 1899; Forster & Wohlfahrt 1981). Habitat: On south-exposed, rocky slopes, on screes with scarce vegetation and on glacial moraines. Vertical distribution from 1300 up to 3100 m, mainly from 1500 up to 2500 m. On limestone and on primary rock. 143

Similar species: Psodos noricana differing from P. alpinata by greenish grey shimmer on wing upperside and by more distinct and more strongly dentate transverse lines. P. coracina (in particular f. pseudonoricana) differing by better marked transverse lines and discal spot, discal spot on underside of hindwing clearly recognizable in P. coracina f. pseudonoricana (very weak or absent in P. noricana). Furthermore, P. noricana slightly larger than P. coracina and with slight green shimmer on wing upperside (missing in P. coracina). For secure identification examination of genitalia is required. Male genitalia of P. noricana clearly differing from all other Psodos species by extremely extended juxta processes. Female genitalia of P. noricana reminiscent of those of P. bentelii, the latter with screw-like sclerotization at anterior part of the antrum (missing in P. noricana), small uncus gap and round margin of lamella postvaginalis (larger uncus gap and heart-shaped lamella postvaginalis in P. noricana). Genetic data: BIN: BOLD:AAI7576 (n=9 from Germany, Austria, Slovenia). Genetically homo­geneous, one specimen from Slovenia diverging by 1.2%. Nearest species: north-­ eastern Italian P. bentelii (BOLD:ACV2332: 1.8%), P. wehrlii (4.9%), P. spitzi (4.9%). Remarks: The view of Leraut (2009), regarding subsp. carpathica Schwingenschuss, 1915 as a subspecies of P. canaliculata, cannot be confirmed. The examined genitalia of the types of carpathica from Tatra mountains, collected by Wehrli, shows close relationship with P. noricana. Nevertheless, both the status of subsp. carpathica and f. kusdasi (=burmanni Tarmann, 1984) is controversial (Malkiewicz 1999; Huemer 2013).

41. Psodos belzebuth Praviel, 1938 Psodos noricana belzebuth Praviel, 1938: Bull. Soc. entomol. France 43 (7/8): 83, fig. 2 (France, Alpes-Maritimes: Mont Mounier). Lectotype ♂ (hereby designated) (ZSM, examined, ­label data: France, Alpes-Maritimes, Mont Demaut, [44°11’N, 07°02’E], 09.viii.1936, leg. J. ­Gazel). Syntypes 22♂4♀ (MNHN, not examined).

External characters and abdomen: Wingspan 19–22 mm. Upperside of wings brown, with few silvery blue scales especially scattered towards wing costa. Ante- and postmedial lines serrated, terminal line interrupted. Antemedial line absent from hindwing. Underside almost uniform, terminal area of all wings slightly lighter, postmedial line present. Dark brown discal spot visible on all wings and from both sides. Antenna dorsally chequered black and white. Variation: Little. Male genitalia: Medium sized. Uncus small, pointed. Gnathos tongue-shaped with slightly rounded tip, almost reaching up to tip of uncus. Costal process of valva well sclerotized, apically not separated from fore-edge of valva, covered with more or less sclerotized spines; a triangular spine on anterior margin of juxta. Aedeagus almost arched at centre, with very few, tiny apical spines. Female genitalia: Antrum slender, funnel-shaped, strongly tapering at base. Lamella postvaginalis with sclerite, widely heart-shaped; uncus gap tiny, without anteriorly extending fissure. Lamella antevaginalis developed as narrow, grooved band with medial gap and strongly sclerotized, dorsally directed, arched corners. Colliculum long, membranous. Ductus bursae very short, sclerotized. Corpus bursae pear-shaped, signum mouth-shaped. Distribution and abundance: (South-west) Alpine. Restricted to a limited area in the French (Alpes-de-Haute-Provence, Alpes-Maritimes) and Italian Alps (Piedmont in the Maritime

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and southern Cottian Alps). Tentatively attributed taxon panticosea in the Pyrenees (see Remarks). – Usually in limited numbers. Phenology: Univoltine. From mid-July to mid-August. Flight activity starting early in the morning, depending on sunlight conditions, with only a few specimens flying in the afternoon (Gianti 2012). Biology: Larval stages unknown. Adults flying during daytime, especially in the morning, on ground or on rocks, rarely on flowers (Flamigni et al. 2016). Habitat: Adults flying on south-facing slopes, in habitats with occurrence of Leontopodium alpium (Gianti 2012). Vertical distribution from 1300 up to 2900 m, mainly from 2400 up to 2500 m (Gianti 2012). On limestone (Wehrli 1953). Similar species: Psodos belzebuth differing from P. bentelii, P. noricana and P. alpmaritima by uniform greyish green ground colour. Examination of genitalia required for secure identification. In male genitalia, sickle-shaped juxta process very similar to that in P. noricana, but without medially directed spines. Female genitalia of P. belzebuth clearly distinguishable from those of P. noricana and P. bentelii by smaller size, slender funnel-shaped antrum and dorsally directed arched corners of lamella antevaginalis. Genetic data: BIN: BOLD:AAF5563 (n=7 from Italy, eastern France). Genetically homo­ geneous. Nearest species: P. alpmaritima (1.7%), P. noricana (4.3%). Remarks: For detailed report on morphology and habitat of P. belzebuth see Gianti (2012). For the taxon Psodos bentelii panticosea Wehrli, 1945 (Mitt. Schweiz. ent. Ges. 19 (9): 336 (Spain, Aragon: Bad Panticosa, 3000 m) we hereby designate the Lectotype: ♂, ZFMK (examined), label data: Aragon, Bad Panticosa, 20.vii.1928, 3000 m, leg. E. Wehrli, prep. G7587). Paralectotype 1♂, hereby designated (ZFMK, examined, label data: France, 145

Pyrénées-Orientales, Haute-Vallé-de-Montet, 26.vii.1909, prep. G0208). The taxon was validated as subspecies of ‘Glacies’ bentelii by Scoble (1999), synonymized with P. pyrenaica by Leraut (2009). The examination of the type specimens of panticosea, however, indicates a close relationship to P. belzebuth, even closer than to P. bentelii (under which it was subordinated in the original description), considering the shape of valva and the spines of the juxta processes. The latter show a big tooth on the proximal and on the distal end of the diagonal inner margin (as in P. belzebuth), whereas the juxta processes of P. bentelii do not show a diagonal inner margin and distally have several stronger teeth. Nevertheless, this taxon is attributed to P. belzebuth here only tentatively. Requiring further research, including DNA barcoding.

42. Psodos pyrenaica Schawerda, 1919 Psodos trepidaria pyrenaica Schawerda, 1919: Zeits. Österr. Ent.-Ver. 4 (7): 68 (France, HautesPyrénées: Pic du Midi). Syntypes ♂♀ (NHMW, not examined). Validated as subspecies of ‘Glacies’ canaliculata by Scoble (1999), raised to species rank by Leraut (2009), here accepted.

External characters and abdomen: Wingspan 20–23 mm. Ground colour of wing upperside brown with golden brown to yellow tinge in terminal area. Discal spots distinct. Underside with whitish terminal area, sometimes with slight sign of two white terminal lines. Variation: Unknown. Male genitalia: Genitalia medium-sized. Uncus small, pointed. Juxta process distally usually only with one strong tooth. Female genitalia: Unknown. Distribution and abundance: Pyrenean, endemic. Distributed in the eastern and western Pyrenees in both France and Spain.

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Phenology: Flight period in July. Biology: Larval stages unknown. Habitat: Vertical distribution from 2400 up to 3000 m. Similar species: According to Leraut (2009) wing colour and pattern are similar to that of P. canaliculata but widely suffused with yellowish hue, underside yellow, olive-tinged in terminal area. For secure identification examination of genitalia is necessary. Male genitalia with juxta process similar to that in P. belzebuth, but P. pyrenaica with shorter diagonal inner margin between proximal and distal tooth. Genetic data: Not yet DNA barcoded. Remarks: Leraut (2009) listed the taxon panticosea Wehrli, 1945 as a synonym of P. pyrenaica. For identity see P. belzebuth (no. 41).

43. Psodos alpmaritima Wehrli, 1924 Psodos alpmaritima Wehrli, 1924: Dt. Ent. Z. Iris 38 (2): 90, pl. 1, fig 8. (Italy, Piedmont, Alpi Marittime, Prov. Cuneo: top of Mt. Argentera, 3300 m). Lectotype ♂ (hereby designated) (ZFMK, examined, label data: France, Alpes-Maritimes: Cime d’Argentera, 3300 m, gen. – slide. G259).

External characters and abdomen: Wingspan 19–22 mm. Upperside of wings dark brown, with dark brown ante-, postmedial and terminal lines, not serrated as in most Psodos species; supported with shadow; terminal line not interrupted (antemedial line absent on hindwing). Underside dark brown, costal area and terminal area of forewing slightly darker; postmedial and terminal lines partially visible, white edging of these lines clearly visible on both wings. All wings with discal spot. Variation: Shape of juxta in male genitalia sometimes variable (Flamigni et al. 2016). Male genitalia: Uncus pointed. Gnathos long, slender, pointed, almost reaching up to tip of uncus. Valva broad, costa well sclerotized, not separated from valva, covered with numerous sclerotized spines. Juxta process cylindrical, reaching up to base of costa of valva; terminally covered with one stout, medially directed, and a few smaller but strongly sclerotized spines; basally conical. Aedeagus arched, vesica with numerous small spines. Female genitalia: Unknown (female specimen of type series belonging to P. bentelii). Distribution and abundance: (South-west) Alpine. Restricted to Alpes Maritimes (Mount Argen­tera, Saint-Martin, Vésubie) on both sides of the Italian-French border. – Very local, in very limited numbers. Phenology: Univoltine. Early July to early August. Biology: Larval stages unknown. Habitat: Vertical distribution from 2500 up to 3350 m. On screes below ridges of gneiss (Gianti 2012) and granite tock (Flamigni et al. 2016). According to Flamigni et al. (2016) P. alpmaritima often occurs sympatrically with P. canaliculata or P. bentelii.

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Similar species: Wing pattern similar to that of P. bentelii, differing from the latter by darker grey colour, tending more to brown, and by more indistinct pattern, in particular on underside. Examination of genitalia required for secure identification. Based on shape of juxta processes and low number of spines in male genitalia this species can be easily distinguished from other Psodos species. Genetic data: BIN: BOLD:ADK2954 (n=1 from south-eastern France; by the courtesy of G. Fiumi). Nearest species: P. belzebuth (1.7%), P. noricana (3.5%), P. bentelii (4.6%). Remarks: Psodos alpmaritima was originally described as a subspecies of P. bentelii by Wehrli (1924). However, sympatric distribution of both taxa along with a genetic distance to P. bentelii of 4.6%, confirm the validity at species level. This hypothesis was confirmed earlier by Leraut (2009), Erlacher (in Tautel 2011) and was followed by Flamigni et al. (2016).

44. Psodos alticolaria Mann, 1853 Psodos alticolaria Mann, 1853: Verh. zool.-bot. Ges. Wien 3: 75 (Austria, Hohe Tauern, Großglockner: Gamsgrube). Syntypes 1♂1♀ (NHMW, not examined, could not be traced). Psodos alticolaria var. gedrensis Rondou, 1903: Actes Soc. Linn. Bordeaux 57: 321 (France: Pyrenees, Gèdre). Syntype(s), no type material mentioned in the original description. Validated at subspecies rank by Scoble (1999), here, subspecies rank tentatively accepted, requiring more research. Psodos alticolaria var. faucium Favre, 1905: Mitt. Schweiz. ent. Ges. 11 (2): 147 (Switzerland, Valais: on Forcletta and Augstbordpass). Syntype(s), no type material mentioned in the original description. Validated at subspecies rank by Scoble (1999), here, subspecies rank tentatively accepted, requiring more research. Psodos frigidata Vorbrodt & Müller-Rutz, 1913: Schmetterlinge Schweiz 2 (2): 185 (Switzer­ land, Valais: Col du Torrent, Col de Sorebois, Augstbordpass, Turtmanntal, Grubener-­

148

Schwarzhorn, Col Forcletta, Gornergrat, Dents du Midi. France, Haute-Savoie, Col de Balme am Croix de Fé). Syntype(s), no type material mentioned in the original description. Synonym of P. a. faucium (Scoble 1999). Psodos chalybaeus Zerny, 1916, in Rebel: Verh. zool.-bot. Ges. Wien 66 (3–5): 115 (Italy, South Tyrol, Ortler Mts: Stilfser Joch). Lectotype ♂ (hereby designated) (NHMW, examined, label data: Italy, South Tyrol, Ortler Mts, Stilfser Joch, 21.vii.1896, leg. Habich). Validated at species rank by Scoble (1999) downgraded to subspecies rank by Flamigni et al. (2016). Herewith downgraded to synonymy of P. a. alticolaria (syn. n.) based on the overlapping distribution patterns of the forms denoted as “chalybaeus” and on the lack of significant genetic differences with respect to the nominotypical subspecies.

External characters and abdomen: Wingspan 21–26 mm. Upperside of wings with ground colour varying from light brown to dark brown, and grey. Ante- and postmedial lines brown, sometimes serrated (antemedial line absent from hindwings). Postbasal, medial and postmedial bands cream to light brown. Terminal line almost continuous, with creamy edging. Underside of wings cream to light brown; only white postmedial and terminal lines distinctly visible in terminal area of underside. Discal spot large and conspicuous. Variation: See subspecies. Specimens from several regions (e.g. Rhaetian Alps, Switzerland, northern Italy) with rounded, not dentate postmedial lines and more contrasted, i.e. with dark fasciae on light areas (f. chalybaeus). Subsp. faucium (western Alps, Switzerland, France, north-western Italy): upperside of wings less contrasted, underside with same pattern as other subspecies but on darker background; terminal line less continuous. Subsp. gedrensis (French Pyrenees): wings less contrasted, medial lines weakly serrated, terminal line straight; underside of wings darker; the two white lines thinner, terminal line usually interrupted. Male genitalia: Uncus triangular, slender, pointed; gnathos widely triangular with slender, rounded tip, at most reaching up to base of uncus. Valva broad; well sclerotized costa apically separated, distally curved, with numerous very long spines arranged in one or two rows. Juxta process slender, short, not reaching up to base of costa of valva, apically with few medially directed spines; margins of juxta process almost parallel; basally funnel-shaped elongated. A hyaline membrane connecting juxta with a long vase-shaped caulis, apically notched. Saccus digitiform projecting. Aedeagus straight, without spines. This latter character along with dentation of apex of juxta characteristic for this species. Female genitalia: Antrum funnel-shape, slender, short. Lamella postvaginalis with distinctly heart-shaped sclerite; ‘uncus gap’ widely opened without anteriorly extending fissure. Lamella antevaginalis developed as narrow, grooved band. Colliculum very short, membranous. Ductus bursae strongly sclerotized, merged with pear-shaped corpus bursae. Signum mouth-shaped. Distribution and abundance: European, alpine, with peri-alpine disjunction. Widely distributed in the Alps from eastern Alps (Hohe Tauern, eastern Tyrol) to Lechtal Alps (western Austria), from Ötztal Alps (Austria) to Rhaetian Alps (Switzerland, Italy), as well as in western Alps (Switzerland, Italy, France) (subsp. faucium) and in French Pyrenees (subsp. gedrensis). – Local, in limited numbers.

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Phenology: Univoltine. From early July to late August, mainly from mid-July to mid-August. Adults active mainly in the morning hours and then again, in small numbers, in the afternoon (S. Erlacher own observation) Biology: Larva polyphagous. Found on lichens and small plants (e.g. Saxifraga oppositi­ folia, Alchemilla xanthochlora, Taraxacum sp., Leontodon sp. and Fragaria sp.) (see Millière 1869–1874; Koch 1908; Lhomme 1923–1935; Thurner 1948; Leraut 2009; Flamigni et al. 2016). Larva overwintering twice, found under stones from early August to early September (Flamigni et al. 2016). Rearing from larva described by Wehrli & Imhoff (1922). Larva found at 2940 m altitude and reared to pupa by Ripper (1927). Egg similar to that of P. wehrlii, oval, light brown (Forster & Wohlfahrt 1981). Larva light brown with two white lateral lines and brown head (Wehrli & Imhoff 1922; Ripper 1927; Forster & Wohlfahrt 1981). Pupa shiny brown, cremaster with paired tapering processes (Ripper 1927; Forster & Wohlfahrt 1981). Habitat: Vertical distribution from 2000 up to 3500 m, exceptionally up to 3800 m (Großglockner, Austria). On rocky slopes with scarce vegetation and scree; on limestone and on primary rocks. Similar species: Upperside of wings of P. alticolaria potentially confused with that of P. noricana, the latter differing by absence of white lines on wing underside. Psodos baldensis and P. spitzi with white postmedial and terminal lines on wing underside but P. alticolaria not occurring sympatrically. In male genitalia juxta processes of P. alticolaria reminiscent of those of P. alpinata, the latter differing by broad and blunt uncus (in P. alticolaria more slender and considerably more pointed). Shape of antrum in P. alticolaria reminiscent of that of P. alpinata and P. coracina. In contrast to both last mentioned species, P. alticolaria with antrum more slender, disc of lamella antevaginalis heart-shaped sclerotized, ductus bursae larger.

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Genetic data: Two BINs: BOLD:ACE6515 (n=13 from Austria (Tyrol to Carinthia), Switzer­ land (Valais), France, north-western Italy), BOLD:AAH7951 (n=16 from northern Italy (Lombardy, South Tyrol), western Austria (Vorarlberg, Tyrol), Switzerland (Valais)). Genetically heterogeneous, maximum variation 4.6%, both BINs at a distance of 2.0%, requiring further integrative study based on more data. Nearest species: P. coracina (5.2%), P. baldensis (5.5%), P. spitzi (5.5%). Remarks: Zerny (1916) described P. chalybaeus as a species with a rounded postmedial line whereas the postmedial line is serrated in P. alticolaria. Vorbrodt (1918) confirmed these external differences, followed by Wehrli (1953). However, Povolny & Moucha (1955) already questioned the status of P. chalybaeus as a separate species claiming that P. chalybaeus seems to be only a form of P. alticolaria and that the differences given by Vorbrodt (1918) and Wehrli (1921) were “entirely unspecific” (Povolny & Moucha 1955: 163). In fact, there are no constant differences in the genitalia of both taxa and the coloration and wing pattern show high variability between the populations which often goes along with the geological substrates. Nevertheless, Wolfsberger (1966) and Scoble (1999) listed P. chalybaeus as species. Finally, Leraut (2009) and Flamigni et al. (2016) treated P. chalybaeus as a subspecies of P. alticolaria although there are some conflicts concerning wing pattern, geographic distribution and results of DNA barcoding especially in the contact zone of the two taxa in the Ötztal Alps (for explanations see Flamigni et al. 2016). Huemer (2013) suspects an edaphic adaptation, as occurs in many alpine species. Further investigations are necessary to clarify the status of the two taxa.

45. Psodos wehrlii Vorbrodt, 1918 Psodos wehrlii Vorbrodt, 1918: Int. Ent. Zeits. 12 (19): 148 (Switzerland, Wallis, Zermatt: Mettelhorn, 3400 m). Lectotype ♀ (hereby designated) (ZFMK, examined, label data: Switzerland, Wallis, Zermatt, Mettelhorn, [46°01‘ N, 07°45‘ E], 3400 m, 22.vii, gen. prep. CJ–110). – ­Paralectotypes 4♂8♀ (hereby designated) (ZFMK, examined, same data).

External characters and abdomen: Wingspan 23–27 mm. Upperside of wings sooty grey. Ante- and postmedial lines white, rounded. Terminal line continuous. Medial fascia dark brown. Basal and terminal areas lighter than rest. Antemedial line absent from hindwing. Medial and terminal lines visible. Underside of wings lighter. Terminal area with two more or less distinct light lines. Discal spots visible only from underside. Antennae dorsally chequered black and white. Variation: Medial area of forewing upperside usually dark but sometimes interrupted.

Male genitalia: Uncus broad, pointed. Gnathos triangular, pointed, reaching up to base of uncus. Valva ventrally broadened, rectangular. Costa of valva apically separated, distally curved, with numerous long and strongly sclerotized spines arranged in several rows. Juxta process characteristically broad, reaching up to base of costa of valva, distally curved, apically covered with a row of stout spines; basally funnel-shaped. Hyaline membrane connecting juxta with heart-shaped caulis, apically notched. Saccus digitiform. Aedeagus weakly arched, with tiny apical spines. Female genitalia: Antrum funnel-shaped, slender, short. Lamella postvaginalis almost rectangular, lateroventral part anteriorly bordered by straight line. Posterior margin of sclerite almost not arched, ‘uncus gap’ not developed. Lamella antevaginalis developed as narrow, grooved band. Colliculum, ductus bursae and corpus bursae not sclerotized. Signum absent. 151

Text-figs 68–71. Diagnostic characters (indicated) of selected European Psodos species. Upperside of wings. Text-fig. 68. P. alticolaria Mann, 1853, male. Austria: North Tyrol, Obergurgl, Rotmoostal, 1.viii.1994 (coll. Erlacher). Text-fig. 69. P. wehrlii Vorbrodt, 1918, male. Switzerland: Valais, Täsch Alp, 3.viii.1987 (coll. TLMF). Text-fig. 70. P. baldensis Wolfsberger, 1966, male. Italy: Verona, Monte Baldo, mid vii.1966 (TLMF), paratype. Text-fig. 71. P. spitzi Rebel, 1907, male. Slovenia: Triglav, 3.viii.1921 (coll. TLMF), cotype.

Distribution and abundance: Central alpine. Highly restricted to small distribution areas in the Alps. Earlier only known from Zermatt and Arolla in the Valais Alps (Switzerland) until Kitschelt (1935) recorded P. wehrlii in the Ortler mountains (northern Italy) (Flamigni et al. 2016). P. wehrlii seems to have disappeared from the Mettelhorn, the locus typicus, in the 152

Text-figs 72–75. Diagnostic characters (indicated) of selected European Psodos species. Underside of wings. Text-fig. 72. P. alticolaria Mann, 1853, male. Italy: South Tyrol, Stilfserjoch, 21.vii.1896 (NHMW), lectotype. Text-fig. 73. P. wehrlii Vorbrodt, 1918, male. Switzerland: Valais, Täsch Alp, 3.viii.1987 (coll. TLMF). Text-fig. 74. P. baldensis Wolfsberger, 1966, female. Italy: Verona, Monte Baldo, Cima Valdritta, mid vii.1969 (ZSM). Text-fig. 75. P. spitzi Rebel, 1907, female. Slovenia: Mount Kanin, 4.viii.1995 (TLMF).

last decades but it still occurs around Taesch (Zermatt) (M. Sommerer pers. comm.). – Highly local and in very limited numbers. Phenology: Univoltine. Flight period from July to mid-August. Biology: Larval food-plants unknown. Reared on Taraxacum (Flamigni et al. 2016). At flight localities of adults there are only low-growing plants, moss and lichens as potential larval food-plants. Under rearing conditions pupation between spun leaves of the food-plant. 153

Rearing to larval stage published by Wehrli & Imhoff (1922). Egg oval, colour turning from light yellow to reddish brown. Larva dark olive-grey, head black with two brownish lines, dorsal line black and interrupted, lateral line indistinct, bright and dark (Wehrli & Imhoff 1922; Forster & Wohlfahrt 1981). Pupa unknown. Habitat: Vertical distribution from 3000 up to 3800 m, P. wehrlii showing the highest distribution of all Psodos species. Inhabiting rocky islands in the middle of glaciers and “firn” fields (corn snow areas) (“Felsen und Granitinseln inmitten von Gletschern und ­Firnfeldern”, Wehrli 1953: 630). Similar species: Similar to P. alticolaria f. chalybaeus if transverse lines in P. wehrlii are not dentate. In male genitalia, P. wehrlii differing from all other Psodos species by ventrally broadened valva and broad juxta processes, and in female genitalia, by almost rectangular shape of lamella postvaginalis and absence of signum. Genetic data: BIN: BOLD:AAX7547 (n=1 from Switzerland). Nearest species: P. baldensis (3.9%), P. noricana (4.9%).

46. Psodos baldensis Wolfsberger, 1966 Psodos baldensis Wolfsberger, 1966: Estr. Mem. Mus. Civ. Storia Nat. Verona 14: 451, pl. 1, figs 1–4; pl. 2, figs 1, 2; pl. 3, figs 1, 2; text-fig. 1 (Italy, Trentino: Monte Baldo, 2100–2200 m). Holotype ♂ (MNV, not examined; Paratypes in ZSM, examined).

External characters and abdomen: Wingspan 19–24 mm. Upperside of wings dark brown, basal area yellow-brown. Transverse lines and discal spot very weak, but with three distinct silvery blue lines in terminal area of forewing (diagnostic). Underside with lighter yellow-­ brown basal field and two silvery blue lines. Discal spots present, shadowed with silvery blue scales. 154

Variation: Little. Male genitalia: Uncus triangular, pointed. Gnathos tongue-shaped with rounded tip, not reaching up to base of uncus. Costal process of valva slightly sclerotized and not separated, with few thin spines. Juxta process triangularly elongated, not reaching up to base of costa of valva, apically truncate and covered with numerous medially directed, triangular spines. Hyaline membrane connecting juxta to vase-shaped caulis, apically cup-shaped. Saccus digitiform, elongated. Aedeagus long, thin, slightly s-shaped, with numerous tiny apical spines. Female genitalia: Antrum funnel-shaped, extremely dilated towards ostium, narrowing towards ductus bursae. Lamella postvaginalis widely heart-shaped; uncus gap developed as deep notch. Lamella antevaginalis a wide, grooved band. Colliculum short, membranous; ductus bursae strongly sclerotized and bearing multiple folds, as large as membranous corpus bursae. Signum mouth-shaped. Distribution and abundance: Endemic to northern Italian Monte Baldo (Veneto) and to Bergamasque Alps (Lombardy) (Flamigni et al. 2016). Geographically isolated from all other Psodos species. – In the summit region of Monte Baldo not rare, but very difficult to catch, because almost exclusively flying on the extremely exposed north-west facing rock edges (S. Erlacher own observation).

Phenology: Univoltine. From mid-July to late July. Flight activity starting with sunrise and ending not later than noon, often at 10 o’clock in the morning (Wolfsberger 1966; S. ­Erlacher own observation). Biology: Larva most likely feeding on Potentilla nitida, adults repeatedly observed flying around that plant species. Egg, larva and pupa unknown. 155

Habitat: Vertical distribution from 2000 up to 2300 m. Habitat on the highest rocks at Monte Baldo with limited vegetation on limestone, in the area of Monte Demignone on arenaceous and porphyritic schist (Flamigni et al. 2016). Similar species: Dark brown wings and silvery blue lines on both sides of wings distinguish P. baldensis from all other Psodos species. Male genitalia of P. baldensis reminiscent of those of P. spitzi. However, identification of these two species easy based on basal elongation, and apical dentation of juxta, and form of aedeagus (slightly curved in P. spitzi and s-shaped in P. baldensis). For differences in female genitalia of P. baldensis and P. spitzi, see under P. spitzi. Genetic data: BIN: BOLD:AAL1649 (n=4 from Italy). Genetically homogeneous. Nearest species: P. spitzi (3.2%), P. wehrlii (3.9%).

47. Psodos spitzi Rebel, 1907 Psodos spitzi Rebel, 1907: Jahresb. Wien. Ent. Ver. 17: 54 (Slovenia: Triglav, 2400 m). Syntypes 1♂1♀ (NHMW, not examined).

External characters and abdomen: Wingspan 18–26 mm. Upperside of wings dark brown to dark grey. Pre- and postmedial lines serrated, terminal line almost continuous, edged with creamy scales. antemedial line absent from hindwing). Underside of wings dark brown with two thin, white lines in the terminal area, the distal line thinner, partially interrupted. Discal spot present on all wings, large on forewings. Variation: Little. Male genitalia: Uncus pointed. Gnathos broad, tongue-shaped, not reaching up to base of uncus. Costa of valva sclerotized, not separated from fore-edge of valva, with few thin bristles. Juxta process triangularly elongated, reaching up to costa of valva, apically with row of medially directed, triangular spines. Aedeagus weakly arched. Female genitalia: Female genitalia very similar to those of P. baldensis, but in P. spitzi funnel-­ shaped antrum clearly smaller and shorter. Lamella postvaginalis much smaller, latero­ ventrally swollen, ‘uncus gap’ very small, without anteriorly extending fissure. Signum much smaller. Distribution and abundance: Endemic to a small area from Dolomites to Carniolian and Julian Alps (north-eastern Italy, southern Austria, Slovenia: Huemer et al. 2009; Flamigni et al. 2016). – Very local, in limited numbers. Phenology: Univoltine. From July to August, mainly from mid-July to mid-August; one female recorded early September. Biology: Larva most likely feeding on Potentilla nitida judged from repeatedly observed adults flying around these plants (pers. observation). Larval stages unknown. Habitat: Vertical distribution from 2000 up to 2900 m. On rocks and scree on calcareous substrate. Similar species: Differing from P. baldensis in dentate postmedial line, in female genitalia by narrower and shorter antrum, narrower lamella postvaginalis, ‘uncus gap’ without anteriorly extending fissure.

156

Genetic data: BIN: BOLD:AAL1660 (n=3 from Italy and Slovenia). Genetically homogeneous. Nearest species: P. baldensis (3.2%), P. noricana (4.9%), P. alticolaria (5.5%).

48. Psodos quadrifaria (Sulzer, 1776) Phalaena quadrifaria Sulzer, 1776: Abgekürzte Geschichte der Insecten 1: 162; ibidem 2: pl. 23, fig. 4 (Alps, probably from Splügenberg, Graubünden, Switzerland (Wehrli 1953: 639). Phalaena equestrara Fabricius, 1776: Genera Insectorum: 288 (type locality unknown). Psodos alpinata pyrenaea Oberthür, 1884: Etudes ent., 8: 42 (France, Hautes-Pyrénées: Cirque de Gavarnie). Syntype(s), no type material mentioned in the original description. Validated at subspecies rank in Scoble (1999), here confirmed. Psodos quadrifaria [forma] stenotaenia Schwingenschuss, 1909: Verh. zool.-bot. Ges. Wien 66 (3–5): 330 (Austria, Glockner, Gemsgrube). Syntypes ♂♀ (NHMW, examined). Psodos quadrifaria sudetica Sterneck, 1918: Zeits. Österr. Ent.-Ver. 3 (7): 70 (Sudety mountains and Carpathians). Syntypes (NHMW, not examined). Validated at subspecies rank in Scoble (1999), here confirmed.

External characters and abdomen: Wingspan 20–26 mm. Upperside of wings dark brown, with yellow to orange, thick postmedial band, not reaching costa. Discal spot weak, almost invisible. Fringes brown. Dorsal scaling of antennae varying from black and white to brown or white. Variation: Little. See subspecies. Subsp. pyrenaea (Pyrenees): Slightly larger, orange postmedial band narrower than in nominotypical subspecies. Subsp. sudetica (Tatra and Sudety mountains): Orange postmedial band wider than in nominotypical subspecies.

157

Male genitalia: Uncus long, slender, apically pointed, thinner and longer than in all other Psodos species. Gnathos small and membranous. Costa of valva strongly sclerotized, medially pointed with spines. Cucullus apically round. Saccus digitiform. Juxta process slender, basally broadened, medially flattened, apically with a group of short spines. Aedeagus relatively straight, short, thick (thicker and shorter than in all other Psodos species). Vesica with long, numerous, strongly sclerotized spines. Male genitalia of P. quadrifaria clearly differing from all other Psodos species by membranous gnathos and strongly sclerotized spines on vesica. Female genitalia: Apophyses posteriores twice length of apophyses anteriores. Antrum reduced to small wide tongue-shaped structure (shorter than in all other Psodos species). Lamella postvaginalis lateroventrally very small, sclerite reduced, ‘uncus gap’ and lamella antevaginalis weakly developed. Ductus bursae slightly cone-shaped, longer than in all other Psodos species, strongly sclerotized, forming several folds. Corpus bursae membranous, signum oval to round. Female genitalia of P. quadrifaria differ from congenerics by reduced structures of lamella post- and antevaginalis. Distribution and abundance: European, alpine. Widely distributed in the European Alps, Massif Central of France and in Vosges mountains. In Pyrenees replaced by subsp. pyrenaea, in Tatra and Sudety mountains by subsp. sudetica.

Phenology: Univoltine. Flight period from mid-May to late August, with main flight period from early July to mid-July. Biology: Larva polyphagous. Found on Rhododendron spec. (Italy, South Tyrol: D. Fritsch pers. comm.). Recorded on low-growing plants, e.g. Lotus, Astragalus, Coronilla, Hippocrepis, Rhododendron, Taraxacum, Leontodon etc. (Redondo et al. 2009; Flamigni et al. 2016). Larva brownish with dark dorsal line and yellow lateral line (Prout 1915; Forster & Wohlfahrt 1981). Pupa light red with yellow, dark striped (Forster & Wohlfahrt 1981). 158

Habitat: Vertical distribution from 600 up to 3000 m, most specimens recorded from 900 up to 2400 m. Adults preferring habitats of high mountain meadows (Povolny & Moucha 1955). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAE8507 (n=16 from Germany, Austria, France, Italy, Croatia). Genetically heterogeneous, maximum variation 1.7%. Nearest species: Charissa obscurata (5.7%), Elophos caelibaria (5.8%), thus COI data not suggesting closer relationship to other Psodos species, for synonymy of genus names Psodos and Glacies, however, see also ­Remarks. Remarks: Leraut (2009: 242) validates ‘subsp. equestrara (Fabricius, 1777) (= sudetica Sterneck, 1918)’ for ‘populations extending from the Giant Mountains (southern Poland; northern Czech Republic) and from Germany to western Switzerland and France (Jura, Vosges, Massif Central, Haute-Savoie, Isère, Savoie, Pyrénées-Orientales)’ while considering nominotypical subspecies to be restricted to “the rest of French Alps”. Not accepted here because of lack of knowledge about type locality for equestrara and supposed type locality of nominotypical subspecies (Switzerland: Graubünden). Sterneck (1918: 71) and Schwingens­ chuss (1923) have considered the taxon stenotaenia Schwingenschuss, 1909 (with particularly narrow, orange postmedial band) to be “a hereditary race”. Presumed hybrid of Psodos quadrifaria and P. alpinata described by Wehrli (1920), supporting synonymy of genus names Psodos and Glacies.

Sciadia Hübner, 1822 Sciadia Hübner, 1822, Syst.-Alphab. Verz.: 40–46, 48–51. Type species: Geometra torvaria H ­ übner, 1813. [Sciadia was the spelling adopted by Hemming, 1937 (the principle of first reviser), from a multiple original spelling in which both Sciadia and ‡ Scidia were used on page 42 (see Fletcher 1979)]. ‡ Sciadia Hübner, 1818, Zuträge Samml. exot. Schmett. 1: 27, [34]. [From Fletcher 1979 (p. 186): ‘Not nomenclaturally available from 1818 as a genus-group name. Hübner used Sciadia on page 27 in a suprageneric sense for one of the names of this tribes (Namen der Stämme) listed on pages [33], [34]. Hübner listed his genus-group names on pages [35], [36]. ‡ Scidia Hübner, 1822, Syst.-Alphab. Verz.: 42. [An incorrect original spelling]. Orphne Hübner, 1825, Verz. bekannter Schmett.: 314. Type species: Geometra torvaria Hübner, 1813. Dasydia Guenée, 1845, in Duponchel, Cat. méth. Lépid. Eur. 281. Type species: Geometra torvaria Hübner, 1813.

Diversity and distribution: The revised concept of Sciadia, as introduced here, contains seven species in the West Palaearctic region, the occurrence is restricted to the South European mountains in the Alps, Pyrenees and Dinaric Alps. Sciadia was for a long time considered a monotypic genus, containing diurnal S. tenebraria Esper only, until it was discovered recently to contain five species (Leraut 2008; Huemer & Hausmann 2009). We combine two additional species to Sciadia but more species from the related genera may need to be transferred to Sciadia, see Remarks. External characters and abdomen: Wingspan of males approximately 26–35 mm, females approximately 21–28 mm. Females smaller, forewings more rounded. Genus contains two species groups: in S. tenebraria group wings are rather uniform dark brown with greyish 159

tint. Medial and postmedial lines dentate, bordering slightly darker medial area. Discal spots distinct. Underside of wings characterized by whitish to yellowish, curved subterminal line, otherwise wings brown. In S. zelleraria group wings are grey-brown, markings light brown. Underside of wing with large discal spots and terminal area with dark brown band. Male antennae bipectinate, female antennae filiform. Hindlegs of both sexes with 2+2 spurs. Male tympanal organs medium-sized, female organs very small. Male 8th sternite small, about half the size of 8th tergite. Male genitalia: Uncus triangular, apex blunt. Socii absent. Gnathos with small lateral sclerites and granulate medial plate. Valva wide at base, apex slightly upturned. Costa base with few stout setae, apex setose. Posterior arms of juxta wide lobes, simple, apex smooth. Anterior arms of juxta short, triangular. Vesica with numerous small, spine-like cornuti, one large horn-shaped cornutus present or absent. Posterior arms of juxta offer species-level diagnostic characters, see genitalia plates. Female genitalia: Papillae anales rounded, setose. Margin of lamella antevaginalis concave. Ductus bursae wide, sclerotized. Corpus bursae wrinkled anteriorly. Signum very small, stellate, located at anterior end or absent. Biology, habitat and phenology: Due to the recent, updated species taxonomy of the S. tenebraria species group (Leraut 2008; Huemer & Hausmann 2009), it is often impossible to know to which taxon the literature data on S. tenebraria refers. The assumed larva of S. tenebraria lives under stones, feeding on herbs such as Saxifraga caesia, S. oppositifolia, Silene acaulis, Ranunculus glacialis and on lichens and mosses (cryptogams) (Vorbrodt & Müller-Rutz 1914; Huemer & Hausmann 2009). Probably the larval development of S. tenebraria requires two years. Larva of S. zelleraria is polyphagous on low alpine plants, potentially on Pimpinella and Campanula, in captivity it has fed on Taraxacum and Lactuca (Flamigni et al. 2016). All species are restricted to rocky habitats and alpine scree above the treeline, ranging roughly from 1500 m up to 3300 m. Species in S. tenebraria group are diurnal, flying only in full sunshine. Males are more active, but both sexes visit cushion plants such as Silene acaulis (Huemer & Hausmann 2009). Species in S. zelleraria group are nocturnal and attracted to light, females of S. zelleraria have rarely been seen visiting flowers of Silene acaulis during daytime. Immature stages: Due to the recent, updated species taxonomy of the S. tenebraria species group (Leraut 2008; Huemer & Hausmann 2009), it is often impossible to know to which taxon the literature data on Sciadia refers to. The assumed larva of Sciadia tenebraria is brown-grey, with partially interrupted beige longitudinal stripes. Each segment laterally has small, distinct red-brown spot. Head is brown, with blackish mottling (Ziegler 2019). The assumed pupa of S. tenebraria has cremaster with four pairs of curved setae, apical pair is longest (Patočka & Turčani 2005). Remarks: The genus-level relationships of structurally, and partly ecologically, similar Elophos Boisduval, 1840, Sciadia Hübner, 1822, Psodos Treitschke, 1825 and potentially other related genera need further research because the relationships have not been investigated in an extensive analytical context so far. The close relationship of Sciadia and S. zelleraria group (transferred from Elophos to Sciadia here) is supported by similar morphology, and by DNA barcodes also. S. zelleraria share or almost share their barcodes with S. tenebraria, S. innuptaria, S. dolomitica and S. slovenica and Bavarian vouchers from southern Germany share barcodes with S. tenebraria from Austria, Italy and Switzerland (Hausmann et al. 2011b). Further, DNA barcodes of S. unicoloraria (transferred from Elophos here) are tangled with S. slovenica and S. tenebraria (Mutanen et al. 2016). As regards S. tenebraria, 160

Mutanen et al. (2016, Supplementary Table 1) wrote: ‘Model case of likely introgression with several species of Elophos’. The barcode sharing of Sciadia zelleraria (as Elophos zelleraria) and Sciadia tenebraria wockearia was noted also by Huemer & Hebert (2016). It is worth noting that despite externally different facies, the male and female genitalia of S. tenebraria and S. zelleraria are similar, without diagnostic differences (our observation). Hausmann et al. (2011b) suggested to unite all these species pairs with supposed close relationship into one single genus. The view is agreed as regards S. zelleraria species group, which is morphologically very similar to Sciadia. It is shown in this book that Elophos includes several, morphologically distinct species groups (diagnosable particularly by the genitalia, see the relevant male and female plates) and it is premature to transfer this hetero­ geneity into a formal classification that contains a single genus only. Genus Sciadia, as introduced here, is morphologically, ecologically and genetically a uniform group but its relationships to other taxa in wider context need further research. We retain several species in genus Elophos, following earlier classifications, but note that when extensive analytical evidence is provided, reclassification is likely to be needed. One of these may be Elophos caelibaria (Heydenreich, 1851), which shares barcodes with two Sciadia specimens (S. tenebraria and S. slovenica) from Italy and Slovenia (Hausmann et al. 2011b). Elophos caelibaria and the closely related E. zirbitzensis are structurally different from the core Sciadia, how­ ever. Another example of close relationship is that of S. slovenica, which is polyphyletic and DNA barcodes are tangled with several Sciadia and Elophos species (Mutanen et al. 2016). Recently Sciadia has been classified in the Gnophini (Urbahn 1967; Vives Moreno 1994; Leraut 1997) and in the Boarmiini (Hausmann et al. 2004; 2011a). Sciadia is structurally similar to numerous Gnophini genera and it shares a number of Gnophini characters as defined by Viidalepp et al. (2007) and as illustrated in this book. Systematic position: ­Gnophini.

49. Sciadia tenebraria (Esper, 1806) Geom[etra] tenebraria Esper, 1806: Die Schmett. 5: plate 51, fig. 6 (Europe). Neotype ♂ (designated by Huemer & Hausmann 2009, [southern Germany] Allgäu, Mädelegabel, 2400– 2600 m, (ZSM), DNA barcoded. Geometra torvaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 71, figs 366–369 (Europe). Neotype ♂ (designated by Huemer & Hausmann 2009, [Austria] Tirol, Innsbruck Saile, ZSM), DNA barcoded. Junior synonym. Dasydia tenebraria var. wockearia Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 169. Syntypes, Italy: Southern Tyrol, Trafoi. Valid at subspecific rank (Huemer & Hausmann 2009). Parascotia olivacea Warren, 1895: Novit. zool. 2: 129 (India [north-west]: Spiti [incorrect locality]). Holotype ♂ (NHMUK, examined externally). Junior synonym. Sciadia sabaudiensis Leraut, 2008: Bull. Soc. entomol. France 113 (2): 181, figs 16–19, 24, 28 (France: Savoie, Pralognan, Petit-Mont-Blanc). Holotype male (MNHN), paratypes 3♂4♀ (MNHN, examined externally, based on photos in Leraut (2008), one paratype DNA bar­ coded). Junior synonym, synonymy established by Hausmann & Huemer (2009). Sciadia tenebraria taurusica Huemer & Hausmann, 2009: Zootaxa 2117: 19 (Austria: Osttirol, Gross­ glockner, 2700 m). Holotype ♂ (TLMF). Valid at subspecific rank (Huemer & Hausmann 2009). Unavailable names (infrasubspecific): interrupta: Wehrli (1920) (f.); vernagtensis: Schawerda (1933) (ab.).

External characters and abdomen: Wingspan 22–31 mm in males, 24–28 mm in females. Females often slightly smaller, more narrow-winged and apex of forewing rounded. Wings 161

dark greyish brown, weakly glossy. Medial area darker, bordered by dentate antemedial and postmedial lines, the latter bordered by slightly paler margin. Terminal line dark brown. Fringes concolorous with wings or slightly paler, unicolorous. Large, dark brown discal spots present on fore- and hindwings. Underside of wings dark brown, with well developed though diffuse beige fascia, curved inwards on forewing costa, and often slightly narrower on hindwings (Text-fig. 76). Discal spots weaker or absent on underside. Frons, collar, thorax and abdomen concolorous with wings. Thorax and basal segments of legs covered with long hairy scales. Female abdomen considerably larger than male. Male antennae bipectinate, female antennae filiform. Hindtibia with 2+2 spurs in both sexes. Male tympanal organs medium-sized, female organs very small. Male 8th sternite slightly smaller than 8th tergite. Other sternites and tergites of both sexes unmodified. Variation: Pale brown fascia on underside can be narrow or wide, usually diffuse. Male genitalia: Uncus large, triangular, apex round. Socii absent. Gnathos with small lateral sclerites, granulate medial plate indistinct. Valva wide and short, basal part rough, costa sclerotized and with several stout setae basally, subcostal area setose. Posterior arms of juxta short, rather wide, apex round, inner margin irregular (rough). Anterior arms of juxta hornshaped. Saccus elongated, roundish. Aedeagus stout, slightly curved. Vesica opens at about 90 degree angle, evenly curved, with 10–25 cornuti of various lengths. Ductus ejaculatorius open from vesica apex. Apex of aedeagus with additional, sclerotized and dentate cornutus. There is tendency to a reduction of the additional cornutus towards western Alps, the structure is absent in populations from Switzerland: Graubünden and Wallis, and the ventral lobes of juxta are usually shorter in western populations (Huemer & Hausmann 2009). Female genitalia: Papillae anales rounded, setose. Margin of lamella antevaginalis concave. Ductus bursae sclerotized, narrower at anterior end. Posterior part of corpus bursae wrinkled, medial part narrow, anterior part roundish. Signum small, stellate, located at anterior part, sometime reduced or absent. There is tendency to a reduction of the signum towards western Alps, the structure is absent in populations from Switzerland: Graubünden and Wallis (Huemer & Hausmann 2009). Subsp. wockearia has strongly contrasting, yellowish or even slightly golden, sharp-edged (not diffuse) and narrow fascia on the underside of the wings (Text-fig. 77). Fascia is strongly curved inwards on costa and apex of forewing has additional yellowish area. Additional cornutus varying from dentate plate to medium-sized to small spine, signum small to more or less absent (Huemer & Hausmann 2009). Taxon is found in a small area in southern Alps (Austria, Switzerland, Italy), ranging from the Ötztaler mountains in the north-east to the Ortler mountains, Adamello and Alpi Orobie in the south-west (Huemer & Hausmann 2009). Subsp. taurusica is without fasciae on the underside of the wings (Text-fig. 78). Additional cornutus varying from small dentate plate to small spine, minute signum is always present (Huemer & Hausmann 2009). Taxon is found in a small area in the central Alps of Austria (Hohe Tauern, Niedere Tauern), extending to the main ridge of the Alps in the north (­Huemer & Hausmann 2009). Distribution and abundance: European, alpine. Widely distributed in the Alps in Germany, Austria, Switzerland, Liechtenstein, France and Italy. According to Huemer & Hausmann (2009), the species has an isolated record in eastern Austria in Hochschwab mountains, where it occurs sympatrically with S. innuptaria. – A very locally occurring species that can be abundant in its localities. 162

Phenology: Univoltine. From late June to late August. Adults active during daytime, flying in sunshine, visiting flowers of Silene acaulis or other plants and settling on the damp ground (Flamigni et al. 2016). In poor or windy weather the moths take shelter under stones. Larval development in S. tenebraria group reputedly lasting two years or more, awaiting confirmation (Huemer & Hausmann 2009). Biology: Larva mono- or oligophagous. Recorded on Saxifraga caesia, S. oppositifolia, ­ ilene acaulis, Ranunculus glacialis, Ranunculus parnassifolius and lichens, however, all S these data awaiting verification due to recent taxonomic revision of the group (Huemer & Hausmann 2009; Flamigni et al. 2016) and unknown attribution of literature data to the different taxa in the S. tenebraria group. Adults of S. tenebraria observed feeding on flowers of Noccaea cepaeifolia (P. Skou own data). Habitat: Alpine. Old records need verification. In Italy found in rocky habitats, often with loose debris and with sparse vegetation above the treeline, on both limestone and siliceous substrates. Rarely in open alpine meadows (Flamigni et al. 2016). In Italy from 2300 m up to 3300 m, the lowest record being from 1050 m, in Switzerland from 1800 m up to 4000 m (Flamigni et al. 2016). Similar species: All species in the Sciadia tenebraria species complex are externally similar, therefore we recommend that genitalia and/or DNA barcodes are used for identification. Reliable identifications will help to understand the true distributions and life histories of these species. Both male and female genitalia structures allow reliable identification. ­Sciadia tenebraria and S. septaria are similar. Male of S. tenebraria has posterior arms of juxta with round apex, inner margin is irregular, additional cornutus is present [exception: in Switzer­ land additional cornutus absent] (posterior arms of juxta with acute apex, inner margin is rather straight and additional cornutus is absent in S. septaria). Female of S. tenebraria has heavily wrinkled posterior part of the corpus bursae, large antrum and signum is present 163

Text-figs 76–78. Diagnostic characters (indicated) of three subspecies of Sciadia tenebraria (­Esper, 1806), all specimens shown from underside. Text-fig. 76. S. tenebraria tenebraria male. Austria: Nordkette, 14.viii.1970 (coll. TLMF). Text-fig. 77. S. tenebraria wockearia male. Italy: Adamello, Rif. Mandron, 2900 m, viii.1958 (coll. TLMF). Text-fig. 78. S. tenebraria taurusica male. Austria: Kärnten, Glockner-Hochtor, 2600 m, 9.vii.1988 (ex pupa), paratype (coll. TLMF).

[exception: in Switzerland signum is absent] (less wrinkled posterior part of the corpus ­bursae, antrum is small and signum is absent in S. septaria). According to current knowledge the distribution areas do not overlap, S. tenebraria occurs in the Alps, S. septaria in the Pyrenees. Despite different external appearances, male and female genitalia of S. tenebraria and S. zelleraria are similar, without diagnostic differences. 164

Genetic data: Four BINs: BOLD:ACE3561, BOLD:ACE3562, BOLD:ACE3563, BOLD:AAB4956 (n=28 from Germany, Austria, France, Italy). These four BINs clustering together at distances of 1.2–2.7%, with just two cases of introgression into BOLD:AAB4956 (by specimens of S. slovenica). Nearest species: S. septaria (1.5% distance from French populations of S. tenebraria), S. unicoloraria (1.3% distance from south-western Swiss populations of S. tenebraria). S. tenebraria frequently (n=17) showing mitochondrial introgression into other species, see genetic data under S. zelleraria and E. caelibaria. See also Huemer & Hebert (2016).

50. Sciadia septaria (Guenée, 1858) Dasydia septaria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 316 (Pyrenees?). Holotype male (ZFMK, examined externally).

External characters and abdomen: Wingspan 27–29 mm in males, 22–23 mm in females (S. septaria smallest species in S. tenebraria species group), more narrow-winged and apex of forewing rounded. Similar to S. tenebraria, but, with following differences: male wings almost unicolorous dark brown, medial area almost same colour as rest of wing (medial fascia darker than rest of wing in S. tenebraria). Underside of wings with barely visible, narrow and diffuse beige fascia (underside with well developed though diffuse beige fascia in S. ­tenebraria). ­Female greyish brown, small, approaching brachyptery, underside of wings light brown except terminal area (female dark greyish brown, with larger wings, underside with narrow and diffuse beige fascia in S. tenebraria). Other characters generally as in S. tenebraria. Variation: All examined specimens are similar (n=4). Male genitalia: Generally as in S. tenebraria, but with following differences: genitalia capsule small, posterior arms of juxta with acute apex, inner margin is rather straight and additional cornutus is absent in S. septaria (genitalia capsule large, posterior arms of juxta with round apex, inner margin is irregular, additional cornutus present [exception: in Switzerland additional cornutus absent] in S. tenebraria). Female genitalia: Generally as in S. tenebraria, but with following differences: posterior part of the corpus bursae is slightly wrinkled, antrum is small and signum is absent in S. septaria (posterior part of the corpus bursae is heavily wrinkled, antrum is large and signum is present [exception: in Switzerland signum is absent] in S. tenebraria). Distribution and abundance: European. Only known from the Pyrenees in France, Spain and Andorra. According to current knowledge S. septaria has an allopatric distribution, not occurring sympatrically with any other taxon in the S. tenebraria species group. – Very local and rare, especially females, being almost absent from museum collections. Phenology: Univoltine: Scarce data from late June to early August. Most likely the flight period is similar to other Sciadia. Males active by day in sunshine, females rarely seen. Biology: Unknown. Habitat: In open habitats above treeline, from 2700 m up to 3000 m. Similar species: All species in the S. tenebraria species complex are externally similar, therefore we recommend that genitalia and/or DNA barcodes are used for identification. Reliable identifications will help to understand the true distributions and life histories of these species.

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Both male and female genitalia structures allow reliable identification. S. tenebraria and S. septaria are similar, see text under S. tenebraria. Genetic data: BIN: BOLD:AAI0834 (n=2 from central Pyrenees). Nearest species: S. tenebraria from south-eastern France (1.5%). No introgression observed, so far.

51. Sciadia innuptaria (Herrich-Schäffer, 1852) Gnophos innuptaria Herrich-Schäffer, 1851: Syst. Bearb. Schmett. Eur. 3: pl. 83, fig. 508 [non binominal], 1852 (6 (55)): 73 (Europe). Lectotype ♀ (designated by Huemer & Hausmann 2009, [Austria, Grossglockner region?], gen. prep. Hausmann G 12010, MNHU). See Huemer & Hausmann (2009) for details about the type specimen(s).

External characters and abdomen: Wingspan 28–30 mm in males, 24–29 mm in females. Generally as S. tenebraria (see description above), but females are about same size as males, forewing apex is pointed (not rounded), underside fascia weaker and wings glossy. Most similar species are S. slovenica and S. dolomitica, and due to absence of clear diagnostic external features examination of the genitalia is recommended. Variation: All examined specimens are similar (n=6). Male genitalia: Generally as in S. tenebraria, most similar species are S. slovenica and S. dolomitica: posterior arms of juxta with round apex, cornuti large in S. innuptaria (posterior arms of juxta with rather pointing apex, cornuti small in S. slovenica and S. dolomitica). Female genitalia: Most similar species are S. slovenica and S. dolomitica: sclerotization at posterior part of corpus bursae rather round, covering anterior part of ductus bursae in S. innuptaria (sclerotization with elongation on right side (when viewed ventrally) in S. slovenica, sclerotization not covering anterior part of ductus bursae in S. dolomitica). 166

Distribution and abundance: European, alpine. Only known from the north-eastern Alps in Austria. According to current knowledge S. innuptaria occurs sympatrically with S. tenebraria in eastern Alps in Austria. – Local and rare, occasionally occurring in numbers. Phenology: Univoltine: From late June to mid-August, most records from mid- and late July. Larval development in the S. tenebraria group reputedly lasting at least two years or more, awaiting confirmation for S. innuptaria (Huemer & Hausmann 2009). Males active in sunshine, females rarely seen. Biology: Unknown. Larva of S. tenebraria group recorded on Saxifraga caesia, S. oppositifolia, Silene acaulis and Ranunculus glacialis, however, all data awaiting verification and exact assignment to species after recent taxonomic revision of the group (Huemer & Hausmann 2009). Habitat: Alpine. From 1700 m up to 2100 m, occasionally even lower. Sciadia innuptaria occurs at lower altitudes than other species in the S. tenebraria group (Huemer & Hausmann 2009). Similar species: All species in the Sciadia tenebraria species complex are externally similar, therefore we recommend that genitalia and/or DNA barcodes are used for identification. Reliable identifications will help to understand the true distributions and life histories of these species. Both male and female genitalia structures allow reliable identification. Sciadia innuptaria has posterior arms of juxta with rounded apex, juxta arms are rather close to each other (parallel, not v-shaped) and cornuti are large, posterolateral parts of antrum are weakly sclerotized, posterior part of corpus bursae is weakly wrinkled and sclerotization in the posterior part of corpus bursae is rather round, covering anterior part of ductus bursae. Genetic data: BIN: BOLD:AAB4957 (n=3 from central Austria). Genetically homogeneous. Nearest species: S. zelleraria (1.2%), BIN-sharing with the latter. The whole population of 167

S. innuptaria probably performed a short, separate evolutionary history after introgression into S. zelleraria whilst in the 13 observed cases of introgression of S. tenebraria into S. zelleraria their mitochondria remained exactly barcode-sharing with the latter.

52. Sciadia slovenica Leraut, 2008 Sciadia slovenica Leraut, 2008: Bull. Soc. ent. France 113 (2): 179, figs 10, 11, 22 (Slovenia: Triglav mountain, 2500 m. Holotype ♂ (MNHN, examined externally, based on photos in Leraut (2008)).

External characters and abdomen: Wingspan 28–30 mm in males, 26–28 mm in females. Generally as S. tenebraria (see description above), with weak diffuse fasciae on the underside of the wings. Most similar species are S. innuptaria and particularly S. dolomitica, and due to absence of clear diagnostic external features examination of the genitalia is recommended. Variation: All examined specimens are rather similar. Male genitalia: Generally as in S. tenebraria, most similar species are S. innuptaria and particularly S. dolomitica: posterior arms of juxta with with rather pointed apex, arms v-shaped, not extending beyond margin of costa in S. slovenica (posterior arms of juxta with rather pointed apex, arms v-shaped and narrow, extending slightly beyond margin of costa in S. dolomitica). In S. innuptaria posterior arms of juxta with rounded apex, juxta arms close to each other (parallel, not v-shaped), cornuti large. Female genitalia: Most similar species are S. innuptaria and S. dolomitica: sclerotization at posterior part of corpus bursae with elongation on right side (when viewed ventrally) in S. slovenica (sclerotization at posterior part of corpus bursae round, covering anterior part of ductus bursae in S. innuptaria and sclerotization not covering anterior part of ductus bursae in S. dolomitica). Distribution and abundance: European, alpine. Only known from the eastern Alps (Julian Alps, Kamnik/Steiner Alps) in Slovenia, Austria and Italy. A doubtful record from Albania possibly referring to a separate taxon (Huemer & Hausmann 2009). According to current knowledge S. slovenica not occurring sympatrically with any other taxon in the S. tene­ braria species group. − Local and rare, usually found in limited numbers. Phenology: Univoltine: From early July to late August, in Italy recorded between 6th of July and 23rd of August (Flamigni et al. 2016). Larval development in the S. tenebraria group possibly lasting two years or more, awaiting confirmation for this species (Huemer & Hausmann 2009). Males active in sunshine, females rarely seen. Biology: Unknown. Larva of the S. tenebraria group recorded on Saxifraga caesia, S. oppo­ sitifolia, Silene acaulis and Ranunculus glacialis, however, all data awaiting verification and exact assignment to species after recent taxonomic revision of the group (Huemer & Hausmann 2009). Habitat: Alpine. Open rocky habitats, including rock debris at the foot of a rocky wall or steep slope, with sparse vegetation above treeline and only on calcareous soil. From 2000 m up to 2500 m (Flamigni et al. 2016).

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Similar species: All species in the Sciadia tenebraria species complex are externally similar, therefore we recommend that genitalia and/or DNA barcodes are used for identification. Reliable identifications will help to understand the true distributions and life histories of these species. Both male and female genitalia structures allow reliable identification. S. slovenica has posterior arms of juxta with with rather pointed apex, arms v-shaped, not extending beyond margin of costa and sclerotization at posterior part of corpus bursae is with an elongation on the right side (in ventral view). Genetic data: Three BINs: BOLD:AAC5403, BOLD:AAD2991, BOLD:AAB4956 (n=5 from Slovenia), apparently the BINs of all examined specimens being a product of introgression into other species (Elophos caelibaria, Sciadia zelleraria, S. tenebraria).

53. Sciadia dolomitica Huemer & Hausmann, 2009 Sciadia dolomitica Huemer & Hausmann, 2009: Zootaxa 2117: 24, figs. 29–32, 40, 48 (Italy: Ter. merid. Boespitze 3000 m). Holotype ♂ (TLMF, examined, including genitalia), paratype DNA barcoded.

External characters and abdomen: Wingspan 25–30 mm in males, 26–29 mm in females. Generally as S. tenebraria (see description above), but S. dolomitica is often more blackish. Most similar species are S. innuptaria and particularly S. slovenica, and due to absence of clear diagnostic external features examination of the genitalia is recommended. Sciadia dolomitica is often dark brown to blackish, compared to other species in S. tenebraria species group that are more brown. Variation: All examined specimens are rather similar. Male genitalia: Generally as in S. tenebraria, most similar species are S. innuptaria and particularly S. slovenica: posterior arms of juxta with pointed apex, arms v-shaped, extending 169

slightly beyond margin of costa and vesica with numerous small cornuti in S. dolomitica (posterior arms of juxta with with rather pointed apex, arms v-shaped, not extending beyond margin of costa and vesica with fewer small cornuti in S. slovenica). In S. innuptaria posterior arms of juxta with rounded apex, juxta arms close to each other (parallel, not v-shaped), cornuti large. Female genitalia: Most similar species are S. innuptaria and S. slovenica: sclerotization at posterior part of corpus bursae not covering anterior part of ductus bursae in S. dolomitica (sclerotization at posterior part of corpus bursae round, covering anterior part of ductus bursae in S. innuptaria and sclerotization with elongation on right side (when viewed ventrally) in S. slovenica. Distribution and abundance: European, alpine. Only known from the southern Alps in Italy, in the Dolomites. Confirmed records confined to Trentino-Alto Adige (Trento and Bolzano) and Veneto (Belluno). See Flamigni et al. (2016) for exact localities; one of these – Le ­Tofane, Rifugio Tiussani in Belluno – being erroneous and referring to S. tenebraria (C. Flamigni pers. comm.). Earlier records (before description of S. dolomitica) from the Dolomites reported as ‘S. tenebraria’. – Local and rare, occasionally occurring in numbers.

Phenology: Univoltine: From mid-July to early August. Larval development in the S. tene­ braria group possibly lasting two years or more, awaiting confirmation for this species (­Huemer & Hausmann 2009). Males active in sunshine, females rarely seen. Biology: Unknown. Larva of the S. tenebraria group recorded on Saxifraga caesia, S. oppo­ sitifolia, Silene acaulis and Ranunculus glacialis, however, all data awaiting verification and exact assignment to species after recent taxonomic revision of the group (Huemer & Hausmann 2009). 170

Habitat: Alpine. Open rocky habitats with sparse vegetation above treeline, only on calcareous soil. From 2300 m up to 3000 m, one specimen collected at 2000 m (Flamigni et al. 2016). Similar species: All species in the Sciadia tenebraria species complex are externally similar, therefore we recommend that genitalia and/or DNA barcodes are used for identification. Reliable identifications will help to understand the true distributions and life histories of these species. Both male and female genitalia structures allow reliable identification. S. dolomitica are often dark brown to blackish, posterior arms of juxta are with pointed apex, arms v-shaped, extending slightly beyond margin of costa and vesica is with numerous small cornuti. Further, sclerotization at posterior part of corpus bursae is not covering anterior part of the ductus bursae. Genetic data: BIN: BOLD:ACE3560 (n=2 from northern Italy, Dolomite Mts.), BIN-sharing with S. zelleraria, but slightly diverging (1.0%).

Sciadia zelleraria – S. unicoloraria species pair Sciadia zelleraria (Freyer, 1836), comb. n. and S. unicoloraria (Staudinger, 1871), comb. n. form a taxonomically difficult species pair that lacks distinct external diagnostic features, both species vary geographically, and according to current knowledge they do not occur sympatrically. Sciadia zelleraria has more eastern distribution area and S. unicoloraria more western, apparently coming closest to each other in Switzerland (Zermatt in West and Umbrail Pass in East, which are about 200 km apart). Short-winged females are rarely seen and those are very scarce in collections, actually the female genitalia of S. unicoloraria was illustrated for the first time very recently (Flamigni et al. 2016), who concluded that female genitalia of these two species are without diagnostic differences. DNA barcodes have not clarified the issue either because S. zelleraria is genetically heterogeneous (maximum intraspecific variation is 5.61%) and it is paraphyletic (tangled with) in relation to S. unicoloraria, S. dolomitica, S. septaria, S. slovenica and S. tenebraria (Mutanen et al. 2016). Based on the material examined, we conclude that S. zelleraria and S. unicoloraria are valid at species level. External appearance of the most typical specimens of each species are diagnostic, see Text-figs 79–80, but both species vary a lot and the appearance is somewhat dependent on the collecting locality and many intermediate specimens exist. This may be anticipated because both species live in mountain habitats and migration abilities of shortwinged females are restricted, potentially resulting in poor gene flow between populations and phenotypic differences over time. Male genitalia allow unambiguous species-level identification. Only two female genitalia of S. unicoloraria have been studied, therefore the observed presence/absence of signum character awaits confirmation. Due to difficulties in identification, literature on both species may be confusing and distribution records must be treated with caution. More research is needed, particularly it would be important to find out if these two species co-occur in sympatry in any locality. Selected diagnostic characters are shown in Table 2 and in Text-figs 79–80.

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Text-figs 79–80. Diagnostic characters (indicated) of Sciadia zelleraria (Freyer, 1836) and S. unicoloraria (Staudinger, 1871) males from above (left column) and from below (right column). Many specimens in this species group cannot be identified on the basis of external characters alone because external characters (density of patterns and wing colour) vary and correlate to some extent with the collecting locality. Examination of the genitalia is recommended for identification, whereas the DNA barcodes are problematic, see text. Wings of these species are shiny and reflections make them difficult to photograph. Text-fig. 79. S. zelleraria male. Italy: Völs am Schlern, 27.vii.2007, 2300–2440 m (coll. TLMF). Text-fig. 80. S. unicoloraria male. France: Punta Marguareis, 21–23.vii.1990, 2100–2200 m (coll. TLMF). Table 2. Selected diagnostic genitalia and distribution characters of Sciadia zelleraria (­Freyer, 1836) and S. unicoloraria (Staudinger, 1871). Character

S. zelleraria

S. unicoloraria

aedeagus + vesica max. width juxta cornutus at base of vesica sclerotization at apex of aedeagus vesica opens female signum

1.0–1.3 mm inner margin rough large does not reach apex at about 90° angle present, minute (n=3) or absent (n=1) from eastern Switzerland (Umbrail Pass) to the east

0.6–0.8 mm inner margin smooth absent reaches apex at about 45° angle absent (n=2)

distribution

from south-western Switzerland (­Zermatt) to the south-west

54. Sciadia zelleraria (Freyer, 1836), comb. n. Gnophos zelleraria Freyer, 1836: Neuere Beitr. Schmettkde 2 (32): 154, pl. 192, figs 2, 3 (Slovenia: Steiner Alps near Ljubljana). Syntype(s). Transferred here from Elophos Boisduval, 1840 (for instance Scoble 1999; Flamigni et al. 2016) to Sciadia Hübner, 1822, quoting structural similarities (comb. n.). Gnophos zelleraria f. robusta Wehrli, 1922: Societas ent. 37 (5): 18 (Austria: Tirol). Syntype(s) ♂ (ZFMK and paratype ♂ in NHMUK, examined externally). Treated valid at subspecific

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rank recently (for instance Scoble 1999; Flamigni et al. 2016), downgraded here (syn. n.), based on absence of constant differential features, falling within intraspecific variation. See Remarks. Gnophos zelleraria teriolensis Müller, 1927: Z. öst. EntVer. 12 (12): 122 (Austria, Tyrol: Ötztal). Holotype. Junior synonym of robusta according to Scoble (1999) and Flamigni et al. (2016), here confirmed. Unavailable names (infrasubspecific): designata: Feichtenberger (1962) (ab.); grisea: Feichtenberger (1962) (ab.); lutea: Feichtenberger (1962) (ab.); reducta: Müller (1927) (ab.); s ­ ignata: Feichtenberger (1962).

External characters and abdomen: Male: wingspan 30–34 mm (note: measured from apex to apex; Flamigni et al. (2016) give wingspan 35–43 mm, potentially measured from the maximum distance between wing margins, below the apex). Wing coloration in male glossy, yellowish grey, heavily irrorated with grey, markings grey-brown. Antemedial line absent. Medial line rather straight, weakly turned inwards on costa, wider on costa and inner margin. Postmedial line weakly dentate, angled inwards before costa and weakly concave before inner margin, most pronounced on costa and inner margin. Terminal area darker, diffuse. Fringes concolorous with wings or slightly paler, unicolorous. Hindwings slightly paler. Discal spots large, smaller on hindwings. Underside of wings paler, weakly irrorated with pale brown. Discal spots distinct and wing termen with dark fascia well developed. Female: wingspan 25–27 mm. Female distinctly smaller, forewings more round, forewing medial line evenly curved inwards on costa. Underside of wings with sharper markings than in male, dark fascia narrower. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Antennae bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Tympanal organs medium-sized in male, small in female. Male 8th sternite reduced in size, about 3/4 size of 8th tergite. Male 8th tergite concave laterally. Other sternites and tergites of both sexes undifferentiated. Variation: Variable. Wings yellowish grey to yellowish pale brown to almost grey with yellowish irroration, terminal area darker. Medial lines may be almost absent, but usually present at least as dots on costa and inner margin or medial lines may be broken into row of dots or be fully present, dark brown. Discal spots always present. Area between medial lines rarely dark. Dark fascia on wing underside may be wide and extend into wing margin and its inner margin may be bordered with pale light band. Veins on wing underside usually weakly visible, but occasionally brown and distinct. Genetically variable also, see Genetic data and Remarks. Male genitalia: Generally as in S. tenebraria, without diagnostic differences. Vesica with 13–16 cornuti of various lengths and one large cornutus at apex of aedeagus. Female genitalia: Generally as in S. tenebraria, without diagnostic differences. Signum very small (n=3) or absent (n=1). Potentially with tendency to reduction of signum towards the north, the structure being well developed in Italy: Monte Baldo (Sihvonen slide 2160, in coll. TLMF), very small in northernmost Italy: Völs am Schlern (Sihvonen slide 2123, coll. TLMF) and at the Italian/Swiss border in Umbrail Pass (Sihvonen slide 2162, coll. TLMF) and absent in Austria: Nordkette (Sihvonen slide 2125, coll. TLMF). Distribution and abundance: European, alpine. From southernmost Germany to northern Italy and from eastern Switzerland to north-eastern Slovenia and eastern Austria. Reported from Bosnia and Herzegovina (Rebel 1904), but not confirmed by authentic specimen(s). – Local, usually found in limited numbers. 173

Phenology: Univoltine. From early July to mid-August. Larva from August to June, hibern­ ating. Males active at night, attracted to light. Females apparently also nocturnal and attracted to light, but much more rarely than males. Females sometimes flushed up by day, flying short distances, sometimes found feeding on alpine flowers. Biology: Larva polyphagous. Found on Salix reticulata and S. serpyllifolia, and sitting between very low growing plants (M. Leipnitz pers. comm.). Recorded on Pimpinella and Campanula (Flamigni et al. 2016). Reared on Taraxacum and Lactuca (Flamigni et al. 2016). Pupation in a light silken web among plant litter and gravel on surface of bottom layer in rearing container (M. Leipnitz pers. comm.; rearing observation).

Text-figs 81–88. Diagnostic characters (indicated) of Sciadia zelleraria (Freyer, 1836) (figs 81– 84) and S. unicoloraria (Staudinger, 1871) (figs 85–88). Figures are not in scale. The genitalia and aedeagus of S. zelleraria are larger, see the genitalia plate for scaled figures and Table 2 for additional diagnostic characters. The sclerotized part of aedeagus (margin highlighted in picture) is poorly visible in grey-scale photograph, but distinct when the structure is stained and viewed in full colour. Text-fig. 81 (apex of aedeagus). Italy: Monte Baldo, 14.vii.1969, 2150 m, slide Sihvonen 2159 (coll. TLMF). Text-fig. 82. Italy: Völs am Schlern, 27.vii.2007, 2300–2440 m, slide Sihvonen 2122 (coll. TLMF). Text-fig. 83. Italy: Monte Baldo, 14.vii.1969, 2150 m, slide Sihvonen 2159 (coll. TLMF). Note: cornutus is absent in this specimen, but the rupture is visible where it should originate (cornutus deformed/detached?). Text-fig. 84. Austria: Mühlauer-Klamm, 2.vii.1933, 1000 m, slide Sihvonen 2161 (coll. TLMF). Text-fig. 85 (apex of aedeagus). France: Col de la Cayolle, 29–30.vii.2009, 2200 m, slide Sihvonen 2164 (coll. Skou). Text-fig. 86. Switzerland: Zermatt, 28.vii.11980, 2580 m, slide Sihvonen 2163 (coll. TLMF). Text-fig. 87. France: Col de la Cayolle, 29–30.vii.2009, 2200 m, slide Sihvonen 2164 (coll. Skou). Text-fig. 88. Italy: % Cima Fauniera, 28.vii.2009, 2450 m, slide Sihvonen 2165 (coll. Skou).

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Habitat: Alpine. Above treeline in a variety of habitat types. Recorded from 1600 up to 2500 m. Similar species: S. unicoloraria (Staudinger, 1871), see Text-figs 79–80 for adult characters, 81–88 for the male genitalia and Table 2 for selected diagnostic characters. S. caelibaria is smaller, without dark fascia on wing underside and the male and female genitalia show that it belongs to a different species group. Surprisingly, the externally very different S. ­tenebraria and S. zelleraria have very similar genitalia structures, we have not noticed diagnostic differences. Genetic data: Four BINs: BOLD:AAB4957 (n=8 from southern Germany, western Austria and northern Italy), BOLD:AAD2991 (n=8 from north-eastern Italy and south-eastern Austria), BOLD:ACE3560 (n=4 from northern Italy), BOLD:ABX0095 (n=1 from Italy: Veneto). Genetically heterogeneous, distances between the BINs 1.7–4.4%. Repeated cases of genetic introgression with Sciadia tenebraria, S. slovenica, S. innuptaria and S. dolomitica observed in the first three S. zelleraria clusters, according to the sympatric occurrence of S. zelleraria with the introgressed sister species. Altogether 20 specimens of Sciadia have so far been found to show mitochondrial DNA of one of the S. zelleraria BINs. One additional specimen of S. zelleraria with separate BIN (BOLD:ACJ3674) from northern Italy close to the Swiss border showing the opposite phenomenon, i.e. introgression of a zelleraria singleton into the broader pool of a “Sciadia” cluster, being close to one of the BINs of S. ­tenebraria from Switzerland. The phenomenon of tangled DNA barcodes is dealt with by Hausmann et al. (2011b), Flamigni et al. (2016), Huemer & Hebert (2016), and Mutanen et al. (2016). Remarks: Sciadia zelleraria is a variable species, both phenotypically and genetically. This variation is richly illustrated and explained in Flamigni et al. (2016). External appearance is somewhat dependent on the collecting locality, but not exclusively. This may be anticipated because S. zelleraria lives in mountain habitats and migration abilities of short-winged females are restricted, potentially resulting in poor gene flow between populations and phenotypic differences over time. Adults often rest on rocks and wing coloration is adapted to local substrate coloration. Taxon robusta has been characterized to include specimens from Austria (Tyrol) and Switzerland (Graubünden) that are generally pure grey, more rarely light yellowish grey, contrary to the nominal taxon that is characterized by specimens that are small and purer yellowish grey. The nominal taxon has been considered to be restricted to the most eastern part of the Alps (Austria: Hochschwab and Ennstal Alps, Slovenia: Kamnik and Julian Alps). The eastern Austrian populations are allopatric according to current knowledge. As DNA barcodes have accumulated, it has become more evident that S. zelleraria is a genetically heterogeneous species and if each genetic node is attributed a name this would result in abundant subspecies concepts. Genetically divergent specimens may occur close to each other, for instance in Stelvio Pass (Italy) and Sölden (Austria), which are only about 10 km away from each other (Flamigni et al. 2016). Sihvonen (current work) has dissected genitalia of numerous specimens across the distribution area of S. zelleraria. No constant morphological differences were found that would correlate with the observed genetic clusters, in agreement with Flamigni et al. (2016: p. 354): ‘The significance which is to be attributed to [the genetic data] is not clear and must be investigated further, because the genetic differences do not seem to correspond to constant differences in the genitalia and the wing pattern is also variable.’ For these reasons we do not consider taxon robusta valid at subspecies level and do not formally describe the genetic variation at subspecies level. 176

55. Sciadia unicoloraria (Staudinger, 1871), comb. n. Note: Identity of Enconista unicoloraria Staudinger, 1871 is clear (see below), but identity and distribution of other taxa associated with the name unicoloraria are uncertain. We have not had access to adequate material, so the taxonomy listed below is tentative and needs more research. For this reason all records in the distribution map are grouped under name unicoloraria. Enconista unicoloraria Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 173 (Spain: Andalusia?, see Distribution). Holotype ♂ (coll. Herbulot [Herbulot acquired the type from coll. Mabille, see Herbulot (1956)] in ZSM, examined, including genitalia). Transferred here from Elophos Boisduval, 1840 (for instance Leraut 1997; Scoble 1999; Vives Moreno 2014; Flamigni et al. 2016) to Sciadia Hübner, 1822, quoting structural similarities (new combination). Old literature gives uniformly Rambur, 1866 as the author of taxon unicoloraria (for instance Staudinger & Wocke 1871; Staudinger & Rebel 1901; Prout 1912–1916; Wehrli 1939–1954), whereas more recent literature, including Scoble (1999), refers to Staudinger, 1871 as the author, unicoloraria Rambur, 1866, Cat. Lep. And. pl. 19:6 being a non-binominal name, (for details see Herbulot 1956). The name was first validated by Staudinger (1871). Gnophos zelleraria f. bubaceki Schawerda, 1915: in Rebel, Verh. zool.-bot. Ges. Wien 65 (5/6): (88), fig. 5 (France: East Pyrenees, Vernet). Syntypes 4♂2♀. Taxon bubaceki is treated in the recent literature under Elophos unicoloraria, but we have not found reference from the literature who has proposed to change the combination and on what grounds. The original article illustrates a male and a female, and those could belong to S. zelleraria (which, how­ever, is considered to be restricted to the Alps). Taxon bubaceki is treated as valid at subspecific rank (for instance Leraut 1997; Scoble 1999; Hausmann et al. 2004; 2011a) or as a junior synonym of Elophos unicoloraria Staudinger, 1871 (for instance Leraut 2009). We have not had access to specimens from the eastern Pyrenees. More research is needed. Dasydia unicoloraria occidentalis Oberthür, 1913: Études Lépid. comp. 7: 321, pl. 183, figs 1796, 1797 (France: Larche and Lac d’Allos [Alpes de Haute Provence] and Spain: Picos de Europa [Asturia]). Syntypes 10♂. Identity of taxon occidentalis is problematic because according to the original description the type series contains specimens from both the western Alps and western Pyrenees. Redondo et al. (2009) classify the Spanish record from Picos de Europa as subsp. occidentalis, following Oberthür (1913). Taxon is considered valid at subspecific rank (for instance Vives Moreno 1994; Scoble 1999; Hausmann et al. 2004; 2011a), but more research is needed. Gnophos unicoloraria f. vallesiaria Wehrli, 1920: Mitt. ent. Ges. Basel 12: 3, fig. 12 (Switzerland: Valais, Hohwänghorn, Zermatterhörnli, 2500–3000 m. Syntypes 2♂ (ZFMK, examined externally). Treated valid at subspecific rank (for instance Leraut 1997; Scoble 1999; Leraut 2009; Hausmann et al. 2004; 2011a; Flamigni et al. 2016). Based on the material examined and its geographical locality within unicoloraria distribution area, we consider vallesiaria to fall within intraspecific variation and a junior synonym of unicoloraria Staudinger. More research is needed, see Remarks. Unavailable names (infrasubspecific): subnivea: Wehrli (1924) (f. of vallesiaria).

External characters and abdomen: Male: wingspan 28–34 mm (note: measured from apex to apex; Flamigni et al. (2016) give wingspan 31–38 mm, potentially measured from the maximum distance between wing margins, below the apex). Female: wingspan 21–26 mm (n=3). Generally as S. zelleraria, but with following quantitative differences in male: wing markings weak giving a more uniform appearance, delicately striated with brown in S. unicoloraria (wing markings often more pronounced giving a less uniform appearance, delicately irrorated with brown in S. zelleraria). Underside of wings with weak marginal fascia in S. unicoloraria (underside of wings with dark marginal fascia in S. zelleraria). ­Female 177

­ ccasionally whitish, with brown-grey markings in S. unicoloraria (female occasionally greyo ish, with pale brown markings in S. zelleraria). Examination of the genitalia and/or DNA barcodes is often necessary to confirm the identity. Also distribution data is important because according to current knowledge S. zelleraria and S. unicoloraria do not occur sympatrically. Variation: Forewing can have distinct medial and postmedial lines and hindwing post­medial line, both are usually dentate (typically in Switzerland), or medial lines can be almost absent (typically in France, Italy), appearing weakly on forewing costa only. Discal spots are distinct, with pale centre in some specimens. Ground colour can vary from whitish (often almost without any markings, typically in France: Alpes-de-Haute-Provence) to yellowish white (often with weak markings or markings almost absent, typically in France, Italy) to yellowish (often with distinct brown markings or almost entirely brown wings, typically in Switzerland) to ochreous (often with distinct brown, dentate medial lines and heavy striation, typically in eastern Pyrenees). Fascia on underside wing margin is usually weak but visible, but it can be almost absent. Male genitalia: Generally as in S. tenebraria and S. zelleraria, but with following differences (see Text-figs 81–88): Juxta margin smooth in S. unicoloraria (margin rough in S. ­zelleraria). Aedeagus small, length about 1.5 mm in S. unicoloraria (long, length about 2.5 mm in S. zelleraria). Vesica opens at 45 degree angle, aedeagus apical cornutus absent, lateral area of aedeagus apically weakly dentate, and aedeagus sclerotization reaches apex in S. unicoloraria (vesica opens at 90 degree angle, aedeagus apical cornutus present, lateral area of aedeagus apex smooth, and aedeagus sclerotization does not reach apex in S. zelleraria). Female genitalia: Only two female genitalia slides of S. unicoloraria have been available for study (Italy: near Lago Sommeiller, 7.viii.1998, 2950 m, prep. and coll. Flamigni; France: Hautes-Pyrénées, Gèdre, August [no other date data], MNHN-EL, prep. n. 14, coll. MNHN, Paris). Generally as in S. tenebraria and S. zelleraria, signum is absent in S. unicoloraria (present or absent in S. tenebraria and S. zelleraria). Distribution and abundance: European. Mountain species occurring only in south-eastern France, north-western Italy and south-western Switzerland. Single records from eastern Pyrenees and from Picos de Europa in north-western Spain from 1946 (Agenjo 1953). Male genitalia of the latter specimen, despite rather superficially drawn (Agenjo 1953), agreeing with the current concept of S. unicoloraria. The specimen is illustrated in Redondo et al. (2009, fig. 159a) also. Type locality of S. unicoloraria is assumed to be Spain: Andalusia (after Rambur 1866, presumably the mountains of Sierra Nevada), but the holotype does not bear any locality label and no other specimens have been recorded from that locality. Staudinger & Wocke (1871) questioned the type locality, and for this reason we show the Andalusian record with a question mark. Leraut (2009) illustrates two S. unicoloraria females, but in our opinion both are misidentifications: Plate 66, fig. 8 from France: Gèdre is likely to be Sciadia septaria (Guenée, 1858) and Plate 66, fig. 15 from France: Col du Petit Saint-Bernard is likely to be Elophos andereggaria (de la Harpe, 1853). – Local, often found in numbers. In the Pyrenees apparently very rare. Phenology: Univoltine. From early July to mid-August. Larva from August to June or July, hibernating. Males active by night, readily attracted to light. Very little is known about the females, with very few specimens traceable in collections. The rarity of the S. unicoloraria female is much in contrast to how relatively often the female of the sister species S. zelleraria is found. 178

Biology: Larva polyphagous. Recorded on various low plants (Flamigni et al. 2016). Habitat: Alpine. In a variety of habitat types above treeline from 1900 up to 2900 m. Similar species: S. zelleraria (Freyer, 1836), see Text-figs 79–80 for adult characters, Textfigs 81–88 for the male genitalia and Table 2 for selected diagnostic characters. S. caelibaria is smaller, without dark fascia on wing underside and the male and female genitalia show that it belongs to a different species group. Female genitalia of S. tenebraria and S. zeller­ aria are similar, without diagnostic differences. Genetic data: BINs: BOLD:ABZ6312 (n=2 from south-eastern France: Provence-Alpes-Côte d’Azur) and BOLD:ACE3564 (n=2 from north-western Italy: Piedmont), diverging by 3.3% (Mutanen et al. 2016). No specimens from the Pyrenees have been DNA barcoded so far. No introgression observed, so far. DNA barcodes are tangled with S.  zeller­aria, S. slovenica and S. tenebraria (Mutanen et al. 2016). Nearest species: S. tenebraria (1.1–1.9%). Remarks: S. unicoloraria is a variable species, both phenotypically and genetically, although the genetic data is still limited and accumulating. External variation is richly illustrated in Flamigni et al. (2016). As in S. zelleraria, external appearance is somewhat dependent on the collecting locality, but not exclusively. Adults often rest on rocks and wing coloration is adapted to local substrate coloration. P. Sihvonen has dissected the genitalia of numerous specimens across the distribution area of S. unicoloraria. No constant morphological differences were found that would correlate with the observed phenotypic and genetic clusters. It is possible that when more DNA barcodes become available, a more heterogeneous genetic structure will emerge.

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The Elophos and Yezognophos complex of genera The genus-level relationships of structurally, and partly ecologically, similar Elophos ­Boisduval, 1840, Yezognophos Matsumura, 1927 (=Parietaria Leraut, 1981 here regarded as synonym), Sciadia Hübner, 1822, Psodos Treitschke, 1825 (=Glacies Millière, 1874 here regarded as synonym), and potentially other related genera need further research because the relationships have not been investigated in an extensive analytical context so far. Foundations for the current classification were laid down by Wehrli (1951; 1953 in Wehrli 1939–1954), Herbulot (1963) and Sauter (1990), the last attempted to diagnose and classify the various Gnophini genera using phylogenetic systematics, but the conclusions were based on rather limited morphological data. Sauter (1990) did not specify which species from which genera were examined. In recent literature, the genera Elophos Boisduval, 1840 and Yezognophos Matsumura, 1927 include approximately 15 species in the Palaearctic region. The taxa have earlier been classified into two separate genera (Elophos and Parietaria in Müller 1996; Leraut 2009), in genus Elophos (Ebert 2003), in Yezognophos (Wolf 1988), in genus Elophos with Yezo­ gnophos and Parietaria as synonyms (Viidalepp 1996), or in genus Elophos sensu stricto and Elophos subgenus Yezognophos (Flamigni et al. 2016). It is shown in this book that Elophos and Yezognophos each possess unique morphological features (see Table 3) and each genus includes several morphologically distinct species groups (diagnosable particularly by the genitalia, see the relevant male and female plates). Therefore we do not transfer the observed heterogeneity into a classification that contains a single genus only, particularly when it is not clear what other taxa might be involved in the complex. We recognize two genera as valid: Elophos Boisduval, 1840 and Yezognophos Matsumura, 1927 but note that an extensive, modern phylogenetic research is needed to show whether the view adopted here will render one of the genera paraphyletic, whether the proposed diagnostic characters are unique and if reclassification is needed. Such research would also be needed to study evolutionary questions such as brachyptery as a potential unique synapomorphy for certain taxa or as having arisen through parallel evolution. Table 3. Diagnostic genus-defining characters and the included species of Elophos Boisduval, 1840 and Yezognophos Matsumura, 1927. Sauter (1990) presents an overview of diagnostic morphological characters of the Gnophos group and explains the difficulty of classifying Phalaena vittaria Thunberg, 1788, because it is transitional in many ways. Character

Elophos

Yezognophos

Female wings Female tympanal organs Male forewing costa Male tegumen Male 8th sternite Male vesica Habitat Species included

reduced in length reduced weakly concave large reduced, about 1/2 of 8th tergite with cornuti high mountain areas E. caelibaria E. zirbitzensis E. operaria E. andereggaria

fully developed normal straight or convex normal normal, 8th sternite and tergite about same size mostly without cornuti diverse, including lowland habitats Y. dilucidaria Y. sproengertsi Y. serotinaria Y. vittaria

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Elophos Boisduval, 1840 Elophos Boisduval, 1840, Genera Index meth. eur. Lepid.: 199. Type species: Geometra operaria Hübner, 1813. Acuspis Gumppenberg, 1893, Nova Acta Acad. Caesar. Leop. Carol. 59: 412, 430. Type species: Gnophos caelibaria Heydenreich, 1851.

Diversity and distribution: The genus Elophos, as applied here, contains four species in the eastern Palaearctic region, restricted to mountain habitats. One additional species, Elophos barbarica Prout, 1915, from South Africa has been placed in the genus (Scoble 1999). We have examined the holotype (in NHMUK and it is misplaced, as noted already by Prout (1915). External characters and abdomen: Wingspan of males approximately 25–37 mm, females are brachypterous or if capable of flying, wings are distinctly shorter than in males. Male forewing costa weakly concave. Wings grey to grey-brown to yellowish brown, often heavily irrorated with different shades of brown. Forewing medial and postmedial lines often weakly expressed, dentate. Hindwing with dentate postmedial line. Discal spots large. Markings weaker on wing underside. Male antennae bipectinate, female antennae filiform. Hindlegs of both sexes with 2+2 spurs. Male tympanal organs medium-sized, female organs very small. Male 8th sternite small, about half size of 8th tergite. Male genitalia: Uncus weakly triangular. Socii absent. Gnathos with small lateral sclerites and granulate or smooth medial plate. Tegumen massive (Text-fig. 89). Valva wide at base, apex upturned. Costa sclerotized, base with few stout setae, apex setose. Anterior arms of juxta dentate or spinose. Posterior arms of juxta short, triangular. Saccus elongated medially. Vesica with several cornuti. Anterior arms of juxta offer species-level diagnostic characters, see genitalia plates. Female genitalia: Papillae anales wide, setose. Lamella antevaginalis membranous, large. Lamella postvaginalis membranous, small, wrinkled. Posterior part of ductus bursae (colliculum) wide, concave, sclerotized. Ductus bursae wide, sclerotized or weakly sclerotized. Anterior part of corpus bursae membranous, wrinkled. Signum very small, dentate or absent. Biology, habitat, phenology: Larva of E. caelibaria and E. andereggaria polyphagous on low plants, former for instance on Rumex, Ranunculus montanus, Sedum, Saxifraga oppositi­ folia, latter reared on Plantago and Taraxacum officinale (Flamigni et al. 2016). Larval development of E. operaria and E. zirbitzensis apparently taking two years (Thurner 1945), adults of E. zirbitzensis occurring in uneven years only (Lichtenberger 1988, 1992). Italian Elophos species living at high altitudes, roughly between 1800 and 3200 m on screes and other rocky habitats, low grasses and debris (Flamigni et al. 2016). Immature stages: Larva of E. caelibaria stout, grey-brown or rather dark brown, with row of pale brown, almost square markings dorsally (Lepiforum 2018). Pupal cremaster of E. caeli­ baria with stout setae D2 only (similar to equivalents in G. obfuscata (Denis & Schiffer­ müller, 1775), see Text-fig. 27), which are remote and bent towards each other, apex hooked (Patočka & Turčani 2005). Remarks: Elophos species tend to vary externally quite a lot, as regards wing colour and intensity of markings. External appearance is somewhat dependent on the collecting locality, and this may be anticipated because Elophos species live in mountain habitats and migration abilities of short-winged females are restricted, potentially resulting in poor gene flow 181

Text-fig. 89. Many Sciadia and Elophos species have the male tegumen very massive. The structure is illustrated here for Elophos operaria (Hübner, 1813). Switzerland: Lucerne, Mount Pilatus, 2050 m, 4.viii.1977, slide Sihvonen 2120 (coll. TLMF). Tegumen is often distorted by cover glass when the genitalia are mounted in ventral view on an objective glass. Photographed in water.

between populations and phenotypic differences over time. Adults often rest on rocks and wing coloration is adapted to local substrate coloration. This has resulted over the time in the description of numerous subspecies, variations and forms. P. Sihvonen has dissected the genitalia of Elophos species across their distribution areas and often no constant morphological differences were found that would correlate with the observed phenotypic clusters. Genetic data is still somewhat limited but accumulating. For these reasons, the (historical) subspecies are often repeated as they appear in the literature, but it is acknowledged that further research may prove many of those to be synonyms. See the chapter ‘On species and subspecies’ in Skou & Sihvonen (2015), which is applicaple to many taxa treated in the current volume. Recently Elophos has been classified in the Gnophini (Vives Moreno 1994; Leraut 1997; Hausmann et al. 2004; 2011a). Elophos has not been included in a multi-gene analysis so far. Systematic position: Gnophini.

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56. Elophos caelibaria (Heydenreich, 1851) Hibernia caelibaria Heydenreich 1851: Lepid. Eur. cat. meth. (Edn. 3): 54 (Austria: Tyrol, Grossglockner, Bolzano). Syntype(s), including ♀. Originally described as a non binominal name by Herrich-Schäffer, 1848, Syst. Bearbeitung Schmett. Eur. 3: pl. 68, fig. 421. Gnophos spurcaria de la Harpe, 1853: Faune Suisse Lépid. 4: 63, [pl. 1], fig. 1 (Switzerland: at the foot of Mount Rosa). Syntype(s). Scoble (1999) and László et al. (2018) treat spurcaria valid at species rank without justification. We consider it tentatively valid at subspecies level, thus agreeing with recent literature (for instance Leraut 2009; Rezbanyai-Reser & Aistleitner 2013; Flamigni et al. 2016), although Redondo et al. (2009) consider it a form of E. caelibaria. Gnophos caelibaria var. jugicolaria Fuchs, 1901: Ent. Zeitschrift 62: 377 (Italy and Switzerland border: Stelvio pass, 2200 m). Syntype ♂ (type specimen depositary not given in the original description). Validated at subspecific rank (Scoble 1999), but retained as junior synonym of Gnophos spurcaria for instance by Flamigni et al. (2016), the latter here confirmed. Gnophos caelibaria var. senilaria Fuchs, 1901: Ent. Zeitschrift 62: 375 (Germany: Oberstdorf, Nebelhorn). Syntype ♂ (type specimen depositary not given in the original description). Herewith downgraded from subspecific rank of Elophos caelibaria senilaria (Scoble 1999; Rezbanyai-Reser & Aistleitner 2013) to synonymy (syn. n.), quoting absence of distinct morphological and molecular characters. See Rezbanyai-Reser & Aistleitner (2013) for a recent treatment. Gnophos intermedia Kautz, 1930: Z. Öst. EntVer. 15 (3): 36 (Austria: Grossglockner Mts.). Syntypes 38♂2♀ (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany). Junior synonym of Hibernia caelibaria. Unavailable names (infrasubspecific): insignata: Kautz (1930) (junior subjective synonym of spurcaria de la Harpe); scalettaria: Millière (1864) (junior subjective synonym of spurcaria de la Harpe). – (incorrect subsequent spellings): caelebaria: Herrich-Schäffer (1852); coeli­ baria: Guenée (1858); scalletaria: Vives Moreno (1994) and Leraut (1997).

External characters and abdomen: Male wingspan 27–30 mm, female 12–17 mm (brachy­ pterous). Male ash-grey, irrorated with brown, terminal area often darker. Forewing medial and postmedial lines brown, unsharp, latter curved inwards on costa, often most pronounced and widened on costa. Outer margin of postmedial line often bordered with paler area. Subterminal line weak, often visible near costa only, dentate. Terminal line very narrow, brown, continuous. Fringes unicolorous, pale brown. Forewing discal spots large, unsharp, hindwing discal spots smaller. Hindwings grey-brown, postmedial line most pronounced, its outer margin bordered with paler area. Wings below almost uniform grey-brown, without markings, apart from well-developed discal spots, and wing margin without dark fascia. Female brachypterous, forewings pointed, wing colour similar to males. Forewing medial and postmedial lines dark brown to blackish, narrow, sharp. Hindwings with postmedial line most pronounced. Discal spots small. Wings below almost uniform grey, discal spots and postmedial line weakly expressed. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Antennae bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Tympanal organs medium-sized in male, small in female. Male 8th sternite reduced in size, about 3/4 size of 8th tergite. Male 8th tergite concave laterally. Other sternites and tergites of both sexes undifferentiated. Variation: Variable species, external appearance correlates with locality. Male wings ashgrey, irrorated with brown with rather weak and unsharp markings (nominotypical subspecies) to yellowish brown, occasionally with tint of orange, with strong and sharp markings (subspecies spurcaria). Female wingspan larger and markings more pronounced in subspecies spurcaria. The specimens at extreme ends of variation could be considered separate species, but the specimens are structurally (including the genitalia) and genetically (DNA barcodes) similar. Taxon senilaria Fuchs, 1901, from Germany: Oberstdorf, Nebelhorn, is 183

not considered valid at subspecies level, based on absence of distinct morphological and molecular characters. See Rezbanyai-Reser & Aistleitner (2013) for a recent treatment. Male genitalia: Uncus triangular, setose, apex sharp. Socii absent. Gnathos arms sclerotized, wide, medial plate large, granulate. Tegumen large. Valva very wide at base, short, costa sclerotized and with several stout setae on small lobe, subcostal ridge near apex setose. Valva margin evenly curved or weakly concave (Text-fig. 91). Anterior arms of juxta large, partially sclerotized with sharp teeth, and with membranous sac, arms separate basally. Posterior arms of juxta small, hollow, horn-shaped. Saccus short, with round apex, curved dorsally. Aedeagus very small (compared to the genitalia capsule), stout, caecum short, apex with large, curved horn. Vesica opens at about 90 degree angle, very wide at base, apex with row of about 10 cornuti. Ductus ejaculatorius opens from medial part, its base wide. Female genitalia: Papillae anales round, setose. Apophyses anteriores short. Lamella antevaginalis concave, sclerotized, less in medial area. Lamella postvaginalis membranous, with small, frilled part medially. Ductus bursae short, wide, partly sclerotized. Posterior part of corpus bursae partly sclerotized (left side when viewed ventrally), partly membranous and wrinkled. Anterior part of corpus bursae membranous, wrinkled. Signum small, weakly stellate. Subsp. spurcaria (Text-figs 92, 93, 95) is yellowish brown, occasionally with tint of o ­ range, with strong and sharp markings. Female wingspan is larger and markings are more pronounced than in nominotypical subspecies. In male genitalia the costa lobe, which bears stout setae, is distinct and margin of valva is concave. No diagnostic differences have been observed in the female genitalia. Distribution not known in detail, but recorded from south-western Austria, northern Italy, south-western Switzerland and in border area between France and Italy. Considered valid tentatively at subspecies level, more research is needed. Distribution and abundance: European. Distributed in the Alps from south-eastern France to Austria and north-western Slovenia and from southern Germany to northern Italy. There is a single confirmed record from the Pyrenees, in addition to old records from the Catalonian Pyrenees (see Redondo et al. 2009 for details) and the species is reported from Montenegro (Durmitor) also (Rebel 1913; Kautz 1930), but we have not seen authentic material. In western Austria (Vorarlberg) the nominotypical subspecies (treated as subsp. senilaria by Rezbanyai-Reser & Aistleitner 2013) and subspecies spurcaria almost meet (Rezbanyai-Reser & Aistleitner 2013). – Local, not common, usually seen in limited numbers. Phenology: Univoltine. From late June to mid-August (Kerschbaum & Pöll 2010), in Switzer­land throughout July (D. Bolt pers. comm.). Larva from August to June, hibernating. Males active at night, attracted to light, sometimes flushed up by day (D. Bolt pers. comm.). Females found in sunshine sitting on flowers of Achillea (M. Leipnitz pers. comm.). Biology: Larva polyphagous. Recorded on Rumex, Ranunculus montanus, Sedum, Saxi­ fraga oppositifolia, Plantago and Campanula (Flamigni et al. 2016). Reared on Sedum ­album, Thymus serpyllum, Campanula scheuchzeri, Taraxacum and Lactuca (Flamigni et al. 2016). Pupation below stones (D. Bolt pers. comm.). Habitat: Alpine, petrophilous. In a variety of habitat types above treeline, especially stony slopes (D. Bolt pers. comm.) and rocky places on limestone or silicate with sparse vegetation (N. Pöll pers. comm.). From 1900 up to 2600 m, exceptionally lower (1500 m in coll. Skou). In Switzerland from 2200 up to 3300 m (D. Bolt pers. comm.).

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Similar species: E. zirbitzensis males smaller (wingspan 24–25 mm), more contrasted, bipectinate male antennae with shorter branches and male aedeagus with short and rather straight apical spine (E. caelibaria males larger, wingspan 27–30 mm, less contrasted, bipectinate male antennae with longer branches, male aedeagus with long and curved apical spine (Text-figs 98–99). E. zirbitzensis females smaller, wingspan 11–12 mm, wings narrower, signum absent in female genitalia (wingspan 12–17 mm, wings wider, and signum present in E. caelibaria). In addition, Sciadia zelleraria and S. unicoloraria males larger, with dark fascia on wing margin on wing underside (male smaller and dark fascia absent in S. caelibaria). S. zelleraria and S. unicoloraria females with short wings and dark fascia on wing underside (female brachypterous and dark fascia absent in S. caelibaria). Genitalia allow reliable identification. Genetic data: BINs: BOLD:AAC5403 (n=20 from southern Germany, Austria and northern Italy), BOLD:ACJ4261 (n=11 from Austria and Switzerland), BINs diverging by 1.4%, genetically heterogeneous also within the BINs. Nearest species: E. zirbitzensis (4.2%), S. zelleraria (4.4%), Y. sproengertsi (4.4%). So far, three cases of introgression (S. tenebraria, S. slovenica) into BOLD:AAC5403 observed, but no E. caelibaria in other species clusters. See also Mutanen et al. (2016). Remarks: Female genitalia of Elophos caelibaria and E. zirbitzensis are structurally similar to Sciadia, having membranous lamella antevaginalis, wide and sclerotized ductus bursae, and anterior part of corpus bursae is membranous and wrinkled. Male genitalia of E. caelibaria and E. zirbitzensis, however, have differently structured anterior arms of juxta with numerous spines/teeth and a membranous sac. More research, preferably combining extensive morphological and DNA data, in addition to the life history and ecological data, is needed to show whether these two species should be transferred to Sciadia.

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Text-figs 90–95. Diagnostic characters (indicated) of Elophos caelibaria caelibaria (Heydenreich, 1851) and E. caelibaria spurcaria (de la Harpe, 1853) (figures are not in scale). The differences are not always as distinct as illustrated here, and there is variation between these extremes. Text-fig. 90. E. caelibaria caelibaria adult male. Austria: Nordkette, 2300 m, 2.vii.1963 (coll. TLMF). Text-fig. 91. E. caelibaria caelibaria male genitalia, from the same specimen, slide Sihvonen 2126 (coll. TLMF). Text-fig. 92. E. caelibaria caelibaria adult female. Austria: Kärnten, Glockner-Hochtor, 2400– 2600 m (ex pupa 27.vi.1988, emerged 5.vii.1988) (coll. TLMF). Text-fig. 93. E. caelibaria spurcaria adult male. Italy: South Tirol, Timmelsjoch, 2600 m, vii.1974 (ex pupa) (coll. TLMF). Text-fig. 94. E. caelibaria spurcaria male genitalia, Switzerland, Umbrail pass, 2200 m, 30.vii.1975, slide Sihvonen 2168 (coll. TLMF). Text-fig. 95. E. caelibaria spurcaria adult female. Italy: South Tyrol, Timmelsjoch, 2600 m (ex pupa, emerged vii.1974 (coll. TLMF). Female genitalia of E. caelibaria caelibaria and E. caelibaria spurcaria are similar, without distinct differences, so those are not illustrated.

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57. Elophos zirbitzensis (Pieszcek, 1902) Gnophos caelibaria var. zirbitzensis Pieszcek, 1902: in Rebel, Verh. zool.-bot. Ges. Wien 52 (1): 11 (Austria: Zirbitzkogel). Syntype(s). Unavailable names (incorrect subsequent spelling): zirbitsensis: Leraut (2009).

External characters and abdomen: Similar to E. caelibaria (Heydenreich), but wingspan smaller (24–25 mm in males and 11–12 mm in females), wings more contrasted (forewing medial area darker than rest of wing, postmedial line often bordered with pale grey to whitish), bipectinate male antennae with shorter branches (0.16–0.17 mm) and wings below with weak discal spots in E. zirbitzensis (in E. caelibaria male wingspan 27–30 mm, female wingspan 12–17 mm, wings more uniform, i.e. forewing medial area and margin of post­medial line often same colour as rest of wing, male antennae with longer branches (0.22–0.26 mm) and wings below with rather distinct discal spots). One of the examined males (slide Sihvonen 2135, coll. TLMF) has an unusual elongated structure on the membrane of 8th segment between sclerites, similar in appearance but smaller to that found in Dicrognophos siciliana Skou & Sihvonen sp. n. (see D. siciliana for illustration). Variation: Very little. In males the dark colour between forewing postmedial line and subterminal line often more pronounced near inner margin only, but occasionally dark area extends into costa also. Similarly on hindwing, terminal area sometimes concolorous with rest of wing, or darker. Forewing discal spot often solid, but sometimes pale-centred. Male genitalia: Similar to E. caelibaria, but male aedeagus with short and rather straight apical spine in E. zirbitzensis (aedeagus spine long and curved in E.  caeli­baria) (Text-figs 96–99).

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Text-figs 96–99. Diagnostic characters (indicated) of Elophos caelibaria (Heydenreich, 1851) and E. zirbitzensis (Pieszcek, 1902). Text-fig. 96. E. caelibaria, Austria: Kärnten, Glockner-­Hochtor, 2400–2600 m (ex pupa 27.vi.1988, emerged 5.vii.1988), slide Sihvonen 2126 (coll. TLMF). Text-fig. 97. E. caelibaria, Italy: South Tyrol, Timmelsjoch, 2600 m, ex pupa vii.1974, slide Sih­ vonen 2168 (coll. TLMF). Text-fig. 98. E. zirbitzensis, Austria: Zirbitzkogel, Lavanttaler Alpen, Styria, 2200 m, 27.vi.1949, slide Sihvonen 2135 (coll. TLMF). Text-fig. 99. E. zirbitzensis, Austria: Styria, Zirbitzkogel, Juli (no other date data), slide Sihvonen 2169 (coll. TLMF).

Female genitalia: Similar to those of E. caelibaria, but in E. zirbitzensis signum absent, ductus bursae symmetrical and longer (in E. caelibaria small stellate signum present, ductus bursae asymmetrical and shorter). Distribution and abundance: European, alpine. Known from Austria only, from Zirbitzkogel mountain (Styria: Lavanttaler Alps). Huemer et al. (2009) show localities of numerous old records from north-west of the current distribution area. – Very local and rare.

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Phenology: Univoltine, bi-annually. From late June to early August in uneven years. By rearing in a refrigerator hibernation as small caterpillars and development from egg to adult taking only one year (M. Leipnitz pers. comm.). Males active at night, attracted to light. Also found as larva or pupa under stones. Biology: Larva polyphagous. Reared on Saxifraga moschata and S. decipiens (M. Leipnitz pers. comm.). Pupation in a light silk web between plant litter (M. Leipnitz pers. comm.; rearing observation). Rearing is difficult and needs to be performed in a refrigerator (M. Leipnitz pers. comm.). Habitat: Alpine, petrophilous. Above treeline on slopes with rocks and stones. From 2000 up to 2500 m (Fajčik 2003). Similar species: E. caelibaria, see that species. Genetic data: BIN: BOLD:AAX7668 (n=1 from south-eastern Austria). Nearest species: E. caelibaria (4.2%), S. zelleraria (4.8%). Remarks: See text under E. caelibaria. E. zirbitzensis was originally described as a variation of E. caelibaria. Species status is supported by several morphological structures, DNA barcodes, and by sympatric occurrence with E. caelibaria in parts of its distribution area (Huemer et al. 2009).

58. Elophos operaria (Hübner, 1813) Geometra operaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 69, fig. 359 (­Europe). Syntype(s) lost. With regard to the existence of related, similar taxa the following male neotype is designated to promote nomenclatural stability, to fix identity and type locality: NEOTYPE, Geometra operaria ♂, Hübner, 1813 [red rectangle label]; Designated by Pasi Sihvonen, in Geometrid Moths of Europe, vol. 6; Austria, [Niederösterreich/Steiermark (Lower Austria/Styria) border, approximately 47°41‘N 15°41‘E] Rax 1800 m, 13.vi.1967, H. Meier leg. (dissected, slide 2116 Sihvonen), coll. TLMF. Gnophos operaria var. hoefneri Rebel, 1903: Verh. zool.-bot. Ges. Wien 53 (8/9): 426 (Austria: Zirbitzkogel). Syntype(s). Considered valid at subspecific rank (Scoble 1999), requiring revision, potentially with clinal character transitions towards nominotypical subspecies. Elophos operaria subsp. necopinatus Rezbanyai-Reser, 1992: Mitt. Schweiz. ent. Ges. 65 (3–4): 217, figs 2, 3 (c1–c5), 4 (a5–b5) (Switzerland: Pilatus-Kulm, 2060 m). Holotype ♂ (NML, examined externally, based on a photograph in the original description). Considered valid at subspecific rank (Scoble 1999), requiring revision, potentially with clinal character transitions towards nominotypical subspecies. Unavailable names (infrasubspecific): anastomosis: Hoffmann & Klos (1913) (ab).

External characters and abdomen: Male wingspan 33–35 mm, female 7–10 mm (micropterous). Male pale grey, weakly irrorated with brown or blackish brown, medial area concolorous or slightly darker than rest of wings. Forewing medial and postmedial lines indistinct, darkest on veins, dentate. On hindwing postmedial line most pronounced. Postmedial line bordered with paler colour, wing terminal area slightly darker, irrorated with brown. Terminal line discontinuous, strongest between vein endings. Fringes concolorous with wings. Discal spots small, roundish, solid or with pale centre. Wings below slightly darker, almost uniform grey, only discal spots, forewing medial area and postmedial line somewhat visible, particularly near forewing costa, without dark fascia on terminal area. Female micropterous, forewings narrow and pointed, grey-brown with indistinct markings. Hindwings short stumps. Frons, collar, ­thorax 189

and abdomen of both sexes concolorous with wings. Antennae bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Tympanal organs medium-sized in male, very small in female. Male 8th sternite about 1/2 size of 8th tergite, posterior margin distinctly narrower. Male 8th tergite concave laterally. Other sternites and tergites of both sexes undifferentiated. Variation: Very variable, external appearance correlates with locality. Male wings vary from pale grey with weak markings and wingspan 33–35 mm (nominotypical subspecies) to yellowish with distinct markings and heavily irrorated with brown, often with dark medial area, and wingspan 37–38 mm (subspecies hoefneri) to almost uniform brown and wingspan 37–38 mm (subspecies necopinatus). Female wing length varies from very short stumps (wingspan 7–10 mm) to brachypterous (wingspan up to 12 mm). Subsp. hoefneri is larger, male wingspan about 37–38 mm, wings are yellowish, heavily ­irrorated with brown to dark brown, medial lines are dark and distinct, particularly post­ medial line. Forewing medial area often distinctly darker than rest of wing and discal spots often with pale centre. Male and female genitalia as in the nominotypical subspecies. Described from Austria: Zirbitzkogel, but distribution not known in detail. Considered valid at subspecific rank (Scoble 1999), requiring revision, potentially with clinal character transitions towards nominotypical subspecies. DNA barcodes not available so far. Subsp. necopinatus is larger, male wingspan about 37–38 mm, wings are almost uniform brown, yellowish ground colour weakly visible. Medial lines are weakly visible, forewing ­medial area often concolorous with rest of wing and discal spots often solid. Male and female genitalia as in the nominotypical subspecies. An allopatric population is described from Switzerland: Pilatus-Kulm, and apparently not known from elsewhere. Considered valid at subspecific rank (Scoble 1999), requiring revision, potentially with clinal character transitions towards nominotypical subspecies. DNA barcodes not available so far. Subspecies is richly illustrated in Rezbanyai-Reser (1992). Male genitalia: Uncus apex blunt. Socii absent. Gnathos arms fused, medial plate large, smooth, apex triangular. Tegumen large (Text-fig. 89). Valva very wide at base, directed upwards at 45 degree angle, costa sclerotized, basal lobe with few stout setae, outer margin of lobe evenly curved, subcostal ridge near apex weakly setose. Anterior arms of juxta large, bases symmetrical, apex densely setose, arms bent at about 90 degrees. Posterior arms of juxta small, hollow, apex acute. Saccus weakly elongated medially. Aedeagus very small (compared to the genitalia capsule), curved, apical portion covered with dentate spines. Vesica opening at about 90 degree angle, base enlarged, with row of about 5–9 large cornuti. Variation of dentate aedeagus spines and number of cornuti on vesica illustrated in Rezbanyai-Reser (1992). Female genitalia: Papillae anales wide, setose. Apophyses anteriores short. Lamella antevaginalis concave, sclerotized. Lamella postvaginalis membranous, membranous, frilled. Ductus bursae wide, rather long, constricted laterally, weakly sclerotized. Base of ductus seminalis enlarged. Corpus bursae pear-shaped, membranous sac (illustrated in the genitalia plate) or posterior part with weakly sclerotized ridges, medial area constricted and anterior part wrinkled (illustrated in the genitalia plate). Signum small, weakly granulate, or very small, almost absent. Distribution and abundance: European. Mountain species occurring in isolated populations in the Alps from Switzerland to eastern Austria, in the Carpathians from Poland, Czech Republic and Slovakia to Romania. – Local and rare, nearly always found in few or single specimens. 190

Phenology: Univoltine. From late June to mid-August. In the Czech and Slovak Republics, caterpillar from August to June, hibernating (Z. Las˘tu˚ vka pers. comm.). Males active at night, attracted to light. Males and rarely females occasionally found by day, sitting on rocks (Z. Las˘tu˚ vka pers. comm.). Biology: Larva polyphagous. Recorded on low plants, including Campanula (Vorbrodt 1914). Habitat: Subalpine or alpine. In the Czech and Slovak Republics in stony alpine meadows (Z. Las˘tu˚ vka pers. comm.). In the Alps from 1700 up to above 2000 m, exceptionally lower (1300 m in coll Skou). In the Czech Republic from 1500 up to 1600 m, in Slovakia from 1500 up to 2000 m (Z. Las˘tu˚ vka pers. comm.). Similar species: Structurally E. andereggaria is the most similar species, see Table 4 for an overview of selected diagnostic characters and Text-figs 100–101 for aedeagus and vesica characters. Wehrli (1939–1954) writes that E. andereggaria male has longer antennal branches than E. hoefneri. There appears to be a small difference, but the character must be used carefully because the length of setae varies greatly according to position on the antenna. The longest branches of E. operaria measure 0.39–0.42 mm, and those of E. andereggaria 0.55–0.58 mm. Genetic data: One DNA barcode from ‘Austria’ (no exact locality given on the labels, no BIN assigned, 458bp). Nearest species: E. caelibaria (6.1%). Genetically far from E. andereggaria (9.0%). Remarks: Scoble (1999) lists Geometra nitelaria Esper, 1806 as a synonym of Elophos operaria Hübner, 1813. According to Hacker (1999), taxon nitelaria is a junior synonym of Gnophos obfuscatus (Denis & Schiffermüller, 1775). The holotype male of taxon nitelaria is in Landesmuseum Wiesbaden (Germany).

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Table 4. Overview of diagnostic characters of European Elophos operaria (Hübner) and E. andereggaria (de la Harpe). Some male specimens may be very difficult to identify from external features alone, but genitalia, particularly male genitalia, provide numerous reliable characters. The mentioned characters are indicated by arrows in the relevant plates. Character

E. operaria

E. andereggaria

Male forewing subterminal area Male wingspan Female wing length Uncus apex Lobe on valva costa Base of anterior juxta arms Apex of anterior juxta arms

without silvery grey hue 33−38 mm micropterous, about 4 mm blunt outer margin round symmetrical bent about 90 degrees

sometimes with silvery grey hue 37−42 mm short, about 11 mm round outer margin angled asymmetrical bent about 45 degrees

Saccus Apical part of aedeagus Ductus bursae

weakly elongated medially covered with dentate spines wide, constricted laterally

distinctly elongated medially bare (without spines) narrow, not constricted laterally

Text-figs 100–101. Diagnostic characters (indicated) of Elophos operaria (Hübner, 1813) and E. andereggaria (de la Harpe, 1853). Text-fig. 100. E. operaria (insert: cornuti from the rear side), Austria: Rax, 1800 m, 13.vi.1967, slide Sihvonen 2116 (coll. TLMF). Text-fig. 101. E. ander­ eggaria (insert: cornuti from the rear side), Switzerland: Wallis, without date, slide Sihvonen 2172 (coll. TLMF).

59. Elophos andereggaria (de la Harpe, 1853) Gnophos andereggaria de la Harpe, 1853: Faune Suisse Lépid. 4: 67, [pl. 1], figs. 3, 4 (Switzerland: Haut Valais, near Brigg). Syntypes ♂♀. Unavailable names (infrasubspecific): mauricauda: Oberthür (1913) (ab.) (see Wehrli 1939– 1954, p. 621, for detailed description of the taxon).

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External characters and abdomen: Male wingspan 37−42 mm. Male similar to E. operaria, particularly to those of putative subspecies E. operaria hoefneri and E. operaria necopinatus, without distinct diagnostic differences. Largest species in genus Elophos (wingspan 37−42 mm), outer margin of forewing postmedial line often with silvery grey tint in E. andereggaria (wingspan 33−38 mm, outer margin of forewing postmedial line without silvery grey tint in E. operaria). Wings yellowish or yellowish grey, heavily irrorated with brown. Postmedial line distinct, dentate. Fringes chequered yellowish and brown. According to Wehrli (1939–1954) E. andereggaria male with longer antennal branches than E. hoefneri, apparently valid as a (small) differential character, but to be used carefully because of greatly variable length of branches according to their position on the antenna. Longest branches of E. operaria 0.39–0.42 mm, of E. andereggaria 0.55–0.58 mm. Female wingspan 24–27 mm, wings yellowish white, heavily irrorated with brown or grey-brown, giving sprinkled or marbled appearance. Forewing postmedial line darker, dentate, outer margin bordered with purplish band. Medial and terminal areas paler. Terminal line narrow, dark brown, discontinuous. Fringes almost uniform yellowish white. Discal spots small, solid or pale-centred. Wings below grey-brown, with weaker markings, irrorations finer, discal spots well visible, terminal area concolorous (not darkened). Frons, collar, thorax and abdomen of both sexes concolorous with wings. Other external structures and abdomen as in E. operaria. Variation: Little. Density of irrorations varies, consequently yellowish, yellowish white or brown to grey-brown can be the dominant colour. In males the forewing terminal area can have silvery grey hue, and medial area can have olive-green hue. Male discal spots often pale-centred. Male genitalia: Similar to E. operaria, see Table 4 for an overview of diagnostic characters. Female genitalia: Similar to E. operaria, but in E. andereggaria ductus bursae narrow, not constricted laterally (in E. operaria ductus bursae wide, constricted laterally). Signum small, irregular in shape or absent (or only a weak rupture visible on ductus bursae). Corpus bursae roundish, membranous sac (illustrated in the genitalia plate) or small and almost fully wrinkled (not illustrated). Distribution and abundance: European. Mountain species occurring in a small area in south-western Switzerland (Valais) and the adjoining areas of the Aosta Valley and Piedmont in north-western Italy. We report E. andereggaria from France, based on a female specimen collected by Robert Mazel (France, Savoie, Cormet d’Aréches, 29.vii.1987, about 2100 m, dissected, slide PS2170 Sihvonen, coll. Sihvonen). The discovery was predictable, being 25 km from nearest records on Swiss-Italian border. The French side of Col du Petit Saint-Bernard was visited in summer 2018 and several males were found (Tautel & Colomb 2018). Leraut (2009) illustrates an E. andereggaria female from Switzerland: Simplon (plate 67, fig. 12), in addition to a female of ‘Elophos unicoloraria’ (plate 66, fig. 15) from France: Col du Petit Saint-Bernard (without date). The latter in our opinion is likely to be E. ander­ eggaria, and if so, it is the second known record of E. andereggaria from France. – Rare and local, usually found in limited numbers. Phenology: Univoltine. From late June to mid-August (Flamigni et al. 2016). Larva from late July to early June, hibernating (D. Bolt pers. comm.). Male active at night, readily attracted to light. Female difficult to find, accidentally found by day sitting on stone (J. Gelbrecht pers. comm.). Biology: Larva probably polyphagous like other species of this genus. 193

Habitat: Alpine. Above treeline. Mainly on slopes with rocks, stones and sparse vegetation in between. Also found on slopes without rocks and stones. From 2000 up to 2800 m. Similar species: Some male specimens of Elophos operaria (Hübner), particularly those of putative subspecies E. operaria hoefneri and E. operaria necopinatus externally very similar but with different genitalia. See Table 4 for an overview of selected diagnostic characters and Text-figs 100–101 for aedeagus and vesica characters. According to Wehrli (1939– 1954) E. andereggaria male with longer antennal branches than E. operaria hoefneri, apparently valid as a (small) differential character, but to be used carefully because of greatly variable length of branches according to their position on the antenna. Longest branches of E. ­operaria 0.39–0.42 mm, of E. andereggaria 0.55–0.58 mm. Genetic data: BIN: BOLD:AAE1311 (n=3 from Switzerland, north-westernmost Italy). Genetically heterogeneous, one barcode from Aosta valley diverging by 0.9%. Nearest species: S. tenebraria, E. caelibaria (both 6.1%). Remarks: Fajčik (2003) illustrates a male from Austria: Zirbitzkogel, stating it to be E. andereggaria (de la Harpe, 1853) in coll. TLMF in Innsbruck. E. andereggaria is not known from Austria (Huemer et al. 2009; Wiesmair pers. comm.). The illustrated specimen is in our opinion E. operaria and Zirbitzkogel is a well-known habitat and collection site of E. operaria.

Yezognophos Matsumura, 1927 Yezognophos Matsumura, 1927, Insecta matsum. 1: 187. Type species: Yezognophos kononis Matsumura, 1927. Taxon kononis is considered currently a subspecies of vittaria (Thunberg, 1788). Yezognophos Matsumura, 1927 has been considered a junior synonym of Elophos Boisduval, 1840 (e.g. Viidalepp 1996; Scoble 1999) or a subgenus of Elophos (Flamigni et al. 2016). We consider it a valid genus based on morphological characters as summarized in

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Table 3, but acknowledge that molecular data is needed to resolve its relationships in relation to other taxa. Parietaria Leraut, 1981, Alexanor 12 (1): 37. Type species: Geometra dilucidaria Denis & Schiffer­ müller, 1775. Leraut (1981) proposed Parietaria as a replacement name for Catascia Hübner sensu auct., for dilucidaria and related taxa, including vittaria (Thunberg, 1788) as defined by Wehrli (1951) and Wehrli (1953 in Wehrli 1939–1954). The correct year of description is 1981, not 1983 as given in Kullberg et al. (2001). Flamigni et al. (2016) treat Parietaria Leraut, 1981 as a junior synonym of Yezognophos Matsumura, 1927, but the synonymy may have been applied earlier. Yezognophos and Parietaria are structurally similar and we follow the synonymy. Unavailable names (incorrect subsequent spelling): Yezonognophos: Bang-Haas (1928).

Diversity and distribution: Four species in Europe (two of those occurring more widely in the Palaearctic region), potentially few more described species in the Palaearctic region as listed in Scoble (1999), but taxonomy of Yezognophos and related genera poorly understood. External characters and abdomen (based on European species): Wingspan ­approximately 30–36 mm, females fully winged. Male forewing costa weakly convex. Wings grey to yellowish brown, irrorated with different shades of brown. Postmedial line most visible, distincly dentate. Terminal line discontinuous. Discal spots large, with pale centre. Markings weaker on wing underside. Male antennae bipectinate, female antennae filiform or weakly fasciculate. Hindlegs of both sexes with 2+2 spurs, male hindtibia slightly swollen. Male and female tympanal organs medium-sized. Male 8th sternite and 8th tergite about same size. Posterior margin of male 8th tergite with folded membrane. Male genitalia (based on European species): Uncus round, triangular or pointed. Socii absent. Gnathos rudimentary, arms do not meet ventrally. Tegumen normal (not massive). Valva wide at base, apex upturned. Costa sclerotized, often with stout spine(s), apex setose. Anterior arms of juxta dentate or antler-shaped. Posterior arms of juxta short, triangular. Saccus concave or weakly elongated. Vesica mostly without cornuti. Anterior arms of juxta offering species-level diagnostic characters. Y. vittaria in many aspects different from the other included species. Female genitalia (based on European species): Lamella antevaginalis often massive, curved around ostium bursae, strongly sclerotized. Lamella postvaginalis often small, membranous. Corpus bursae membranous or with sclerotized ridges. Signum stellate, invaginated medially or absent. Y. vittaria in many aspects different from the other included species. Biology, phenology, habitat: Larvae of Yezognophos species are polyphagous. Larva of Y. ­dilucidaria has been recorded to feed for instance on Gramineae, Sempervivum, Lotus, Achillea and Vaccinium, Y. serotinaria on Alnus viridis, Vaccinium, Lonicera xylosteum and Tanacetum vulgare (Flamigni et al. 2016 and references therein). Larva of Y. vittaria on Betula pubescens and B. nana, Vaccinium uliginosum and V. myrtillus (Ebert 2003; Flamigni et al. 2016 and references therein). The larval development of Y. vittaria potentially takes two years (Mikkola et al. 1989), although there are records of adults from all years. Y. vittaria is a boreo-montane species, whereas the other European species are found in montane habitats mostly, roughly from 700 up to 2800 m in larch, pine and spruce forests, bushy areas, rocky valleys, pastures, alpine grasslands with rocky debris and rocky slopes, depending on altitude. Immature stages: Larvae of Y. dilucidaria, Y. sproengertsi and Y. vittaria are stout, pale brown with longitudinal stripes and rows of sparse black markings (Herzet et al. 2011; Lepiforum 2018; Lepinet 2018). Larvae of Y. dilucidaria, Y. vittaria, Y. serotinaria and Y. ­sproengertsi 195

have two triangular, small dorsal horns on segment A9+A10 (Lepiforum 2018; Lepinet 2018). Pupal cremaster of Y. vittaria with four pairs of small, straight setae, whereas apices are hooked in Y. dilucidaria and Y. serotinaria (Patočka & Turčani 2005). Remarks: The genus-level relationships of structurally, and partly ecologically, similar Elophos Boisduval, 1840, Yezognophos Matsumura, 1927 (Parietaria Leraut, 1981 here regarded as synonym), Sciadia Hübner, 1822, Psodos Treitschke, 1825 (Glacies Millière, 1874 here regarded as synonym), and potentially other related genera need further research, see chapter on Elophos and Yezognophos complex before genus Elophos. Y. vittaria and Y. serotinaria are in many aspects different from other included species and their systematic position needs more research. Recently Yezognophos has been classified in the Gnophini (Vives Moreno 1994; Viidalepp 1996 [as a synonym of Elophos]; Leraut 1997 [as a synonym of Elophos]; Hausmann et al. 2004; 2011a [as a subgenus of Elophos]). Yezognophos has not been included in multi-­ gene analysis so far. Systematic position: Gnophini.

60. Yezognophos dilucidaria (Denis & Schiffermüller, 1775) Geometra dilucidaria Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wienergegend: 315 (Austria: Vienna district). Syntype(s), lost. Phalaena myopata Fabricius, 1794: Ent. Syst. 3 (2): 193 (Italy). Syntype(s). Gnophos dilucidaria var. minor Krulikovsky, 1913: Russk. Ent. Obozr. 13 (1): 102 (Russia: Kirov Oblast: Vjatka). Syntype(s). Valid at subspecific rank (Scoble 1999), further research is needed (see Remarks). Gnophos dilucidaria var. alagnensis Nitsche, 1926: Z. öst. EntVer. 11 (7): 61, pl. 2, fig. 26 (Italy: Piedmont, S. Antonio, Sesiatal). Syntypes 3♂3♀. Validated at subspecies level by Scoble (1999), regarded as junior synonym by Flamigni et al. (2016). Gnophos dilucidaria var. helvetica Nitsche, 1926: Z. öst. EntVer. 11 (7): 61, pl. 2, fig. 28 (Switzerland: Zermatt, Simplon, Furkapass, Alps). Syntypes 2♂4♀. Validated at subspecies level by Scoble (1999), regarded as junior synonym by Flamigni et al. (2016). Gnophos dilucidaria var. silesiaca Nitsche, 1926: Z. öst. EntVer. 11 (7): 61, pl. 2, fig. 25 ([Poland], Silesia: Altvater, Karlsbrunn, Freiwaldau). Syntypes 7♂1♀. Valid at subspecific rank (Scoble 1999), further research is needed (see Remarks). Gnophos dilucidaria var. carpathica Soffner, 1932: Ent. Z., Frankf. a. M. 45: 309: 309, figs 1, 2 (eastern Carpathian mountains, Hoverla district). Syntype(s). Valid at subspecific rank (Scoble 1999), further research is needed (see Remarks). Gnophos dilucidaria var. crenularia Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 613 (Switzerland: Tessin [Ticino], Fusio). Syntype(s). Gnophos dilucidaria generosa Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 613 (Switzerland: South Tessin [Ticino]. Syntypes 30♂♀ (2 ♂ paratypes in NHMUK, examined externally). Validated at subspecies level by Scoble (1999), regarded as junior synonym by Flamigni et al. (2016). Unavailable names (infrasubspecific): abbreviata: Nitsche (1926) (ab.); agglomerata: Nitsche (1926) (ab.); albula: Nitsche (1926) (ab.); blanca: Nitsche (1926) (ab.); brunnea: Vorbrodt & Müller-Rutz (1913) (ab.); designata: Nitsche (1926) (ab.); destrigaria: Nitsche (1926) (ab.); externaria: Nitsche (1926); fuscaria: Nitsche (1926) (ab.); infuscaria: Nitsche (1926) (ab.); interrupta: Hirschke (1910) (ab.); latefasciata: Nitsche (1926) (ab.); lineata: Nitsche (1926) (ab.); mendolensis: Dannehl (1928); obscuraria: Nitsche (1926) (ab.); pupillata: Nitsche (1926) (ab.); restincta: Nitsche (1926) (ab.); retrusata: Nitsche (1926) (ab.); semiobscuraria: Nitsche (1926) (ab.); subfasciaria: Nitsche (1926) (ab.); pulchraria: Nitsche (1926) (ab.); unicolorata: Nitsche (1926) (ab.); unilineata: Nitsche (1926) (ab.).

196

External characters and abdomen: Wingspan 27–36 mm, females often slightly smaller and markings weaker. Wings whitish grey, markings brown to blackish, densely irrorated with blackish grey. Antemedial and postmedial lines distinct, weakly dentate, widest on ­costa. Forewing antemedial line weakly curved inwards. Forewing postmedial line concave near inner margin and distinctly angled inwards before costa. Subterminal line weak, ­often visible on forewing costa only, often outer margin bordered with pale grey. Hindwing post­medial line distinct, subterminal line diffuse or not visible. Terminal line weak, discontinuous. ­Fringes concolorous with wings, uniform. Discal spots large, pale centred. Wings below ­rather uniform grey, forewings distinctly darker. Outer margin of postmedial line bordered with light fascia. Frons, vertex, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Hindtibia with 2+2 spurs in both sexes, male hindtibia slightly swollen. Tympanal organs large in male, almost touching each other medially, female organs medium-sized. Male 8th sternite and 8th tergite about same size. Posterior margin of male 8th tergite with folded membrane. Variation: Very variable. Wings are in most specimens whitish grey, but they can be dark grey, rarely even brownish grey. Density of irrorations variable. Position of medial lines can vary, markings vary from very weak to distinct blackish, often postmedial line is the strongest. There appears to be weak correlation between external appearance and geographical locality: similar specimens can be found from distant places and there are differences in appearances within the same locality. Nitsche (1926) lists 25 different individual or geographical forms. Flamigni et al. (2016) explain the great variability and consider the forms described from the Alpine Arch as junior synonyms. Y. dilucidaria is a widespread species with several putative subspecies as listed above. DNA barcodes from central European specimens are rather homogeneous (see Genetic data), but no data was available from the peripheral areas of distribution. These need to be evaluated critically, which was not done here, and those may turn out to be junior synonyms. Therefore we repeat them as given in Scoble (1999), but do not give descriptive accounts and do not recognize them in the distribution map. Male genitalia: Uncus roundish, apex weakly concave. Socii absent. Gnathos plate-like, fused ventrally. Valva wide at base, apex upturned, narrow. Costa sclerotized, with stout spine. Subcostal ridge setose. Anterior arms of juxta very long, apex strongly dentate. Posterior arms of juxta short, triangular. Saccus round. Aedeagus fused to juxta via sclerotized structure. Aedeagus small, curved ventrally, caecum short. Vesica opens at about 135 degree angle, base widened, with longitudinal grooves, without cornuti. Female genitalia: Papillae anales round, setose. Terminal segments very long, telescopic. Apophyses posteriores about twice length of apophyses anteriores, with subapical enlargement. Lamella antevaginalis massive, sclerotized, particularly on margins. Posterior margin weakly concave, sterigma distinctly v-shaped and sclerotized. Lamella postvaginalis small, sclerotized, folded in transverse direction. Ductus bursae short, membranous. Posterior part of corpus bursae long and narrow, anterior part roundish. Signum small, weakly stellate. Distribution and abundance: European. In the Carpathians from southern Poland and northern Slovakia through northern Czech Republic and south-eastern Germany to the Alps and the eastern mountains of France. Also recorded from a few places outside this area, especially in the Balkan peninsula. An old and doubtful record from French eastern Pyrenees (Mazel & Peslier 1997), and several other old records from Spanish Pyrenees, referring to misidentifications according to Redondo et al. (2009). Erroneously recorded for Romania, 197

Portugal, central and southern Spain on the map in Leraut (2009). Y. dilucidaria is mentioned from several places in European Russia (Mironov et al. 2008), but these records are misidentified Y. vittaria (Mironov pers. comm.). – Local, but rather common, sometimes found in numbers.

Phenology: Univoltine. From early June to late August. Caterpillar from late July to June (D. Bolt pers. comm.), hibernation as small larvae (M. Leipnitz pers. comm.). Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Potentilla, Vicia and Lotus, close to the ground (M. Leipnitz pers. comm.). Recorded on Sempervivum, Lotus, Hypericum, Helianthemum, Laserpitium, Gentiana, Linaria, Achillea, Taraxacum, Hieracium, Genista, Cistus and Vaccinium, especially V. myrtillus (Flamigni et al. 2016). Reared on Lonicera tatarica, Saxifraga decipiens, Sanguisorba minor and Taraxacum (M. Leipnitz pers. comm.). Pupation among littered leaves on the ground or in the soil in a thin cocoon (Z. Las˘tu˚ vka pers. comm.). Habitat: Montane, alpine. In a variety of habitat types, both below and above treeline. Below treeline in open places as well as open forests. According to Kerschbaum & Pöll (2010) from 200 up to 2400 m, while reported from 800 up to 2800 m by Flamigni et al. (2016). Similar species: Yezognophos vittaria (Thunberg, 1788) more brownish (subsp. mendicaria Herrich-Schäffer), dots on terminal line more distinct, on underside postmedial line more distinct and its outer margin weakly bordered with paler fascia, frons blackish (in Y. diluci­ daria wings whitish grey, dots on terminal line less distinct, on underside postmedial very weak and its outer margin bordered with rather distinct pale fascia, frons concolorous with wings) (Text-figs 102–103). Genitalia allowing reliable identification. Males of genus Yezo­ gnophos easily recognizable by their bipectinate antennae, which are filiform in the externally similar genera Gnophos Treitschke, Odontognophos Wehrli and Charissa Curtis. 198

Text-figs 102–103. Diagnostic characters (indicated) of Yezognophos dilucidaria (Denis & Schiffer­ müller, 1775) and Y. vittaria (Thunberg, 1788). The vertex and frons can be irrorated with brown or blackish scales in Y. dilucidaria (being concolorous with wings), whereas the frons is distinctly dark (blackish) in Y. vittaria throughout its distribution area. See text for additional diagnostic features. Text-fig. 102. Y. dilucidaria, Austria: St. Jakob im Lesachtal, 1800 m, 8.viii.1999 (coll. TLMF). Text-fig. 103. Y. vittaria, Austria: St. Jakob im Lesachtal, 1650 m, 2.iv.2001 (ex ovo) (coll. TLMF).

Genetic data: BIN: BOLD:AAC9509 (n=20 from Germany, Austria, Italy). Genetically homogeneous. Nearest species: Y. sproengertsi (5.1%), S. unicoloraria (6.0%), Y. vittaria (7.4%).

61. Yezognophos sproengertsi (Püngeler, 1914) Gnophos sproengertsi Püngeler, 1914: Dt. ent. Z. Iris 28: 51, pl. 3, figs 17, 25 (France: Alpes maritimes: St. Martin-Vésubie). Syntype(s), including ♂♀ (examined externally, based on photographs in the original publication). Gnophos canitiaria f. subpullata Wehrli, 1924: Dt. ent. Z. Iris 38: 88, pl. 1, fig. 7 (France: Alpes maritimes. Syntypes 1♂1♀ (ZFMK). Junior synonym (Flamigni et al. 2016). Gnophos sproengertsi dinica Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 614, pl. 51a (France: Alpes-de-Haute-Provence: forest of Entraunes; Larche). Syntypes, including ♂. Valid at subspecific rank (Scoble 1999), but potentially falling within the variation of the nominotypical subspecies. Subspecies not illustrated in the distribution map. More research needed. Unavailable names (incorrect subsequent spelling): spröngeri: Wehrli (1951).

External characters and abdomen: Wingspan 26–30 mm. Wings pale yellowish grey, markings and striations pale (olive) grey to brown. Antemedial line weak, postmedial line distinct, both widened into blackish distinct spot on costa. Postmedial line deeply dentate, angled inwards before costa, its outer margin bordered with paler area. Subterminal line weak, appearing pale in terminal area, which is often rather uniform grey, particularly near costa. Terminal line blackish, discontinuous. Fringes concolorous with wings, uniform. Forewing discal spots large, roundish, hindwing spots weakly elongated, both pale-centred. Wings below uniform dark grey, except outer margin of postmedial line pale, wide. Hindwings below slightly paler than forewings. Discal spots weak, diffuse. Frons, vertex, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Hindtibia with 2+2 spurs in both sexes, male hindtibia slightly swollen. Tympanal organs medium-sized in both sexes. Male 8th sternite and 8th tergite about same size. Posterior margin of male 8th tergite with folded membrane. 199

Variation: Rather little. Density of and extent of irrorations and striations variable. Forewing medial area sometimes paler, concolorous or darker than rest of wing, and if the latter, pale area on its outer margin standing out. Discal spots pale-centred in most specimens, but sometimes solid. Wing colour varying from olive-yellowish grey to pale grey; old specimens tend to loose the olive-yellowish tint, becoming rather uniform grey. Tentative subspecies ­dinica Wehrli paler (from France: Alpes-de-Haute-Provence, near Digne), but more research is needed. Male genitalia: Uncus elongated. Socii absent. Gnathos reduced, only weak membranous lateral arms present. Valva wide at base, with sclerotized margin, apex upturned and narrow. Costa sclerotized, with enlarged lobe in middle, bearing 3–4 stout horns. Subcostal ridge setose. Anterior arms of juxta large, inner margin strongly sclerotized and spined, dorsal part with membranous sac. Posterior arms of juxta short, elongated. Saccus short, concave. Aede­agus small, curved dorsally, caecum round, apex with small tooth. Vesica open at about 135 degree angle, base weakly enlarged, with weak longitudinal grooves, without cornuti. Female genitalia: Papillae anales round, setose. Terminal segments not elongated. Apophyses very narrow, apophyses posteriores about twice length of apophyses anteriores. Lamella antevaginalis very large, made up of two curved sclerotized plates, which are separated by sutura ventrally. Lamella postvaginalis small, heavily sclerotized, dorsal margin roundish. 8th tergite sclerotized, rather widely or along its margin only, extending ventrally and forming sclerotized ring that passes ostium bursae anteriorly (see Genitalia plate for variation). Ductus bursae very short, membranous. Corpus bursae large, elongated. Signum small elongated plate, on dorsal side, with horizontal invagination medially. Distribution and abundance: European. Very limited distribution, found in the south-western Alps only in the border area of south-eastern France and north-western Italy. In France recorded from Alpes-Maritimes, Alpes-de-Haute-Provence, Hautes-Alpes, Isère and Drôme

200

(Herzet et al. 2011; Lepinet 2018), in Italy recorded from Piedmont and Liguria (Flamigni et al. 2016). – Rather rare and local, usually found in limited numbers, under good weather conditions sometimes in numbers. Phenology: Univoltine. Apparently with relatively short flight period, from mid-July to early August. Larva from July, hibernating to April or May (M. Leipnitz pers. comm.; rearing observation). Adults active at night, both sexes readily attracted to light. Biology: Larva potentially polyphagous. Reared on Taraxacum and Daucus carota (M. Leipnitz pers. comm.). Pupation in a light silk web between moss and plant litter on the ground (M. Leipnitz pers. comm.; rearing observation). Habitat: Montane, alpine. Found in open meadows above treeline as well as in localities with scattered trees and bushes, often in places with rocks and stones. From 1700 up to 2400 m. Similar species: E. dilucidaria (Denis & Schiffermüller) with grey wings, postmedial line less dentate, in Y. sproengertsi wings pale yellowish grey and postmedial line more dentate. Male and female genitalia allow reliable identification. Several species of Charissa Curtis, 1826 externally similar, particularly C. glaucinaria (Hübner, 1799), but male antennae filiform in C. glaucinaria (bipectinate in Y. sproengertsi), dark terminal area of wing underside often with pale blotch in C. glaucinaria (without blotch in Y. sproengertsi). Genetic data: BIN: BOLD:AAK9535 (n=3 from France, Italy). Genetically homogeneous. Nearest species: S. zelleraria (4.6%), Y. vittaria (5.1%).

62. Yezognophos vittaria (Thunberg, 1788) Phalaena vittaria Thunberg, 1788: D. D. Mus. nat. Acad. Upsala 6: 74, figured (Sweden: Upp­ land, Upsala). Lectotype ♂ (Universitets Zoologiska Institut, Uppsala, Sweden), paratype ♂ [without abdomen] (NHMUK, examined externally). According to Koçak (1984: 108; 1986: 33) Phalaena vittaria Thunberg, 1788 should be preoccupied by P. vittaria Sulzer, 1776. However, this seems to be a mistake as Sulzer, as far as it has been possible to trace, never did describe Phalaena vittata (M. Scoble in litt. in Müller 1996). Phalaena sordaria Thunberg, 1792: Diss. ent. sistens Insecta Suecica 4: 60, pl. 4, figured (Sweden: Gestricia [=Gästrikland]). Lectotype ♂ (Universitets Zoologiska Institut, Uppsala, Sweden). Gnophos mendicaria Herrich-Schäffer, 1852: Syst. Bearb. Schmett. Eur. 6 (55): 72; ibidem (1851) 3: pl. 80, figs 491–493 [non binominal] (the Alps) Syntype(s). Valid at subspecific rank. Yezognophos kononis Matsumura, 1927, Insecta matsum. 1 (4): 187 (Japan: Hokkaido, Mt Daisetsu, Koizumidake). Holotype ♂. Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): distincta: Strand (1902) (ab. of sordaria); strandiata: Fuchs (1903) (ab. of sordaria); radiata: Hirschke (1910) (ab. of mendicaria).

External characters and abdomen: Wingspan 31–36 mm, males slightly larger on average. Wings pale beige, or very pale yellowish, weakly irrorated with brown, postmedial line and discal spots most pronounced, wider on veins, brown. Antemedial line weak. Postmedial line usually well-developed, weakly dentate, curved inwards before costa, where widened. Forewing medial area concolorous with wings, narrower on inner margin. Terminal line distinct, discontinuous. Fringes concolorous with wings. On hindwings often postmedial line visible only. Discal spots distinct, pale-centred. Wings below suffused, slightly darkened from base to postmedial line, which is pale-bordered. Discal spots diffuse, terminal line distinct. 201

Text-figs 104–107. Males of Yezognophos vittaria subsp. vittaria (Thunberg, 1788) and Y. vittaria subsp. mendicaria (Herrich-Schäffer, 1852). In the nominotypical subspecies (northern Europe) vesica opens at about 135 degree angle and in subspecies mendicaria (the Alps) at about 90 degree angle. Although the vesica angle appears diagnostic between northern European and Alpine specimens, the angle is not always easy to interpret due to the dissection artefacts or because the angle may be transitional between the mentioned extremes, see Remarks. Nominotypical subspecies is characterized by pale beige wings and brown markings, whereas subspecies mendicaria has grey or whitish grey wings and blackish markings in the Alps. Not enough material is studied to understand the geographical variation of the morphological and molecular structures, see also genetic data. Text-fig. 104. Y. vittaria vittaria, Finland: Kaunispää, 4.vii.1994, all genitalia figures from the same specimen, slide Sihvonen 2080 (coll. Sihvonen). Text-fig. 105. Y. vittaria vittaria, Finland: Enonkoski, 6–7.vii.1996, all genitalia figures from the same specimen, slide Sihv­ onen 2181 (coll. Sihvonen). Text-fig. 106. Y. vittaria mendicaria, Italy: Bozen, Ritten, 1750 m, 1.vii.2010, all genitalia figures from the same specimen, slide Sihvonen 2110 (coll. TLMF). Textfig. 107. Y. vittaria (subspecies uncertain), Kasakhstan: South Altai, Burkhat Pass, 2100–2500 m, 11–15.vii.1998, all genitalia figures from the same specimen, slide Sihvonen 2180 (coll. Skou). %

Frons distinctly dark (blackish), vertex, collar, thorax and abdomen concolorous with wings. Antennae bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Tympanal organs small in both sexes. Male 8th sternite smaller than 8th tergite. Other sternites and tergites of both sexes undifferentiated. Variation: Terminal area often darker than rest of wing. Discal spots often pale-centred, but can be solid. See subsp. mendicaria. Male genitalia (Text-figs 104–107): Uncus apex narrow, elongated. Socii absent. Gnathos arms weakly sclerotized, separate. Tegumen large, with sutura mid-dorsally. Valva very wide at base, outer margin weakly concave subapically, apex upturned. Valva costa sclerotized, with several stout setae, with distinct angle subapically, apex setose. Anterior arms of j­uxta large, apex antler-shaped, inner margin of subapical part concave. Posterior arms of ­juxta small, short, hollow, horn-shaped. Saccus elongated, curved dorsally. Aedeagus wide medially, apex with very large tooth. Vesica opens at about 135 degree angle in Fennoscandian specimens, at about 90 degree angle in the Alps and in Kazakhstan (see Text-figs 104–107). Vesica very large, roughly rectangular in shape, apex with row of about 5–6 large cornuti. Ductus ejaculatorius opens from upper end of cornuti row, directed downwards. Female genitalia (Text-figs 108–111): Papillae anales round, setose. Apophyses anteriores short, less than half length of apophyses posteriores. Lamella antevaginalis (homology difficult to interpret) narrow, semi-lunular plate, curved ventrally and fused with sclerotized corpus bursae. Lamella postvaginalis (homology difficult to interpret) small, separate, doughnut-shaped, sclerotized, curved structure around ostium bursae. Ductus bursae very short, almost absent. Corpus bursae elongated, heavily sclerotized, with two or three long distinct ridges, except anterior part, being wrinkled and membranous. No definite signum, but gorge-shaped, elongated concavity mid-ventrally reminescent of such structure. Ductus seminalis opening mid-dorsally from corpus bursae. Subsp. mendicaria slightly bigger, grey, or even whitish grey, heavily irrorated with dark brown or blackish (particularly near wing margins). Antemedial line often well-developed, postmedial line usually more distinct, wider on veins. Discal spots distinct. Wings below heavily irrorated with blackish. Genetically distinct from the Fennoscandian specimens, maximum barcode distance 6.5% (Mutanen et al. 2012), but no morphological differences observed in 202

the genitalia, except the angle at which vesica opens, see Text-figs 104–107. Distribution of taxon mendicaria unknown in detail, described from the Alps and reported to occur in the central European mountains including the Carpathians (Levente 2011; ­Malkiewicz 2012) and the Dinaric Alps (Flamigni et al. 2016). Barcodes available from Fennoscandia and Alps only, so far. More research is needed. 203

Text-figs 108–111. Females of Yezognophos vittaria subsp. vittaria (Thunberg, 1788) and Y. vittaria subsp. mendicaria (Herrich-Schäffer, 1852). Unlike the difference found in the male genitalia, no constant differences were observed in the female genitalia. See also genetic data. Circle indicates the point of origin of the ductus seminalis. Text-fig. 108. Y. vittaria vittaria, Norway: Vestlige Finnmark), Burfjord, 200–300 m, 20.vii.1998, slide Sihvonen 2081 (coll. Sihvonen). Text-fig. 109. Y. vittaria vittaria, Finland: Enonkoski, 10–16.vii.1996, slide Sihvonen 2082 (coll. Sihvonen). Text-fig. 110. Y. vittaria mendicaria, Italy: Bozen, Ritten, 1750 m, 1.vii.2010, slide Sihvonen 2111 (coll. TLMF). Text-fig. 111. Y. vittaria vittaria, Switzerland: Bern, Susten Pass, 2100 % m, 12. vii. 2015, slide Sihvonen 2178 (coll. Skou).

Distribution and abundance: Eurasian, in Europe boreo-montane. From the Ural mountains across northern European Russia to Fennoscandia and in the Carpathians, Alps and Jura mountains, and further south with isolated occurrences in the Balkan peninsula. – Outside Europe east of the Urals across southern Siberia to Magadan Province and Japan (subsp. kononis). – In central and northern Fennoscandia common, often found in numbers, to the south more local and rare. In the Alps somewhat local, usually in limited numbers.

Phenology: Univoltine. In Fennoscandia from early June to late July. In the Alps from early June to late July or early August. Most records are from June to early July. Larva in the Alps from August to May, hibernating. In northern Fennoscandia larvae overwintered first as a small larva, and during second winter as a full-grown larva (Nordman 1941; rearing observation). In south-western Germany a 1 cm long overwintered larva found in March, but without information on the further development and the question of annual or bi­ennial life-­cycle (Ebert 2003). Adults active at night, especially the males attracted to light. In Fenno­scandia, adults mainly found by flushing them up by day. 204

Biology: Larva in Fennoscandia monophagous on Betula, in the mountains on B. nana. In Central Europe polyphagous. Recorded on Betula, Campanula, Clematis, Vaccinium myrtillus and Vicia (Ziegler 2019). Also reared on V. myrtillus (Ebert 2003). Pupation in a light silk web between moss, plant litter and soil (M. Leipnitz pers. comm.). 205

Habitat: Silvicolous. In Fennoscandia in open deciduous forests and mountain heaths. In Central Europe in a variety of habitat types such as moors, mountain river valleys, conifer forests with Vaccinium myrtillus and in the Alps also above treeline. In Fennoscandia from close to sea-level up to 700 m, in the Alps from 1000 up to 2500 m. Similar species: Yezognophos dilucidaria (Denis & Schiffermüller, 1775), see description and Text-figs 102–103 under that species. The genitalia allow reliable identification. Genetic data: BINs: BOLD:AAD3010 (n=20 from Germany, Austria, Switzerland, Italy), BOLD:AAD3009 (n=5 from Finland, Sweden, Norway), both BINs diverging by 5.9%. Genetically heterogeneous, cluster BOLD:AAD3010 with maximum variation 2.0%. Nearest species: Y. sproengertsi (5.1%), E. caelibaria (6.2%). See also Mutanen et al. (2012), who report maximum genetic distance of 6.5% between the Fennoscandian and the Alpine specimens. Remarks: Y. vittaria is in many aspects different from the other species included in the genus Yezognophos, indicating that its systematic position needs further research. Homologies in the female genitalia around the ostium bursae are difficult to interpret. Numerous specimens were examined morphologically to understand the diagnostic value of the vesica angle in separating the nominotypical subspecies and subspecies mendicaria, and to understand whether taxon mendicaria should be treated as valid at species level. In the nominotypical subspecies (northern Europe) vesica opens at about 135 degree angle and in subspecies mendicaria (the Alps) at about 90 degree angle (Text-figs 104–107). In Kazakh­stan the angle is about 90 degrees (see Text-fig. 107, slide Sihvonen 2180, coll. Skou), in Poland: Sudetes close to 90 degree (see Malkiewicz 2012), in Russia: Buryatia close to 135 degrees (slide Sihvonen 2193, coll. ZMH), and in Russia: Altai close to 135 degrees (slide Sihvonen 2194, coll. ZMH). There is some variation between the mentioned extremes, which may be due to dissection artefacts or the character is transitional. No diagnostic differences were found in the female genitalia, specimens were examined from Europe (several specimens, see Text-figs 108–111) to Russia (Irkutskaja oblast, slide Sihvonen 2195, coll. ZMH). From the biogeographical point of view it could be possible that taxon vittaria is widespread from Northern Europe to Siberia, and taxon mendicaria could be restricted to the central European mountains only. The specimen from Kazakhstan, however, blurs the picture because its morphology is more similar to mendicaria (dissection artefact?). Taxon kononis ­Matsumura, 1927 from Japan was not examined. We consider taxon mendicaria valid at subspecies level (alternative would be to consider it valid at species level) until more extensive research is carried out, including DNA barcodes throughout the distribution area of the species.

63. Yezognophos serotinaria (Denis & Schiffermüller, 1775) Geometra serotinaria Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 315 (Austria: Vienna district). Syntype(s). Gnophos serotinaria dognini Thierry-Mieg, 1910: Naturaliste 32: 154 (France: Pyrénées-­ Orientales, Pic du Canigou). Syntype(s) ♂. Flamigni et al. (2016) considered taxon dognini a junior synonym of taxon serotinoides Wehrli, whereas Leraut (2009) considered it valid at subspecies rank. We consider it valid at subspecific rank as Y. serotinaria dognini. Gnophos serotinaria aenearia Oberthür, 1913: Études Lépid. comp. 7: 311, pl. 481, figs 1775, 1776 (France: Pyrénées-Orientales, Cady valley; Casteil; Mariailles). Syntypes ♂♀. Junior synonym of serotinaria Denis & Schiffermüller according to Scoble (1999), but most likely a junior synonym of dognini Thierry-Mieg (as in Leraut 2009; Vives Moreno 2014 and Flamigni et al. 2016), judging from the type localities in the Pyrenees.

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Gnophos serotinaria f. valdesignaria Wehrli, 1922: Schweizer ent. Anz. 1 (4): 28, fig. 3 (Austria: Kaltenberg; Switzerland: Bevers 1570 m; Davos). Syntype(s). Considered valid at subspecies level by Scoble (1999), but recently considered a junior synonym of taxon serotinaria Denis & Schiffermüller, 1775 (Flamigni et al. 2016). Gnophos dognini distinctior Wehrli, 1922: Schweizer ent. Anz. 1 (4): 27, figs 8, 9 (Switzerland: Campolongo pass; Fusio). Syntypes ♂♀ (ZFMK). The specimens from Ticino must be excluded from the type series, because the author refers to them as a distinct variant (by name) (ICZN, Article 74.2.1). Considered valid at subspecies level by Scoble (1999), but recently considered a junior synonym of taxon serotinoides Wehrli, 1922 (Flamigni et al. 2016). Gnophos serotinoides f. griseisparsa Wehrli, 1922: Schweizer ent. Anz. 1 (4): 27, figs 6, 7 (Switzer­ land: Valais, Zermatt, Täsch, Arolla 2000 m, Simplon). Syntypes 15♂8♀ (ZFMK). The specimens from Valais must be excluded from the type series, because the author refers to them as a distinct variant (by name) (ICZN, Article 74.2.1). Considered valid at subspecies level by Scoble (1999), but recently considered a junior synonym of taxon serotinoides Wehrli, 1922 (Flamigni et al. 2016). Gnophos serotinoides Wehrli, 1922: Schweizer ent. Anz. 1 (4): 26 (France: Alpes-de-HauteProvence, Digne; Alpes-Maritimes, Mont Berceau near Menton). Syntypes 13♂7♀. The type localities given in Scoble (1999), including Russia, are incorrect (Flamigni et al. 2016). Valid at subspecific rank as Y. serotinaria serotinoides. Parietaria serotinaria lozerica Leraut, 2009: Moths of Europe, vol. 2: 238, pl. 65, fig. 12 (France: Lozère, Bagnols-les-Bains. Holotype ♂ (MNHN), gen. prep. Leraut # 12240. Leraut does not not illustrate the genitalia of lozerica, which is potentially a junior synonym of taxon sero­ tinoides Wehrli, 1922, judging from the superficial written description and the type locality near western end of the Alps. Parietaria serotinaria vesubiaria Leraut, 2009: Moths of Europe, vol. 2: 238, pl. 65 figs 13–15, fig. 127a (France: Alpes-Maritimes, La Brigue (Morignole)). Holotype ♂ (MNHN), gen. prep. Leraut 10472. Junior synonym of taxon serotinoides Wehrli, 1922 (Flamigni et al. 2016). Potentially unavailable names (infrasubspecific): fumipennaria: Galvagni (1917); guillaumei: Leraut (2009) (f.); tenebraria: Wagner (1910); umbrosaria: Wehrli (1922).

External characters and abdomen: Wingspan 28–38 mm. Wings pale yellowish, irrorated with brown, markings brown. Females often slightly paler. Forewing costa rounded near apex. Ante- and postmedial lines distinct, dentate, curved inwards on costa. Subterminal line weak, often visible near costa only, or absent. Terminal line distinct, row of blackish dots. Fringes concolorous with wings. Discal spots distinct, pale-centred. Wings below more uniform, darker, postmedial line most pronounced. Frons, collar, thorax and abdomen of both sexes concolorous with wings. Antennae densely bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Tympanal organs medium-sized, slightly smaller in females. Male 8th sternite smaller than 8th tergite. Male 8th lateral membrane with two small, elongated structures inside the abdomen. Other sternites and tergites of both sexes undifferentiated, more sclerotized in male. Variation: Variable species, but external appearance rather weakly correlates with locality. Brown irrorations can be rather few, particularly in females that occasionally appear almost uniform pale brown, or irrorations can be dense, especially near terminal area, giving smoky appearance. Although the specimens in a single locality tend to be externally similar to each other, there are aberrant specimens, so the wing colour and pattern do not have diagnostic value in separating the subspecies. Male genitalia: Uncus base wide, apex narrow and pointed. Socii absent. Gnathos absent (or very rudimentary). Tegumen large, elongated. Valva upturned, very wide at base, outer margin distinctly concave. Costa sclerotized, with erect sharp tooth, basal part bare, apical 207

Text-figs 112–120. Diagnostic male genitalia characters (indicated) of Yezognophos serotinaria (Denis & Schiffermüller, 1775). The aedeagus is considerably longer in Y. serotinaria serotinaria than in other subspecies. Figures are not in scale. Text-figs 112–113. Italy: South Tyrol, Vinschgau, 1300 m, 2.iv.1986, ex ovo, slide Sihvonen 2114 (coll. Sihvonen). Text-fig. 114. Austria: Tirol, Umhausen, 4.vii.1945, slide Sihvonen 2189 (coll. TLMF). Text-fig. 115. France: HautesPyrénées, Lac d’Orédon, 1900 m, 24.vii.2014, slide Sihvonen 2185 (coll. Skou). Text-fig. 116. Spain: Lerida, 6 km SE Coll. del Cantó, 1460 m, 16.vii.2003, slide Sihvonen 2187 (coll. Skou). Text-fig. 117. France: Hautes-Pyrénées, Lac d’Orédon, 1900 m, 24.vii.2014, slide Sihvonen 2185 (coll. Skou). Text-fig. 118. Switzerland: Valais, Saastal, 1900 m, 13–14.vii.2015, slide Sihvonen 2183 (coll. Skou). Text-fig. 119. France: Alpes-Maritimes, Vallée des Merveilles, 14.vii.1934, slide Sihvonen 2176 (coll. TLMF). Text-fig. 120. Switzerland: Valais, Saastal, 1900 m, 13–14.vii.2015, % slide Sihvonen 2183 (coll. Skou).

part setose. Ventral margin of costa with row of small teeth. Anterior arms of juxta large, long, apically covered with minute sclerotized teeth, apex acute. Posterior arms of juxta rather long, hollow, tapering towards apex. Saccus small, weakly concave. Apical part of aedeagus very long, narrow, curved ventrally, apex smooth or with minute teeth and separate distinct tooth (Text-figs 112–120). Aedeagus with ventral lobe and basal part wide, its length variable. Vesica opens at 45–135 degree angle (see subspecies below), base weakly widened, with longitudinal grooves, cornuti absent. Size of aedeagus and its curvature vary considerably. Female genitalia: Papillae anales round, setose. Terminal segments elongated. Apophyses anteriores short. Lamella antevaginalis massive, sclerotized, with longitudinal grooves. Its posterior margin with large, incurved hooks and deep constriction medially, latter paralled-­ sided or v-shaped (Text-figs 121–126). Lamella postvaginalis weakly sclerotized medial plate, often broken into two parts, or membranous. Ductus bursae sclerotized, its posterior part long, followed by distinct dorsal angle, its anterior part shorter. Corpus bursae pearshaped, large. Signum small, stellate, situated near anterior end. Size of genitalia varying considerably, lamella antevaginalis rounded or angled, slightly rectangular in shape. Subsp. dognini is diagnosable by the genitalia structures (Text-figs 115–117; 123–124). Male with wide vesica base, opening at 90 degree angle, spike at aedeagus apex almost straight, spike length to aedeagus length ratio approximately 1:4, ventral margin often smooth (without spines), aedeagus narrow, base of aedeagus long, caecum long. Apex of ventral arm of juxta rounded. Female with rounded margin of lamella antevaginalis, ductus bursae narrow and long. Type locality of taxon dognini is in the Pyrenees, but occurrence in the Alps is not excluded. More research is needed. See Remarks. Subsp. serotinoides is diagnosable by the genitalia structures (Text-figs 118–120; 125– 126). Male with wide vesica base, opening at 45 degree angle, spike at aedeagus apex almost straight, spike length to aedeagus length ratio approximately 1:3, ventral margin often spinose, aedeagus thick, base of aedeagus long, caecum long. Apex of ventral arm of juxta acute. Female genitalia with v-shaped margin of lamella antevaginalis, ductus bursae wide and short. According to current knowledge taxon serotinoides is restricted to the western part of the Alps, but occurrence elsewhere in the Alps is not excluded. Flamigni et al. (2016, p. 346) diagnose subsp. serotinoides as “the sclerotized part of the ductus bursae is shorter in serotinoides; in some Apennine specimens it is longer, but its sides are parallel up to the end, without any narrowing.” We find that such specimens fall within variation of subsp. 208

sero­tinaria. Our material indicates that subsp. dognini and subsp. serotinoides are similar, both having short and wide ductus bursae, correlating with length and width of male aedeagus. See Remarks, more research is needed. 209

Text-figs 121–126. Diagnostic female genitalia characters (indicated) of Yezognophos serotinaria (Denis & Schiffermüller, 1775). The ductus bursae is considerably longer in Y. serotinaria serotinaria than in other subspecies. Figures are not in scale. Text-fig. 121. Italy: Vinschgau, Sulden 1900 m, 28.v.1996 ex ovo, slide Sihvonen 2115 (coll. Sihvonen). Text-fig. 122. Austria: Ost­tirol, Ainet 750 m, 18.vii.1985, slide Sihvonen 2175 (coll. TLMF). Text-fig. 123. France: Hautes-Pyrénées, Lac d’Orédon, 1900 m, 24.vii.2014, slide Sihvonen 2186 (coll. Skou). Text-fig. 124. Spain: Lerida, Sort, 7.vii.1993, 1000 m, slide Sihvonen 2188 (coll. Skou). Text-fig. 125. Switzerland: Wallis, Leuk-Guttet, 1000 m, 14.xi.1991 ex pupa, slide Sihvonen 2131 (coll. TLMF). Text-fig. 126. France: Alpes-Maritimes, Thorenc, 18.vii.1934, slide Sihvonen 2177 (coll. TLMF).

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Distribution and abundance: European. Mountain species occurring in the eastern Pyrenees and in the Alps from eastern France to eastern Austria and Slovenia, and from southern Germany to northern Italy. Three old records from Romania doubtful, not having been confirmed (Székely 2011). Old records from the Volga-Ural region misidentified, as noted by Anikin et al. (2000). Subspecies serotinoides tentatively restricted to western part of the Alps and northern Italy. Subspecies dognini tentatively restricted to eastern Pyrenees. – Relatively common but somewhat local, sometimes found in numbers.

Phenology: Univoltine. From mid-June to mid-August, according to Flamigni et al. (2016) until late August. Larva from August to May, hibernating as small larva (M. Leipnitz pers. comm.; rearing observation). Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Recorded on Alnus viridis, Vaccinium, Lonicera, Tanacetum vulgare, Taraxacum officinale, Acer campestre, Lotus corniculatus, Onobrychis, Prunus spinosa, Rosa and Prunus domestica (Flamigni et al. 2016). Reared on Lonicera tatarica, Daucus carota, Sanguisorba minor, Taraxacum (M. Leipnitz pers. comm.) and Genista (D. Fritsch pers. comm.). Pupation in a light silk web between moss, plant litter and soil (M. Leipnitz pers. comm.). Habitat: Montane. In a variety of mountain habitats such as meadows, places with scattered trees and bushes, open forests, often places with open rock or stones. In the southern and western Alps and in the Pyrenees from 1000 up to 2000 m. In Austria from 600 to 1600 m (Kerschbaum & Pöll 2010). Similar species: Pale yellowish wings and brown markings usually diagnostic, allowing unambiguous identification. Worn specimens may be confused with Y. vittaria (Thunberg). Male and female genitalia are diagnostic. 211

Genetic data: BIN: BOLD:AAX7664 (n=12 from northern Italy, Austria, Slovakia) genetically comparatively homogeneous, populations of Y. s. serotinoides from southern Piedmont (‘Appennino piemontese’, northern Ligurian Alps) close to haplotypes of nominotypical subspecies but populations from Val di Antrona, Hautes-Alpes and western Liguria closer to Iberian subsp. dognini and not BIN-sharing. Other BIN: BOLD:AAX7664 (n=5 from Spain, France, north-westernmost Italy) genetically homogeneous, genetic distance between both BINs 1.1%. Remarks: Yezognophos serotinaria-serotinoides-dognini is a taxonomically difficult complex and these taxa have been considered valid at various taxonomic ranks in different publications. Our conclusion, one species with three subspecies, is based on the interpretation of the material throughout the distribution area, and includes examination of the genitalia and DNA barcodes. Other options could have been to classify the complex into one species with two subspecies, namely Y. serotinaria serotinaria and Y. serotinaria dognini (the latter including taxon serotinoides also) or to consider these three taxa valid at species level. We chose to consider the populations in the western Alps (taxon serotinoides) and Pyrenees (taxon dognini) valid at subspecies level, because the populations are allopatric, diagnosable by small differences in the male and female genitalia, but the conclusion is obscured by the fact they share the same BIN and the barcodes are tangled. We note, however, that specimens in Switzerland (and potentially elsewhere) are somewhat transitional, obscuring the boundaries between the mentioned subspecies. To further complicate matters, DNA barcodes do not always correlate with the morphology in this complex. Flamigni et al. (2016) noted this also, and explain such cases in detail. Mutanen et al. (2016) found that Elophos serotinaria and E. dognini were tangled with each other in COI gene trees and considered them a “parapatric species with a narrow hybrid zone in Switzerland and NW Italy”. The genitalia size varies considerably, this is more notable in females (see genitalia plate). Y. serotinaria is structurally different from other Yezognophos species and its systematic position needs to be studied in an analytical context. More research is needed.

Crocota Hübner, 1823 Crocota Hübner, 1823, Verz. bekannter Schmett.: 295. Type species: Geometra tinctaria Hübner, 1799. Cleogene Duponchel, 1829, in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 7 (2): 109. Type species: Geometra tinctaria Hübner, 1799. [A junior objective synonym of Crocota Hübner, 1823].

Diversity and distribution: Crocota contains four species in the western Palaearctic region (Scoble 1999; Leraut 1999, but see Remarks), the occurrence is restricted to the south European mountains, mostly in the Pyrenees and the Alps. External characters and abdomen: Medium-sized moths, wingspan approximately 22–35 mm, females smaller and often with different color than males (sexual dimorphism). Wings uniform yellow, grey or white, finely irrorated with iridescent or dark scales, creating shiny appearance, particularly on underside. Very narrow terminal line present on underside. Distinct discal spots present in one species (C. peletieraria males). Male antennae bipectinate, female antennae filiform. Male hindlegs unmodified, with 2+2 spurs. Male 2nd sternite medially weakly sclerotized, 8th sternite very small (about half size of 8th tergite). Postero­ lateral parts of tergites weakly sclerotized in both sexes (Tex-fig. 127). Female tympanal organs very small.

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Text-fig. 127. Diagnostic abdominal characters (indicated) found in Crocota Hübner, 1823, illus­ trated on C. tinctaria (Hübner, 1799), Italy: Prov. Cuneo, Alpi Cozie, Demonte NW, 2420 m, 2.viii.2010, slide Sihvonen 2101 (coll. TLMF). Small male 8th sternite correlates with dorso­ ventrally large tegumen. These structures are the most pronounced in this genus but not unique as similar features are present in some species of Sciadia Hübner, 1822 and Elophos Boisduval, 1840 also, particlarly in species that have short-winged females. More research on the relationships of these genera is needed.

Male genitalia: Uncus wide with two distinct horns laterally, or triangular with concave apex. Posterior arms of juxta short, hollow, slightly triangular. Anterior arms of juxta absent. Posterior part of juxta plate curved dorsally, fused with aedeagus. Valva wide, costa sclerotized with few stout setae, apex setose. Aedeagus apex with distinct tooth or with numerous micro­teeth. Vesica without cornuti or sclerotizations. Female genitalia: Papillae anales wide, setose. Apophyses posteriores short, apophyses anteriores very short. Lamella postvaginalis sclerotized, irregular in shape. Corpus bursae membranous, thin. Biology, phenology, habitat: Larvae of Crocota species apparently polyphagous on low-growing herbaceous plants in the subalpine zone. Recorded on Polygonum bistorta, Inula helenium, Gentiana acaulis, Leontodon, Achillea, Coronilla minima, Rumex, Plantago and Taraxacum (Flamigni et al. 2007 and references therein; Redondo et al. 2009). Larva of C. tinctaria overwintering, pupating in the ground (Vorbrodt 1930–1931; Rezbanyai-Reser 2000; 2002a; 2002b; 2002c). All species occurring in mountain habitats, mostly in the grasslands and slopes in the subalpine zone. Adults diurnal, but both sexes also attracted to light. Immature stages: Larva of C. niveata brown, with several differently coloured brown longitudinal stripes, broken into slightly triangular wedges dorsally (Lepiforum 2018). Cremaster with three or fours pairs of curved setae, apical pair longest (Patočka & Turčani 2005). 213

Remarks: Crocota is a peculiar genus because the wing colour implies relationships that are not supported by structural features in the male and female genitalia. The bright orange-­ yellow C. tinctaria is structurally very similar to C. peletieraria with dark brown males and creamy white females. Another bright orange-yellow species C. pseudotinctaria, which cannot be separated externally from C. tinctaria, is structurally more similar to C. niveata, which has almost pure white wings in both sexes. So the orange-yellow species (C. tinctaria and C. pseudotinctaria) are not closely related, nor are the species with white females (C. peletieraria and C. niveata). Cleogene ostrogovichi Caradja, 1930, described from Romania, was classified in the genus Crocota (e.g. Scoble 1999) until a new, monotypic genus, Palaeocrocota was described by Stanescu (2002). The specimen has been shown to be a melanic specimen of Isturgia roraria (Fabricius, 1776) (Rákosy et al. 2017). Crocota ricaria Geyer, 1837 from South Africa is potentially a mislabelled specimen, whose identity remains unknown. All known Crocota species are restricted to the South European mountains and occurrence of a single species in South Africa would render the genus distribution really unusual. The species was included in genus Crocota by Scoble (1999), but not in the South African Checklist (Vári et al. 2002) and its appearance is not familiar to Hermann Staude (pers. comm.), an acknowledged South African lepidopterist. According to the original illustration, C. ricaria is brown-grey, with distinct silvery, elongated discal spots on forewings and wedge-shaped, discontinuous terminal line (Geyer in Hübner 1837, plate 153, figs 891, 892). Geyer compared it to C. peletieraria (Duponchel, 1830). The type specimen of C. ricaria is lost. Recently Crocota has been classified in the Boarmiini (Vives Moreno 1994; Leraut 1997; Hausmann et al. 2004; 2011a; Vives Moreno 2014) or in the wide sense of Boarmiini of Holloway (1994). The morphological support for this association is weak (our observation), as illustrated here against some of the Boarmiini synapomorphies (from Holloway 1994): usually pupal cremaster apically with bifid projection, lacking the lateral setae (Crocota: cremaster apex with two separate D2 setae, with three pairs of thin, hooked lateral setae), fovea present in male forewing base (Crocota: fovea absent) and usually a transverse comb of setae are present on third abdominal sternite (Crocota: setae absent). Crocota is structurally similar to numerous Gnophini genera and it shares a number of Gnophini characters as defined by Viidalepp et al. (2007) and as illustrated in this book. Crocota has not been subject to analytical phylogenetic studies yet. Systematic position: transferred from Boarmiini Duponchel, 1845 to Gnophini Duponchel, 1845 (new tribus combination). Crocota shows some structural similarities to Sciadia and Elophos but we retain it as a separate genus because a phylogenetic study on the relationships of these genera is lacking and Crocota has several unique features: Anterior arms of juxta absent, aedeagus attached to juxta via a (semi)sclerotized ridge, aedeagus apex with a single tooth or numerous ‘micro­ teeth’, uncus with two horns (absent in C. pseudotinctaria), corpus bursae without sclerotizations. More research on the relationships of these genera is needed.

64. Crocota tinctaria (Hübner, 1799) Geometra tinctaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 23, fig. 121 (­Europe). Syntype(s) lost. Neotype ♂ (designated by Leraut 1999, France: Hautes-Alpes: Abriès, gen. prep. number 3995, MNHN).

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‡ Phalaena lutearia Fabricius, 1794: Ent. Syst. 3 (2): 143 (Italy). Syntype(s) male(s) lost. Junior primary homonym of Phalaena Geometra lutearia Villers, 1789 (identity: Lythria purpuraria Linnaeus, 1758, Sterrhinae). Crocota tinctaria estachyi Leraut, 1999: Alexanor 20: 470, figs. 2, 8 (France: Alpes-Maritimes, Madone de Fenestre). Holotype ♂ (gen. prep. number 3994 in MNHN) (examined). Tentatively validated at subspecific rank. Crocota tinctaria vintejouxi Leraut, 1999: Alexanor 20: 471, figs. 3, 9 (France: Cantal, Plomb du Cantal). Holotype ♂ (gen. prep. number 3952 in MNHN) (examined). Tentatively validated at subspecific rank. Unavailable names (infrasubspecific): albescens: Leraut (2009) (f.); martinae: Leraut (1993) (f.); quadripunctaria: Thierry-Mieg (1907) (ab.); subatrata: Wehrli (1953) (f.).

External characters and abdomen: Sexually dimorphic. Male wingspan 29–35 mm, female smaller, wingspan 22–26 mm. Male wings uniform bright orange-yellow, veins slightly darker than wings, wings weakly irrorated with brown-grey. Discal spots barely visible, dark orange. Wings below as above, slightly more irrorated with brown-grey. Female wings paler, pale orange-yellow, finely irrorated with brown-grey, hindwings often paler. Female forewing apex sharper. Frons, thorax and abomen concolorous with wings. Base of collar paler yellow, sometimes whitish. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Hindtibia with 2+2 spurs in both sexes. Male tympanal organs large, elongated, female organs very small. Abdomen as under genus description, see also Text-fig. 127. Variation: Amount of irrorations vary, underside often more irrorated. Males can have a tint of olive-green on their wings and females of subsp. estachyi can occasionally be whitish. Subsp. estachyi externally similar to nominotypical subspecies, except the occasional occurrence of whitish females (Leraut 2009). According to Leraut (1999) ventral margin of valva slightly more concave than in nominotypical subspecies and valva costa with two ­basal and one medial spine (spines absent in nominotypical subspecies). According to L ­ eraut (1999; 2009) restricted to Alpes de Haute-Provence and Alpes Maritimes in France and Apennines in Italy. All specimens from the Apennines in Italy with two or more spines on the costa (Flamigni et al. 2007, fig. 38; C. Flamigni pers. comm.). However, nominotypical subspecies and ‘subspecies estachyi’ (as identified by genitalia features) occurring sympatrically in Maritime Alps and Ligurian Alps in Italy, thus questioning the validity of estachyi at subspecies level (Flamigni et al. 2007; see also Flamigni et al. 2016 for additional data). Potential hybridization zone and subspecies hypothesis needs further research. Potentially a junior synonym. Subsp. vintejouxi externally similar to nominotypical subspecies but according to Leraut (1999) with smaller wingspan on average. Male genitalia similar to those of subsp. estachyi, valva costa bearing two basal and one medial spine. According to Leraut (1999; 2009) restricted to Cantal, Loire and Puy-de-Dôme departements of France and allopatric, not in contact with other populations of the species. Potentially a junior synonym. Male genitalia: Uncus wide, posterior margin straight or weakly concave, with two ­distinct horns laterally. Presence of socii unclear, several lateral and ventral stout setae potentially representing rudimentary structure. Gnathos absent. Tegumen large, with distinct mid-­ dorsal sutura. Valva wide, sclerotized, weakly setose, ventral margin weakly concave. Valva costa heavily sclerotized. Submarginal ridge long, membranous, folded, apical portion densely setose. Juxta curved ventrally, narrow, heavily sclerotized, fused with aedeagus near caecum. Dorsal lobes of juxta short, horn-shaped. Saccus wide, slightly rectangular. 215

Aedeagus curved ventrally (about 180 degrees), flattened dorsoventrally. Caecum distinct, round, narrow. Aedeagus apex bilobed, left side (when viewed ventrally) with sharp projection reaching aedeagus apex, right side blunt-ended, turned left, with few minute subapical sclerotized teeth. Aedeagus dorsoventrally flattened and very narrow, therefore vesica not everted, presumably a narrow tube, without sclerotizations. Female genitalia: Papillae anales wide, setose. Apophyses posteriores short, apophyses anteriores very short. Lamella postvaginalis sclerotized, irregular in shape. Margin of lamella antevaginalis rather straight. Ductus bursae evenly wide, membranous. Base of ductus seminalis widened, with small irregular sclerotization. Corpus bursae roundish, membranous, thin. Distribution and abundance: European. In the western part of the Alps, in Massif Central in France and with two more isolated occurrences in the Appenines in Italy. Recorded from seven localities in south-western Ukraine, all records being old (Nowicki 1860; Hormuzaki 1899). Five of the mentioned localities are in the Carpathian Mountains, therefore potentially being suitable habitats for C. tinctaria, whereas two localities are in lowland plain area. The latter two are ex larva specimens, reared in 1910 and 1916, according to their labels. Lviv museum in Ukraine holds seven specimens from five localities, and recently Igor ­Kostjuk (pers. comm.) has studied their identity from the external and genitalia features. He confirms they are C. tinctaria. However, since there are no records of this species from adjoining Slovakia and Poland with higher mountains than in Ukraine, and since there are no recent records, we find it unlikely that C. tinctaria has, or has had, populations in Ukraine. It is also peculiar that some of the Ukrainean records are labelled as reared from caterpillar, because C. tinctaria is much easier to observe by day, when adults are easily flushed up from the vegetation. – Local, rather often found in numbers.

216

Phenology: Univoltine. From mid-June to early or mid-August. Hibernation as a small cater­pillar (M. Leipnitz pers. comm.). Adults active at night, readily attracted to light, males in considerably larger numbers than females. Adults at daytime often found feeding on flowers, being flushed up very easily. Biology: Larva polyphagous. Recorded on Polygonum bistorta, Inula helenium and Gentiana acaulis (Flamigni et al. 2007). Reared on Taraxacum officinale, Rumex, Plantago (Rezbanyai-­ Reser 2002b), Achillea and Thymus pulegioides (M. Leipnitz pers. comm.). Pupation in the ground (Flamigni et al. 2007). Habitat: Alpine. In open meadows above treeline up to 2600 m, rarely lower. Similar species: Externally C. tinctaria without diagnostic differences from C. pseudotinctaria but genitalia allowing easy identification. Genitalia of C. tinctaria and C. peletieraria similar but allowing reliable identification, see the Genitalia plates and text under C. peletieraria. Genetic data: Four BINs: BOLD:AAE4323 (n=4 from France and Italy: Piedmont and Abruzzo), BOLD:ACR3183 (n=2 from Italy: South Tyrol)¸ BOLD:ABY9600 (n=3 from Switzerland: Valais), BOLD:ABY8927 and BOLD:ABY9599 (n=4 from Italy: Val d’Aosta). Genetically very heterogeneous, BINs diverging by 1.9–3.4% from each other. Nearest species: C. peletieraria (2.2%). Remarks: See text under C. peletieraria.

65. Crocota peletieraria (Duponchel, 1830) Cleogene peletieraria Duponchel, 1830: in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 8 (1): 140, pl. 180, fig. 2 (France: near Bareges, near Bagneres sur l’Adour). Syntype(s) not located (see also Leraut 1999), identity not in doubt on the basis of original description and illustration. Unavailable names (infrasubspecific): darwiniana (f.): Rondou (1914); flavescens (f.): Leraut (2009).

External characters and abdomen: Distinctly sexually dimorphic. Male wingspan 31–33 mm. Wings uniform dark brown (veins slightly darker than wings), irrorated with grey-silver, scales partially iridescent. Discal spots barely visible, dark brown. Forewing margin round. Wings below as above. Female wingspan 26–27 mm. Wings uniform creamy white (veins slightly darker than wings), weakly irrorated with brown. Discal spots barely visible. Forewing margin slightly concave below apex. Wings below as above. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Hindtibia with 2+2 spurs in both sexes. Male tympanal organs medium-sized, female organs very small. Abdomen as under genus description, see also Text-fig. 127. Variation: Females irrorated with brown, grey or yellow scales. Male genitalia: Generally as in C. tinctaria (see above), but with following differences: Ventral margin of valva less concave in C. peletieraria (ventral margin more concave in C. tinctaria), valva costa with setose basal lobe in C. peletieraria (valva costa without lobe in C. tinctaria), aedeagus base weakly bent and aedeagus apex with short tooth in C. peletieraria (aedeagus base distinctly bent and aedeagus apex with long tooth in C. tinctaria). Female genitalia: Generally as in C. tinctaria (see above), but with following difference: Lamella postvaginalis is wide and membranous and ductus bursae is funnel-shaped in C. peletieraria (lamella postvaginalis is narrow and sclerotized and ductus bursae is evenly wide in C. tinctaria). 217

Distribution and abundance: Iberian. Only known from the Pyrenees and the Cantabrian mountains in north-western Spain. – Local, usually found in limited numbers. Phenology: Univoltine. In July and August. Hibernation as second or third instar larvae (D. Fritsch pers. comm.; rearing observation). Adults active at night, both sexes attracted to light. According to Redondo et al. (2009) adults diurnal, but females attracted to light. Biology: Natural foodplant(s) apparently unknown. Recorded on Taraxacum officinale (­Redondo et al. 2009). Reared on Plantago (D. Fritsch pers. comm.). Habitat: Montane-alpine. In open places in forests, in meadows amd in places with rocks and stones. From 1500 up to 2500 m. Similar species: Uniform dark brown males of C. peletieraria are unique, there are no similar species in Europe. Females of C. peletieraria resemble C. niveata but wings are creamy white in C. peletieraria (wings satin white in C. niveata). Genitalia of C. peletieraria and C. tinctaria are similar, but minor differences allow identification: valva costa with setose basal lobe in C. peletieraria (valva costa without lobe in C. tinctaria), aedeagus base weakly bent and aedeagus apex with short tooth in C. peletieraria (aedeagus base distinctly bent and aedeagus apex with long tooth in C. tinctaria) and ductus bursae is funnel-shaped in C. peletieraria (evenly wide in C. tinctaria). Genetic data: BIN: BOLD:ABY9602 (n=2 from northern Spain and southern France). Genetically homogeneous. Nearest species: C. tinctaria (2.2%). Remarks: Male and female genitalia of C. peletieraria and C. tinctaria are structurally similar, only minor differences are found. Preliminary analysis of COI barcode genes supports close relationship also.

218

66. Crocota pseudotinctaria Leraut, 1999 Crocota pseudotinctaria Leraut, 1999: Alexanor 20: 473, figs. 5, 11 (Switzerland: Valais). Holotype ♂ (gen. prep. number 3996 in MNHN) (examined). According to Leraut (1999, p. 474) [our translation from French], “the same specimen was illustrated in colour earlier (Leraut 1992, p. 211) under an erroneous name Crocota peletieraria due to transposing legends.” Unavailable names (infrasubspecific): griseostrigata: Vorbrodt & Müller-Rutz (1917) (f.).

External characters and abdomen: Externally, C. pseudotinctaria is without diagnostic differences from C. tinctaria, see text above, but the genitalia are different and allow easy identification. Male wingspan 26–33 mm, females smaller, wingspan 24–26 mm. Male wings uniform bright orange-yellow, veins slightly darker than wings, wings weakly irrorated with brown-grey. Discal spots barely visible, dark orange. Wings below as above, slightly more irrorated with brown-grey. Female wings paler, pale orange-yellow, finely irrorated with brown-grey, hindwings often paler. Female forewing apex sharper. Variation: Amount of irroration on wings varies. Male genitalia: Uncus narrow, tapering to apex, apex variable, with minute emargination or rather distinct concavity. Presence of socii unclear, several lateral and ventral stout setae potentially representing rudimentary structure. Gnathos absent. Tegumen large, with distinct mid-dorsal sutura. Valva wide, narrower towards apex. Valva costa sclerotized, medial part with lobe and several stout setae. Submarginal ridge short, membranous, folded, apical portion sparsely setose. Ventral margin of valva weakly concave. Juxta curved ventrally, narrow, heavily sclerotized, fused with aedeagus near caecum. Dorsal lobes of juxta short, horn-shaped. Saccus rather wide. Aedeagus undulating, apex slightly curved ventrally, subapical part not expanded, heavily covered with sclerotized teeth and stout setae. Caecum short, straight. Vesica weakly expanded at base, straight tube, without sclerotizations. Female genitalia: Papillae anales small, setose. Apophyses posteriores short, apophyses anteriores very short. Lamella postvaginalis sclerotized, large, posterior margin folded midventrally. Lamella antevaginalis sclerotized, irregularly shaped narrow ridge. Ductus bursae short, membranous. Corpus bursae elongated, membranous, thin. Distribution and abundance: European. Restricted to western Alps, with one old specimen from North Tyrol in Austria (Huemer et al. 2009). – Local, rather often found in numbers. Phenology: Univoltine. From early July to late August. Adults active at night, readily attracted to light, males in considerably larger numbers than females. In daytime often found feeding on flowers, very easily flushed up. Biology: Larva supposedly polyphagous. Natural foodplant(s) unknown. Reared on Taraxacum officinale (Rezbanyai-Reser 2002b). Habitat: Montane-alpine. In open meadows, both below and above treeline. From 1300 up to 2600 m. Similar species: Externally C. pseudotinctaria is without diagnostic differences from C. tinctaria, but the genitalia are different and allow easy identification. Structurally C. pseudotinctaria and C. niveata are the most similar. C. pseudotinctaria with narrow uncus and setose/ microteeth-covered aedeagus apex (in C. niveata uncus wider, aedeagus apex expanded and sparsely covered with microteeth). Lamella postvaginalis large, with round margin in C. pseudotinctaria (in C. niveata lamella postvaginalis small, with almost straight margin). 219

Genetic data: BIN: BOLD:AAE9435 (n=12 from Switzerland: Valais). Genetically very homo­ geneous. Nearest species: C. peletieraria (4.0%), C. tinctaria (4.3%), C. niveata (5.6%).

67. Crocota niveata (Scopoli, 1763) Phalaena niveata Scopoli, 1763: Ent. Carniolica, p. 217, fig. 539 (north-western Slovenia: Carniolia). Syntype(s) lost. Scoble (1999) states that syntype(s) are in the Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark. This is an error, confirmed by O. Karsholt (pers. comm.) Geometra illibata Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 116 (Austria: Vienna distrcit). Syntype(s) lost. ‡ Phalaena niveata Fabricius, 1775: Syst. Ent, p. 638 (’America’) [incorrect locality]. Syntypes(s) (Zoological Museum, Natural History Museum of Denmark, Copenhagen, Denmark, four wings glued to cardboard, examined). Homonym of Phalaena niveata Scopoli, 1763. Geometra illibaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2): pl. 40, fig. 207 (­Europe). Syntype(s) male(s) lost. Unnecessary replacement name for illibata Denis & Schifferm ­ üller.

External characters and abdomen: Weakly sexual dimorphic. Male wingspan 26–30 mm, female wingspan 21–26 mm, females smaller and forewing apex sharper. Wings of both sexes uniform white, without creamy or yellowish scales, veins slightly darker than wings, and wings weakly irrorated with pale brown. Wings below as above, except male forewing almost pale brown, distinctly darker than hindwings. Frons, thorax and abomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Hindtibia with 2+2 spurs in both sexes. Male tympanal organs large, elongated, female organs very small. Abdomen as under genus description, see also Text-fig. 127. Variation: Amount of irrorations varying.

220

Male genitalia: Generally as C. pseudotinctaria, but with following differences: uncus wider, margin slightly rounded in C. niveata (uncus evenly narrowing to tip in C. pseudotinctaria). Aedeagus apex expanded, sparsely covered with microteeth, and with sclerotized tooth subapically in C. niveata (in C. pseudotinctaria aedeagus apex not expanded, densely covered with microteeth and stout setae, and without sclerotized tooth subapically). Female genitalia: Generally as C. pseudotinctaria, but with following differences: Lamella postvaginalis small, with almost straight margin in C. niveata (lamella postvaginalis large, with round margin in C. pseudotinctaria). Distribution and abundance: European. Endemic in southern central Austria with a limited distribution area. Fajčik (2003) erroneously mentions the species from Slovenia (S. Gomboc pers. comm.). – Local, sometimes abundant in its localities.

Phenology: Univoltine. From early July to early August. Hibernation as larva. Adults active at night, attracted to light, males more frequently than females. Most specimens, however, found by day when actively flying or flushed up. Biology: Possibly polyphagous. Natural foodplants unknown. Recorded on “low plants” (Fajčik 2003) which probably means herbaceous plants. Reared on Taraxacum (Mack 1985). Habitat: Montane-alpine. In open meadows just below and above treeline, often in somewhat moist places. From 1500 m up to at least 2200 m. Similar species: C. niveata males are whitish, the only species in genus Crocota, allowing easy identification. Unrelated Gypsochroa renitidata (Hübner, 1817) (Desmobathrinae) has pure white wings and rather straight forewing margin (wings whitish and forewing margin round in C. niveata) (see Hausmann 2001). C. niveata females whitish (C. peletieraria 221

females creamy white and unrelated G. renitidata females considerably larger and pure white). Also some species in the unrelated genus Lithostege Hübner, 1825 (Larentiinae: Chesiadini) are whitish, showing certain external resemblance (see Hausmann & Viidalepp 2012). Genitalia of both sexes of C. niveata allow reliable identification. Genetic data: BIN: BOLD:AAI9737 (n=7 from Austria). Genetically very homogeneous. Nearest species: C. peletieraria (4.9%), C. pseudotinctaria (5.5%), C. tinctaria (6.5%).

Synopsia Hübner, 1825 Synopsia Hübner, [1825] 1816, Verz. bekannter Schmett.: 317. Type species: Geometra sociaria Hübner, 1799. There are no genus synonyms.

Diversity and distribution: Monobasic, containing one species in the Palaearctic region. External characters and abdomen: Medium-sized moths, wingspan 28–44 mm. Wings brown with light grey areas. Venation of forewing: R2 anastomising with R3+R4. Antennae bipectinate or partially bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Uncus and gnathos well developed, gnathos ventrally fused. Valva wide, sclerotized, with projection on costa. Valva apex round, setose. Juxta developed as a plate. Aedeagus, see species description. Female genitalia: Papillae anales setose. Apophyses posteriores long, straight, apophyses anteriores shorter. Anterior part of corpus bursae membranous, with signum. Remarks: Gnophini position is supported by multi-gene analysis (Murillo-Ramos et al. 2019), Synopsia being sister to lineage that includes Phthonandria + Angerona and Nearctic genera. Synopsia is listed under Gnophini in Beljaev (2016).

68. Synopsia sociaria (Hübner, 1799) Geometra sociaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 29, fig. 155 (­Europe). Syntype(s). Boarmia propinquaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 197 (France: Corsica, Monspelii). Syntype(s). Validated at subspecific rank by Scoble (1999), downgraded here from subspecific rank (syn. n.), based on absence of constant and significant differential features, falling within intraspecific variation. Boarmia luridaria Freyer, 1845: Neuere Beitr. Schmettkde. 5 (79): 154, pl. 474, figs 3, 4 (Russia: Urals). Syntype(s). Validated at subspecific rank by Leraut (2009), downgraded here from subspecific rank (syn. n.), based on absence of constant and significant differential features, falling within intraspecific variation. Synopsia staudingeraria Martorell, 1874: Cat. Meth. Raz. Lepid.: 140 (as standingeraria, an incorrect original spelling, errata, p. 232) (Spain: Barcelona). Synopsia sociaria var. almasa Schawerda, 1912: Verh. zool.-bot. Ges. Wien 62 (5/6): (143) (Dalmatia: Mostar; Velez; Bisina). Syntypes 8♂1♀. Validated at subspecific rank by Scoble (1999), downgraded here from subspecific rank (syn. n.), based on absence of constant and significant differential features, falling within intraspecific variation. Synopsia sociaria unitaria Prout, 1915: in Seitz, The Macrolepidoptera of the World 4: 364 (Russia: Sarepta (Krasnoarmeysk); Altai). Syntype(s) including ♀. Validated at subspecific rank by Scoble (1999), downgraded here from subspecific rank (syn. n.), based on absence of constant and significant differential features, falling within intraspecific variation.

222

Synopsia sociaria f. acuta Wehrli, 1926: Dt. ent. Z. Iris 40 (2/3): 127 (Spain: Sierra Nevada, 1500 m). Syntypes 6♂.

External characters and abdomen: Wingspan ♂ 26–35 mm, females ♀ 31–43 mm. Ground colour of forewings beige to brown, dark speckled. Medial area of forewing and basal area of hindwing often paler. Terminal area often with a black subapical streak and a double blotch. Terminal line black, discontinuous. Fringes divided by a fine line, chequered beige and grey. Forewing with postmedial line black, sinuous, distally edged brownish, double angled at costa. Antemedial line bent, jagged, proximally edged brownish. Medial line diffuse, indistinct, sometimes absent. Postmedial line of hindwing bent, straight over one-third towards costa, concave towards inner termen. Discal spots weak or absent. Underside paler, with distinct bright apical blotch, blackish postmedial line and discal spots distinct, outstanding. Frons, vertex, collar, tegula and thorax beige, chequered with grey hairy scales. Abdomen dorsally with blackish rings. Antennae bipectinate in male, about 10 apical flagellomeres free of branches, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Medial area of forewing and basal area of hindwing varying from light beige to brown, in the latter case more homogeneous in colour, but always dark speckled. Male genitalia: Uncus sclerotized, triangular base bearing two digitiform, bent tips. Gnathos wide, fused ventrally, forming an upturned, sclerotized, tapered structure. Valva wide, apex round, setose; at the centre of costa a projection with one or two spines at tip. Sacculus wide, obtusely angled, inwards a ridge. Saccus wide, tapered. Juxta a wide plate, distal margin concave. Aedeagus thick, cylindrical, vesica with two groups of cornuti (total about 25 spines). Female genitalia: Papillae anales slightly elongated, setose. Apophyses posteriores long, straight, length on average 2.25 mm, apophyses anteriores shorter, on average 1.4 mm, spatulate at tip. Lamella antevaginalis consisting of two strongly sclerotized, bent plates, lamella postvaginalis a plate with concave posterior margin. Ductus bursae wide, sclerotized. Anterior part of corpus bursae membranous, oval, concentrically striated, signum unevenly star-shaped. Diameter of posterior part of corpus bursae half that of anterior part, sclerotized. Distribution and abundance: European-West Asiatic. Widely distributed in Europe from the southern Ural Mountains through eastern and southern central Europe to France, in the south from Portugal and Spain, across Italy including Sardinia and Sicily to the Balkan peninsula including Greece. Formerly occurring up to north-western Germany from where it has disappeared in recent decades (see open circles on map). – In Europe common, usually occurring in limited numbers, but sometimes common in Mediterranean habitats. – Outside Europe in Turkey, Georgia, Armenia and Azerbaijan, Kazakhstan, Kirghizstan and Uzbekistan (subsp. unitaria). Phenology: Bivoltine. Generally from late May to early September. In Spain from April to June and August to September (Redondo et al. 2009). However, we have also records from Sierra Nevada from early and mid-July. In Crimea from late May to June and from late July to mid-September (Budashkin & Kostjuk 1987). Caterpillars found from April to August (Crimea: V. Savchuk pers. comm.), from early March to early May and from June to September (Spain: G. King pers. comm.), and in mid-March (Hungary: B. Tóth pers. comm.). Overwintering as half grown larva (M. Leipnitz pers. comm.). Last instar larvae found in southern France from early May to early June (J. Gelbrecht pers. comm.). Adults nocturnal, readily attracted to light. 223

Biology: Larva polyphagous. Found on Salvia (D. Fritsch pers. comm.), Clematis, Conso­ lida (T. Drechsel pers. comm.), Artemisia camphorata, flowers of Dianthus, Genista (M. Leipnitz pers. comm.), Salvia nutans, Limonium scoparium, Onobrychis pallasii (V. Savchuk pers. comm.), Conyza canadensis (G. King pers. comm.) and Artemisia campestris (Z. Las˘tu˚ vka pers. comm.). Recorded on Quercus, Thalictrum, Rosa, Sanguisorba minor, Prunus, Cytisus, Spartium, Dorycnium, Lotus, Hippophae, Tamarix, Verbascum nigrum, Scrophularia, Calluna, Ligustrum, Echium, Thymus, Lavandula, Plantago, Aster, Helichrysum italicum, Achillea and Centaurea (Flamigni et al. 2007). Reared on Artemisia campestris (G. King pers. comm.), Taraxacum officinale, Plantago major (B. Tóth pers. comm.) and Heli­chrysum italicum (T. Varenne pers. comm.). Pupation among plant litter in a cocoon (V. Savchuk, B. Tóth, G. King pers. comm.) or in the ground (M. Leipnitz, Z. Las˘tu˚ vka pers. comm.). Habitat: Xerothermophilous. In a large variety of habitat types such as Mediterranean macchia, open and semi-open places with scattered trees and bushes. In Portugal only in limestone areas (M. Corley pers. comm.). From sea-level up to 1700 m in southern Italy (Flamigni et al. 2007) and to 1800 m in Sierra Nevada in southern Spain (Redondo et al. 2009). Similar species: Some species of the genus Peribatodes Wehrli, 1943, all differing by the shape of the forewing postmedial line not being acute-angled below costa. Lightly contrasted forms of Menophra japygiaria without acute-angle below costa in postmedial line, usually with pale blotch in hindwing terminal area. Genetic data: BIN: BOLD:AAD3244 (n=15 from Austria, Hungary, Italy, Bulgaria, Turkey). Genetically slightly heterogeneous, maximum variation 1.4%. Nearest species: Synopsidia phasidaria (Rogenhofer, 1873) from Transcaucasus and Iran (4.1%); Phthonandria stricta­ ria (7.6%).

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Phthonandria Warren, 1894 Phthonandria Warren, 1894, Novit. Zool. 1:434. Type species: Hemerophila atrilineata Butler, 1881.

Diversity and distribution: Eight species in the eastern Palaearctic region, in Europe one species only, occurring in the southern Ural mountains. Genus combination of several taxa in Phthonandria is putative, including one species in Africa, requiring taxonomic re-examination. One species, Synopsia strictaria (Lederer, 1939) is transferred from Megalycinia to Phthonandria. External characters and abdomen (based on Palaearctic species): Medium to large moths, wingspan about 25−54 mm. Wings various shades of brown, medial lines oblique, blackish. Postmedial line distinct on both wings, sharply curved inwards near forewing costa. Hindwing termen undulating. Male antennae bipectinate with long pectinations, female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Few species with setae on male 3rd sternite and hindtibia with scent-brush. Male genitalia (based on Palaearctic species): Uncus small, bifid. Socii absent. Gnathos large, fused, medial element upturned. Transtilla arms separate, very large. Valva wide, weakly constricted at base, costa with spinose lobe, apex densely setose. Valva with medial ridge. Saccus large. Aedeagus with or without cornuti. Female genitalia (based on Palaearctic species): Papillae anales elongated, setose. Apophyses posteriores about twice length of apophyses anteriores. Sclerotizations around ostium bursae weak. Ductus bursae short. Posterior part of corpus bursae narrow, anterior part rounded. Signum oval, elongated laterally, concave at centre, margins sclerotized. Biology, habitat, phenology: Very little data available. Larva of P. atrilineata recorded on Morus alba (Kim et al. 2001). P. strictaria inhabiting various kind of steppes, adults nocturnal, both sexes readily attracted to light (K. Nupponen pers. comm.), adults flying early in summer, but data available for few species only. Immature stages: No data available. Remarks: Phthonandria is part of the Gnophini complex of genera, whose relationships require further study. These genera are characterized for instance by an elongated and medially concave signum, large and separated transtilla arms, and costa with setose lobe. Synopsia and Phthonandria are particularly similar in many features, and other genera include at least Chariaspilates Wehrli, 1953, Angerona Duponchel, 1829, Synopsidia Djakonov, 1935 and Hypoxystis Prout, 1915. Externally, several species of Nychiodes Lederer, 1853 and Menophra Moore, 1887 are similar to Phthonandria, but they do not share the mentioned structural characters. Recently Phthonandria has been classified in the Gnophini (Viidalepp 1996). Phthonandria is structurally similar to numerous Gnophini genera (see above) and it shares a number of Gnophini characters as defined by Viidalepp et al. (2007) and as illustrated in this book. Gnophini position is supported by multi-gene analyses (Jiang et al. 2017; Murillo-Ramos et al. 2019), in the latter Phthonandria is sister to Angerona (Murillo-Ramos et al. 2019). Systematic position: Gnophini.

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69. Phthonandria strictaria (Lederer, 1853), comb. n. Synopsia strictaria Lederer, 1853: Verh. zool.-bot. Ver. Wien 3: 378, pl. 6, fig. 2 (Russia: Siberia). Syntypes 2♀ (MNHU, examined externally). Lederer (1853) mentions in the original description that only two females are known. In coll. MNHU there are 3 males and 3 females with purple labels “Origin.”, and all of them are easily identifiable as P. strictaria. Hemerophila strictaria variegata Djakonov, 1936: Ark. Zool. 27A (39): 47 (China: Kansu (south), Tao-ho, Lu-pa-sze, ca. 2750 m, Ka-tien-kou, Kung-ta, ca. 2850 m. Syntypes 2♂1♀ (Natur­ historiska Riksmuseet, Stockholm, Sweden, examined externally). Treated valid at sub­specific rank recently (Scoble 1999), downgraded here (syn. n.), based on absence of constant differential features, falling within infraspecific variation. See Remarks.

External characters and abdomen: Wingspan 25−34 mm. Wings creamy white, forewing traversed by whitish oblique medial area, bordered by dark brown medial and postmedial lines. Terminal area brown, interrupted by creamy white wavy line. Hindwing base creamy white, irrorated, creamy white medial area most distinct near inner margin, bordered by brown and whitish lines in terminal area. Terminal line blackish, distinct. Fringes weakly chequered or concolorous with wings. Forewing margin round, hindwing termen undulating. Discal spots small, distinct, blackish. Underside almost uniform grey-brown, irrorated with brown, terminal area darker. Discal spots distinct, blackish. Frons, collar, thorax and abdomen concolorous with wings. Male antennae bipectinate with long pectinations, female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Tympanal organs medium-sized, base evenly curved (not enlarged, not bottle-shaped). Abdominal segments without modifications. Variation: Variable. Wingspan varying, apparently in a clinal manner (n=about 100 specimens), ranging from 25 mm (Russian Far East) to 34 mm (southern Ural region in Russia). Oblique medial area in forewing narrow, weakly visible, or wide and dominant, distinctly paler than rest of wing or nearly concolorous. Medial lines suffused, barely recognizable, or sharp, creating striped appearance. Ground colour varying from creamy white to greybrown. Wings heavily irrorated with dark brown, or almost devoid of irrorations. Males often more suffused than females. One examined, aberrant male with forewing almost uniform dark brown and blackish, hindwings dark grey (Russia: Tuva republic, Tannu-Ola in coll. ZMH). Specimens in southern Ural (Text-figs 131−133) larger, wings pale brown, pale oblique medial area wider, medial and postmedial lines strong, wavy line absent or weak. Pale, oblique forewing medial area wide in most specimens, but in few specimens area narrow and almost absent. In two examined male specimens medial and postmedial lines are fused at centre of wing, and widely set apart at costa and inner margin (coll. Junnilainen). Male genitalia with narrow valva base, costal margin upright, saccus wide (Text-figs 129, 132). Female genitalia with elongated corpus bursae (Text-figs 130, 133). See Remarks. Male genitalia: Uncus very small, weakly bilobed. Socii absent. Gnathos large, fused, medial element smooth, upturned. Transtilla arms separate, large, sclerotized. Valva wide, weakly constricted at base, costa with spinose lobe basally, apex setose. Valva with sclerotized medial ridge, apex with stout setae, sacculus weakly sclerotized. Juxta sclerotized plate. Saccus elongated, rather narrow, margin round. Aedeagus straight, apex weakly elongated. Vesica opens at about 135 degree angle, base weakly enlarged, without cornutus. Female genitalia: Papillae anales elongated, setose. Apophyses posteriores about twice length of apophyses anteriores. Lamella postvaginalis membranous. Lamella antevaginalis weakly v-shaped, sclerotized. Ductus bursae short, membranous. Posterior part of corpus

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bursae narrow, anterior part rounded. Signum oval, elongated laterally, concave in middle, margins sclerotized. Distribution and abundance: Euro-Siberian. In Europe only around the southern Ural mountain chain. – In Europe rather poorly known, local, but can be rather abundant. (K. Nupponen pers. comm.). – Outside Europe in the eastern vicinity of the southern Urals and in Kazakhstan, southern Siberia, Buryatia, Dahuria, Far East Russia and Mongolia.

Phenology: Univoltine. Recorded from late May to early July, peaking in mid-June in southern Urals (K. Nupponen pers. comm.). In Altai from early June to mid-July. Adults active at night, both females and males readily attracted to light (K. Nupponen pers. comm.). Biology: Unknown. Habitat: Xerothermophilous. In various steppe habitat types, forest steppe and treeless lowland steppe, as well as calcareous steppe slopes. In southern Urals a lowland species found up to 450 m (K. Nuponen pers. comm.), in the Altai mountains up to 2500 m. Similar species: No similar species in Europe. The non-European Phthonandria emaria (Wehrli, 1941) and some Menophra species share similar oblique and pale area in forewing but those are easily diagnosable. Genetic data: BIN: BOLD:AAH0731 (n=9 from Russia, Kazakhstan, Mongolia, northern China). Genetically homogeneous, maximum intra-specific variation 0.39%. Nearest species: Phthonandria emaria (Wehrli, 1941) from China (6.4%), Synopsia sociaria (7.6%). Remarks: Comparative morphological analysis revealed that Megalycinia is not correct genus combination (as for instance in Ahola et al. 1999; Scoble 1999; Hausmann et al. 2004; 2011a), neither is Synopsia Hübner, 1825 (as for instance in Vojnits 1975; Viidalepp 1996; Mironov et al. 2008) for the taxon strictaria Lederer, 1853. Morphologically P. strictaria is similar to 227

Text-figs 128−133. Diagnostic characters (indicated) of Phthonandria strictaria (Lederer, 1839) in East Asia and in southern Urals. Text-fig. 128. Adult. Russia: Amur Region, Bureinskyi Mountains, Tyrma, 800 m, 1.vii.1997 (coll. Skou). Text-fig. 129. Male Genitalia. Russia: Amur Region, Bureinskyi Mountains, Tyrma, 800 m, 1.vii.1997, slide 2225 Sihvonen (coll. Skou). Text-fig. 130. Corpus bursae (photographed in water during dissection to maintain the shape), Mongolia: West Hangay Mts., 10 km E from Uliastay, 1830 m, 23.vi.2004, slide 2228 Sihvonen (coll. Skou). Textfig. 131. Adult. Russia: S-Ural, Orenburg district, Donskoje village, 11.vi.1998 (coll. N ­ upponen). Text-fig. 132. Male genitalia. Russia: Ural Mountains, Orenburg Region, Kyvandyk Village, 1−11. vi.1995, slide 2229 Sihvonen (coll. Skou). Text-fig. 133. Corpus bursae (photographed in water during dissection to maintain the shape), Russia: Ural Mountains, Orenburg Region, Kyvandyk % Village, 1−11.vi.1995, slide 2230 Sihvonen (coll. Skou), paratype.

Synopsia sociaria (Hübner, 1799), but when all data is considered, genus Phthonandria Warren, 1894 is more appropriate combination (see illustrations of some species in Kim & Beljaev 2001). Further, Phthonandria emaria (Wehrli, 1941) is thus far the nearest species from the DNA barcode point of view. Synopsia and Phthonandria share many similar features, and they are part of a larger Gnophini complex including genera such as Chariaspilates Wehrli, 1953, Angerona Duponchel, 1829, Synopsidia Djakonov, 1935 and Hypoxystis Prout, 1915, and a thorough analysis is needed to better understand their relationships. The elongated, medially concave signum and large and separated transtilla arms (hemitranstilla) are among the shared characters in some of these genera (see illustrations in Skou & Sihvonen 2015). The specimens in southern Ural are different from all other known material, but we do not describe it formally because current material is geographically limited and the observed differences are likely to be gradual when more material becomes available. The DNA barcodes differ by 0.39% only from central and eastern Asian populations. The distinctive appearance of Ural population has been noted by Alphéraky (in Staudinger 1896), but we have failed to find other references to Ural specimens in the literature. Material from southern Urals has been scarce in the collections until recently, perhaps explaining why this apparently distinctive population has not been illustrated in the modern literature. Recent expeditions to southern Ural area have shown that P. strictaria can be locally rather adundant. Extensive materials are housed in private collections of Junnilainen, Kaitila and Nupponen.

Menophra Moore, 1887 Menophra Moore, [1887] 1884–1887, Lepid. Ceylon 3: 409. Type species: Phalaena abruptaria Thunberg, 1792. [Replacement name for Hemerophila Stephens, 1829 [June]]. Genus synonyms after Scoble (1999). ‡ Hemerophila Stephens, 1829 (June), Nom. Br. Insects: 43. Type species: Hemerophila abruptaria Thunberg, 1792. [Junior homonym of Hemerophila Hübner, [1817] 1806 (Lepidoptera: Glyphipterigidae)]. Ephemerophila Warren, 1894, Novit. zool. 1: 434. Type species: Hemerophila humeraria Moore, 1868. Leptodontopera Warren, 1894, Novit. zool. 1: 445. Type species: Selenia decorata Moore, 1868. Ceruncina Wehrli, 1941, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 454. Type species: Hemerophila senilis Butler, 1878. Malacuncina Wehrli, 1941, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 461. Type species: Hemerophila prouti Sterneck, 1928.

Diversity and distribution: The genus Menophra contains 65 species in the Palaearctic and Oriental regions and in Africa. 228

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External characters and abdomen: Medium-sized to large moths. Wingspan 22−44 mm. Ground colour brown, grey or sandy, partly more strongly striated. Post- and antemedial lines usually black, distinct, oblique, in between usually a brownish yellow shade. Some species with terminal area of hindwing near tornus paler. Discal spot absent from hindwing upperside. Underside with distinct postmedial lines sometimes broken into dots, discal spots on both wings present. Fringes weakly chequered. Antennae (partially) bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia (European species): Uncus bell-shaped or triangular often with beak-shaped tip. Gnathos arms fused ventrally, upturned. Valva wide, central part narrower, apex wider, roundish, densely setose. Costa basally sclerotized with projections, diagnostic for separating species. Juxta often shield-shaped with varying (diagnostic) shape of margins. Saccus wide, in centre convex or concave depending on the species. Aedeagus usually short, thicker, vesica with one or more cornuti of different length or without cornuti (M. annegreteae). Female genitalia: Papillae anales slightly sclerotized, densely covered with setae. Apophyses posteriores medium-sized, apophyses anteriores shorter. Corpus bursae usually membranous and partly striated, in some species with diverticulum. Signum present, often stellate. Remarks: Gnophini according to Jiang et al. (2017) and Beljaev (2016), and confirmed by multi-gene analysis (Murillo-Ramos et al. 2019), where Menophra is sister to a group of genera including Synopsia + Phthonandria + Angerona and numerous non-European genera. M. harterti, M. thuriferaria and some north-west African species form an own group separated by their female genitalia structure and by the larval food-plant (conifers). A revision is needed in future.

70. Menophra abruptaria (Thunberg, 1792) Phalaena abruptaria Thunberg, 1792: Diss. ent. sistens Insecta Suecica 4: 59 (Sweden). Lectotype ♂ (UZIU). Phalaena strigata Fabricius, 1794: Ent. Syst. 3 (2): 171 (England). Syntype(s) (ZMUC). Phalaena suberaria Donovan, 1798: The natural history of British insects 7: 91, pl. 251, fig. 1 (United Kingdom). Syntype(s). Geometra petrificata Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2): pl. 52, fig. 267 (Europe). Syntype(s). Boarmia petrificaria Duponchel, 1829: in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 7 (2): 375, pl. 161, fig. 3. Emendation of petrificata Hübner. Hemerophila barcinonaria Bellier, 1862: Annls Soc. ent. Fr. (4) 2 (1): 127, pl. 1, fig. 2 (Spain: near Barcelona). Syntype(s) ♂. Hemerophila abruptaria var. dalmata Galvagni, 1907: Verh. zool.-bot. Ges. Wien 57 (1): (30) (Dalmatia: Meleda Island, Govedjari). Holotype ♀. (synonymy after Leraut 2009, here confirmed). Hemerophila abruptaria theobromaria Turati, 1909: Naturalista sicil. 21: 111, pl. 6, fig. 42 (Italy: San Remo). Holotype ♂. Hemerophila abruptaria f. maura Oberthür, 1913: Études Lépid. comp. 7: 255, pl. 165, fig. 1609 (Tunisia: Tunis). Lectotype ♂ (ZFMK). Valid at subspecific rank (Scoble 1999). Hemerophila abruptaria murina Oberthür, 1913: Études Lépid. comp. 7: 255, pl. 165, fig. 1608 (France: Menton). Syntype(s). Unavailable names (infrasubspecific): aestivalis: Dannehl (1929) (f.); brunneata: Tutt (1898) (ab.); fuscata: Tutt (1898) (ab.).

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External characters and abdomen: Wingspan ♂ 24−37 mm, ♀ 28−40 mm, second generation somewhat smaller. Ground colour lighter or darker reddish brown. Fringes weakly chequered dark and brownish. Forewing: Postmedial line blackish, slightly wavy, oblique, right-angled near costa. Antemedial line blackish, straight or slightly bent, angled near costa. Wavy line often with black dot between veins M3 and CuA2, ending in a blackish streak towards apex. Discal spot distinct, black, small. Hindwing with slightly wavy postmedial line and a weak medial line. Subterminal line diffuse, bent, usually dividing paler terminal area from rest of wing. Discal spot absent. Underside with black, serrate postmedial lines, distinct discal spots on both wings and light apical blotch on forewing. Abdomen, frons and collar dark brownish, thorax concolorous with ground colour. Abdomen with indistinct lighter and darker rings on each segment. Antennae partially bipectinate in male, about 12 apical flagellomeres free of branches, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Very variable. Ground colour varies from light sandy or pale greyish to copper-­ brown. Forewing terminal area of contrasted forms are divided in four parts as in M. berenicidaria: apex und central part usually lighter than ground colour, area between darker, sometimes blackish, tornal part usually darker brownish, never black. Hindwing with terminal area at tornus lighter than ground colour. Intermediate forms less contrasted, other forms nearly unicolorous. Melanic forms blackish brown with brownish yellow subterminal line on hindwing. Male genitalia: Uncus bell-shaped with beak-shaped tip. Gnathos arms fused ventrally, upturned, with wide ending. Valva wide, central part narrower, apex wider, roundish, densely setose. Half length of costa sclerotized with two finger-shaped projections. Proximal projection occasionally with two ends. Juxta plate, anteriorly rounded, lateral margins angled, posteriorly with double tip. Saccus wide, margin usually concave at centre (in M. japygiaria and M. berenicidaria convex). Aedeagus short, thick, vesica with two large cornuti of different length. Female genitalia: Papillae anales oval, slightly sclerotized, densely covered with setae. Apo­ physes posteriores on average 1.1 mm, slightly bent, apophyses anteriores shorter (on average 0.74 mm). Lamella antevaginalis membranous, lamella postvaginalis weakly sclerotized, both inconspicuous. Ductus bursae wide, membranous, anterior half striated. Corpus bursae membranous, partly striated, posteriorly a diverticulum, anterior end rounded. Signum stellate. Distribution and abundance: Atlanto-Mediterranean (Sub-Mediterranean). From England, Wales and Iberian peninsula through France including Corsica, Italy including Sardinia, Sicily and the island Pantelleria, to western Balkan peninsula including Greece and Crete. In Germany restricted to a small area in the south-westernmost part, erroneously recorded for the rest in Leraut (2009). – Common, sometimes found in numbers. – Outside Europe recorded for Turkey (Koçak & Kemal 2009) and in North Africa from Morocco (nomino­ typical subspecies) to Algeria and Tunisia (subsp. maura). Phenology: Uni-, bi- or trivoltine: Generally from late March to mid-July and from early August to mid-October, sometimes even until mid-November. Probably the generations overlap. In south-western Germany from early April to late May (Ebert 2003), in Britain from mid-April to early June (Skinner 2009) and in southern Spain in every month of the year (P. Hale pers. comm.). Larva through most of the spring and summer until late October. Hibernation as pupa. Adults active at night, both sexes readily attracted to light. 231

Biology: Larva polyphagous. Found on Linaria vulgaris (Ebert 2003). Recorded on Quercus, Ulmus, Clematis vitalba, Ribes rubrum, Prunus, Laburnum anagyroides, Cytisus, Genista, Spartium junceum, Ulex, Dorycnium, Tilia, Arbutus unedo, Syringa vulgaris, Jasminum, Ligustrum, Phillyrea, Lonicera, Smilax (Flamigni et al. 2007), Ruta and Rosa (Redondo et al. 2009). Reared on Betula and Fraxinus (Flamigni et al. 2007). Pupation in a tough cocoon spun up on a food-plant twig (Porter 2010). Habitat: Xerothermophilous. In a large variety of habitats such as Mediterranean macchia, open forests, open places with scattered bushes and scrub, semi-deserts. In Britain in woodland, commons, parkland and gardens (Skinner 2009). From sea-level up to 1400 m in the Iberian peninsula (Redondo et al. 2009). Similar species: M. japygiaria and M. berenicidaria (Text-figs 134–136). Genetic data: BIN: BOLD:AAB3582 (n=44 from United Kingdom, Austria, Switzerland, Spain, France, Italy, Croatia, Greece, Morocco). Genetically somewhat heterogeneous without clear geographic patterns, maximum variation 1.3%. One more strongly diverging singleton from Austria (1.9% minimum distance) possibly due to pseudogene-sequencing. Nearest species: M. japygiaria (5.9%).

71. Menophra japygiaria (Costa, 1849) Boarmia japygiaria Costa, 1849: Fauna Regno Napoli (Ent.): 384, pl. (Geom.) 9, fig. 5 (Italy, Puglia: near Lecce; cf. Flamigni et al. 2007). Syntype(s) ♂. Hemerophila fractaria Staudinger, 1859: Stettin. ent. Ztg. 20 (7–9): 218 (Spain: Andalusia, Chiclana). Holotype ♂. ‡ rhizolitharia Rambur, 1866: Cat. syst. Lépid. Andalousie 2: pl. 17, fig. 6 (Spain: Andalusia). Syntype(s). Uninominal, not available.

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Hemerophila japygiaria var. fidelensis Mendes, 1909: Broteria 8 (2): 71, pl. 13, figs 1, 5, 6 (Portugal). Syntypes 1♂4♀ (downgraded to synonymy in Redondo et al. 2009, confirmed by M. Corley pers. comm.). Hemerophila japygiaria f. sagarraria Turati, 1924: Atti Soc. ital. Sci. nat. 63 (1/2): 125, pl. 4, fig. 37 (Libya: Cyrenaica). Syntype(s). Turati (1924, p. 126 (110)) considers sagarraria as belonging to japygiaria, comparing it also with berenicidaria. Turati’s (1924) pictures on plate IV show two typical japygiaria from Algeria and from Calabria. It is not excluded that the taxon sagarraria will need to be moved to the synonymy of M. berenicidaria after examination of the types. Unavailable names (infrasubspecific): capsitanaria: Chrétien (1916) (ab.); periphanaria: Dannehl (1933) (ab.).

External characters and abdomen: Wingspan ♂ 22−33 mm, ♀ 29−34 mm. Ground colour dark ochreous to dark brown, densely striated with grey. Fringes chequered dark grey and light brownish grey. On forewing post- and antemedial lines and discal spots distinct, black. Postmedial line trapeziform near costa, antemedial line bent, both wavy. Wavy line often inconspicuous. Medial area usually lighter, in centre smaller. Black streak below apex. Hindwing with black postmedial line. Wavy line often weak, light, in centre rectangularly broken or obtusely angled. Discal spot usually absent. Underside with black, serrate postmedial lines, on hindwing with more distinct teeth. Discal spots on both wings present. Abdomen, frons, collar and thorax concolorous with ground colour. Abdomen with indistinct lighter and darker rings on each segment. Antennae partially bipectinate in male, about 12 apical flagellomeres free of branches, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variable, especially in contrast. Terminal area can be distinctly lighter than ground colour below apex, near inner margin of the forewing, and above hindwing tornus. Male genitalia: Uncus triangular with beak-shaped tip. Gnathos arms fused ventrally, upturned, with narrow ending. Valva wide, central part narrower, apex wider, roundish, densely setose. Half length of costa sclerotized with two projections, proximal projection tapered, distal projection digitiform with rounded tip. Juxta developed as an anteriorly rounded plate, lateral margin tapered, posteriorly with double tip. Saccus wide, convex at centre (in M. abruptaria concave). Aedeagus short, thick, vesica with two, sometimes three or four cornuti of different length. Female genitalia: Papillae anales slightly sclerotized, densely covered with setae. Length of apophyses posteriores on average 1.48 mm, apophyses anteriores shorter, on average 0.65 mm. Lamella antevaginalis sclerotized, alate, sides tapering. Lamella postvaginalis slightly sclerotized plate. Ductus bursae wide, weakly sclerotized. Corpus bursae membranous, in centre constricted to half of diameter, striated, posteriorly with diverticulum. Signum stellate, teeth anteriorly longer than posteriorly. Distribution and abundance: West Mediterranean. From Portugal through Spain and southernmost France to Italy including Sardinia, Sicily and Malta. Erroneously recorded for Greece by Leraut (2009), no verified specimens from there have been traced. – In Europe relatively common, usually found in limited numbers. – Outside Europe in North Africa: Morocco to Tunisia. Phenology: Bi- or trivoltine. From early April to mid-October, apparently with overlapping generations. In Portugal in all months of the year with peaks in May, July and September, suggesting three generations (M. Corley pers. comm.). Also in southern Spain in every month of the year (P. Hale pers. comm.). Caterpillars probably in most months of the year. 233

Half grown caterpillars found in early March (M. Leipnitz pers. comm.). In rearings a short pupal time of only two weeks (D. Fritsch pers. comm.). Both larva, pupa and the adult moth can be found during the winter. Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Olea europaea (M. Leipnitz pers. comm.). Recorded on Rhus coriaria, Searsia tripartita, Ziziphus lotus and Olea europaea. Reared on Syringa vulgaris (Flamigni et al. 2007). Pupation in a solid cocoon, in rearings placed between plant litter on the ground (M. Leipnitz pers. comm.). Habitat: Xerothermophilous. In warm open places or places with scattered trees and bushes, in macchia, both growing low and higher. Mainly a lowland species, but towards the south also at higher altitudes: in Sicily up to 1200 m (Flamigni et al. 2007). In Spain recorded from 2000 m (Redondo et al. 2009), but specimens from such high altitudes are probably migrants from lower lying localities. Similar species: M. abruptaria and M. berenicidaria (Text-figs 134–136). Genetic data: BIN: BOLD:AAB6706 (n=32 from Spain, Italy, Malta, Morocco, Tunisia). Genetically homogeneous. Nearest species: M. berenicidaria (3.5%).

72. Menophra berenicidaria (Turati, 1924) Hemerophila berenicidaria Turati, 1924: Atti Soc. ital. Sci. nat. 63 (1/2): 121, pl. 4, fig. 35 (­Libya: Cyrenaica, Benghasi). Lectotype ♂ (ZFMK, examined by Fischer & Lewandowski 2004). Hemerophila trypanaria Wiltshire, 1948: Entomologist’s Rec. J. Var. 60: 83, pl. 3, fig. 3 (N. ­Cyprus: Kyrenia Mountains). Holotype ♀. Downgraded from species rank to synonymy by Fischer & Lewandowski (2004).

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Hemerophila trypanaria cretacaria Reisser, 1958: Z. wien. ent. Ges. 43 (7): 119 (Crete: Silva Rouwa, Ida Mountains, 1300 m). Holotype ♂. Valid at subspecific rank (Scoble 1999), herewith downgraded to synonymy (see under Remarks). Unavailable names (infrasubspecific): coffearia: Turati (1924) (f.); uniformis: Reisser (1958) (f.); variegata: Reisser (1958) (f.).

External characters and abdomen: Wingspan ♂ 23−29 mm, ♀ 27−33 mm. Ground colour pale grey to dark brown, partly densely striated with grey. Fringes chequered dark grey and light brownish grey with weak longitudinal line. On forewing post- and antemedial lines distinct, black. Postmedial line right-angled near costa. Black streak below apex, ending in short teeth in medial area. Antemedial line bent, slightly wavy, oblique. Medial area and parts of terminal area usually lighter. Hindwing with black, slightly wavy postmedial line, angled on Rs. Wing paler at termen and basal area. Underside with black postmedial lines, dentate on veins, not angled as on upperside, with pale apical blotch, terminal area often darker. Discal spot on forewing small, absent from hindwing, on underside of both wings distinct. Frons and thorax concolorous with ground colour, collar blackish grey, abdomen blackish brown. Antennae partially bipectinate in male, about 12 apical flagellomeres free of branches, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Very variable in ground colour (see above) and contrast. Contrasted forms with forewing terminal area divided in four parts as in abruptaria: apical and medial area usually lighter than ground colour, area between darker, sometimes blackish, tornal part usually darker brownish, never black. Medial area lighter as well as the terminal blotch of the hindwing. Other forms less contrasted, appearing coppery brown. Discal spot on forewing sometimes absent. Without constant and significant subspecific differences between populations from different East Mediterranean localities.

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Text-figs 134−136. Diagnostic characters (indicated) of the Menophra abruptaria species-group. Text-fig. 134. M. abruptaria (Thunberg, 1792), France, Pyrenees, Olette, 07.x.1988. Text-fig. 135. M. japygiaria (Costa, 1849), Tunisia, Tabarka, 15.vi.1992. Text-fig. 136. M. berenicidaria (Turati, 1924), Turkey, vic. Antalya, 07.iii.1996 (all coll. B. Müller).

Male genitalia: Uncus bell-shaped with beak-shaped tip. Gnathos arms fused ventrally, upturned, with wide ending. Valva wide, central part narrower, apex wider, roundish, densely setose. Half length of costa sclerotized with one band-shaped projection, concave ending in two tips. Juxta developed as anteriorly rounded plate, lateral margins tapered, posteriorly with double tip. Saccus wide, margin convex. Aedeagus short, thick, vesica with two or three cornuti of different length. 236

Female genitalia: Papillae anales slightly sclerotized, densely covered with setae. Length of apophyses posteriores on average 1.45 mm, apophyses anteriores shorter, on average 0.64 mm. Lamella antevaginalis sclerotized, alate, sides tapered. Lamella postvaginalis slightly sclerotized plate. Ductus bursae wide, weakly sclerotized. Corpus bursae membranous, in centre constricted to half of diameter, partly striated, posteriorly with diverticulum. Signum stellate, round with short teeth. Distribution and abundance: East Mediterranean. In Europe only in southern Greece, Crete and some of the Aegean Islands including Samos. Recorded from southern Croatia (Lepiforum 2018: as 'japygiaria') – A poorly known species with rather few and scattered records only. – Outside Europe in Turkey, Cyprus and Libya. Phenology: Bivoltine. From early February to June and from September to October (Fritsch et al. 2014). Larva from early May to mid-August, the moths hatching in mid-September (D. Fritsch pers. comm.; observation from ex ovo rearings). Hibernation probably as pupa. Adults active at night, readily attracted to light. Biology: Larva possibly polyphagous. Reared on Genista and Origanum vulgare (D. Fritsch pers. comm.). Habitat: Xerothermophilous. In warm or very warm open places, macchia and also in cultivated areas. From sea-level up to at least 850 m. Similar species: M. abruptaria and M. japygiaria (Text-figs 134–136). Genetic data: BIN: BOLD:AAC8802 (n=16 from Greece/Crete, Turkey). Genetically homogeneous. One strongly diverging specimen (5.0%) with typical coloration of berenicidaria from Libya, i.e. from the type locality of berenicidaria, questioning the postulated synonymy of trypanaria Wiltshire, 1948 with berenicidaria. Nearest species: M. japygiaria (3.5%). Remarks: Scoble (1999) considers the taxon cretacaria Reisser, 1958 as valid at subspecies rank. We cannot see any differences between specimens from different East Mediterranean localities. Even the variability is the same. Therefore we do not recognize this taxon as valid at subspecies level.

73. Menophra harterti (Rothschild, 1912) Hemerophila harterti Rothschild, 1912: Novit. zool. 19: 127 ([eastern] Algeria: El Kantara). Syntypes 2♂. Hemerophila solieraria f. powelli Oberthür, 1913: Études Lépid. comp. 7: 257, pl. 165, figs 1613, 1614 (Algeria: Aflou). Syntypes ♂♀. Boarmia dresnayi Lucas, 1932: Bull. Soc. ent. Fr. 37 (11): 166 (Algeria: Oran, Frenda; Sebdou; Sidi-bel-Abbés). Syntype(s). Boarmia dubiosa Albers & Warnecke, 1941: Mitt. münchn. ent. Ges. 31 (1): 126, pl. 2, fig. 5 (Morocco: Ouida). Syntypes 2♂. Boarmia undulosa Albers & Warnecke, 1941: Mitt. münchn. ent. Ges. 31 (1): 127, pl. 2, fig. 6; pl. 4, figs 33, 36 (Morocco: Southern Rabat, Korifla; Zimeri). Syntypes 4♂. Menophra harterti atlasica Leraut, 2009: (Morocco, Middle Atlas, Ifrane). Holotype ♂. Unavailable names (infrasubspecific): brunneata: Leraut (2009) (f.).

External characters and abdomen: Very similar to M. thuriferaria. On average smaller (wingspan 22–28 mm), pattern darker. Male and female genitalia are diagnostic (see plate).

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Male genitalia: Generally as in M. thuriferaria (see above). Uncus and gnathos identical. Valva slightly larger, in the centre less constricted. Basal half of costa sclerotized, with one naked extension, apex setose, round, shorter than in M thuriferaria. Juxta and saccus without differences. Aedeagus narrow, vesica usually with two cornuti of different length, aedeagus slightly smaller than in M. thuriferaria. Female genitalia: Generally as in M. thuriferaria. Papillae anales slightly sclerotized, densely covered with setae. Apophyses posteriores and anteriores identical. Sterigma inconspicuous. Ostium bursae wide, membranous, funnel-shaped. Ductus bursae long, elongated, sclerotized, distinctly striated. Posterior part of corpus bursae also with membranous globular structure. Anterior part as in M. thuriferaria, but wider constricted posterior and anterior part. Signum stellate, weakly sclerotized, teeth shorter than in M. thuriferaria. Distribution and abundance: Central Mediterranean. In Europe only in southern Sardinia (Flamigni et al. 2007), earlier records from northern Sardinia misidentified (Flamigni et al. 2016). – In Sardinia apparently rare, just found in single specimens. – Outside Europe in North Africa, from Morocco to Tunisia, for unclear conspecifity of Moroccan populations see under genetic data.

Phenology: Univoltine (bivoltine?). Scarce Sardinian records from September and October. In Morocco in June and October (Rungs 1981). No data available on larval phenology or on hibernation. Adults active at night, attracted to light (in Sardinia). Biology: Unknown. Habitat: Xerothermophilous. Mediterranean shrubland close to the coast where Juniperus phoenicea grows (Flamigni et al. 2007).

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Similar species: See under M. thuriferaria. Genetic data: BIN: BOLD:AAD2912 (n=7 from Tunisia, Italy: Sardinia). Genetically heterogeneous, populations from Sardinia and Tunisia slightly diverging by 0.8%, but those from Morocco (n=12) though BIN-sharing nevertheless diverging by 2.5% (minimum pairwise distance), two further specimens from Libya (Cyrenaica) by 2.3% (and 1.9% from Moroccan populations). The two taxa undulosa Albers & Warnecke, 1941 and dubiosa Albers & Warnecke, 1941 both described from Morocco and both currently retained as synonym with M. harterti (Scoble 1999) possibly requiring to be upgraded to subspecies or species ranks. Nearest species (in Europe): M. thuriferaria (8.0%).

74. Menophra thuriferaria (Zerny, 1927) Boarmia thuriferaria Zerny, 1927: Eos, Madr. 3: 422, pl. 10, figs 27, 28 (Spain: Prov. Teruel, Albarracín). Syntype(s) (ZFMK).

External characters and abdomen: Wingspan ♂ 23−26 mm, ♀ 24−30 mm. Ground colour brownish grey. Fringes grey, only weakly chequered with dark grey. On forewing post- and antemedial lines black, oblique, sub-parallel. Medial area brownish filled with sharp discal spot. Postmedial line near costa zigzagging, here hardly visible. Apical streak distinct. Terminal area brown at postmedial line, towards termen with diffuse light greyish band. Hindwing with black, straight or slightly bent postmedial line, densely striated with grey, terminal area paler, without discal spot. Underside with serrate postmedial lines, discal spots on both wings present. Frons white, collar whitish, tips of scales brownish, thorax grey. Segments of abdomen tricolorous with transverse whitish, mid-brown and dark brown rings. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Uncus triangular with beak-shaped tip, laterally wavy, weakly setose. Gnathos arms fused ventrally, upturned, ending rounded. Valva wide, central part narrower, apex wider, roundish, densely setose. Half length of costa sclerotized, with two naked extensions, basal one larger, apical one very small (in M. harterti only one extension, being longer). Juxta anteriorly concave, lateral margin straight, posteriorly slender with weak double tip. Saccus wide, margin convex at centre. Aedeagus slender, vesica with one or two (exceptionally three) cornuti of different lengths, slightly larger than in M. harterti. Female genitalia: Papillae anales slightly sclerotized, densely covered with setae. Length of apophyses posteriores on average 1.10 mm, apophyses anteriores shorter, on average 0.68 mm. Sterigma inconspicuous. Ostium bursae wide, membranous, funnel-shaped. Ductus bursae long, elongated, sclerotized, distinctly striated. Posterior part of corpus bursae with membranous globular structure. Anterior part membranous, elongate-oval, partly striated around signum. Shortly constricted between two parts (this part longer in M. harterti). Signum stellate, anteriorly with longer teeth. Distribution and abundance: Endemic in Spain. One putative record from southern France (Cannes), considered doubtful by Leraut (2009). – Somewhat local and uncommon, occurring in limited numbers. Phenology: Bivoltine. From March to June and from September to December (Redondo et al. 2009). Additional records in mid- and late July and late August possibly indicating the overlap of the two generations. In southern Spain recorded in all months of the year except September and December (P. Hale pers. comm.). Adults active at night, readily attracted to light. 239

Biology: Oligophagous (1) on Cupressaceae. Recorded on Juniperus, especially J. thurifera and Cupressus. Also recorded in urban Cupressus hedges (G. King pers. comm.). Habitat: Xerothermophilous. In open shrub- and woodland. From close to sea-level up to 1600 m. Similar species: The allopatric M. harterti can hardly be separated from M. thuriferaria by external features, only by examination of the male genitalia (see above) and/or DNA barcoding. Ecleora solieraria and Peribatodes aragonis can be easily separated, different pattern in postmedial line and apical streak are diagnostic, see Text-figs 167–169. Ekboarmia atlanticaria is also similar, see Text-fig. 159. Genetic data: BIN: BOLD:AAD2913 (n=9 from Spain). Genetically homogeneous. Nearest species: M. harterti from Tunisia and Sardinia (8.0%). Remarks: Several authors considered M. thuriferaria as a subspecies of M. harterti (Scoble 1999; Flamigni et al. 2007; Redondo et al. 2009). Genetic data show that M. thuriferaria is not conspecific with M. harterti.

75. Menophra nycthemeraria (Geyer, 1831) Geometra nycthemeraria Geyer, 1831: Samml. Eur. Schmett. 5 Geometrae (1): pl. 108, fig. 564 (Switzerland: C. Wallis). Syntype(s). Hemerophila nycthemeraria dinicola Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 452, pl. 39: e (France: Alpes Maritimes, Digne). Syntype(s). Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features. Unavailable names (infrasubspecific): paucistrigata: Leraut (2009) (f.); perstrigata: Leraut (2009) (f.).

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External characters and abdomen: Largest European Menophra species. Wingspan ♂ 34− 40 mm, ♀ 36−44 mm. Ground colour ash-grey to sandy brown. Fringes chequered grey and brownish. Forewing with broad silver-grey costal band, finely striated with brownish grey. Postmedial line black until apical streak, slightly wavy, oblique, near costa right-angled, pale brown and hardly visible. Distal half of costal band separated by apical streak. Antemedial line nearly straight, black, ending at costal band. Postmedial line on terminal area, antemedial line on basal area dark shadowed. Discal spot distinct, black, small. Hindwing densely striated with brownish grey, terminal area darker, without discal spot. Postmedial line black, nearly straight. Underside with postmedial lines broken into dots, with discal spots on both wings. Frons black, collar blackish ringed, thorax bicolorous, anterior half pale, posterior half blackish grey. Abdomen concolorous with ground colour. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variable in brightness of wings. In darker specimens the shadows on the post- and antemedial lines of the forewing can be extended to the whole medial area, i.e. a broad dark brownish band connecting the basal inner margin with the terminal area below the apex. Populations of south-eastern France (‘f. dinicola’) with light brownish (beige) medial area and reduced striation. Similar forms often in southern Alps. Male genitalia: Uncus triangular, setose, with small bent tip. Gnathos arms fused ventrally, upturned, ending digitiform. Dorsal arm of valva digitiform, curved with small rounded projection, less than half length of ventral arm. Ventral arm of valva wide, central part narrower, apex wider, roundish, densely setose. Juxta developed as plate, anteriorly rounded, lateral margins straight, posteriorly concave, wavy. Saccus wide, central margin concave. Aedeagus thick, vesica with group of about six short cornuti of different lengths. Female genitalia: Papillae anales slightly sclerotized, densely covered with setae. Length of apophyses posteriores on average 1.25 mm, apophyses anteriores shorter, on average 0.90 mm. Lamella antevaginalis small, membranous, alate, ‘wings’ upright. Lamella ­postvaginalis slightly sclerotized triangular plate. Ductus bursae wide, sclerotized. Ductus seminalis leads to wide, more strongly sclerotized diverticulum. Corpus bursae drop-shaped, membranous, weakly striated. Signum oval, without teeth, anteriorly more strongly sclerotized. Distribution and abundance: West Mediterranean. In Europe only known from Valais in Switzerland, southern France, north-westernmost Italy, Spain and Portugal. Three specimens recorded in Thuringia, Germany from 1936 and earlier (see open circles on map). – Rather local and rare, usually found in limited numbers. – Outside Europe only in Morocco. Phenology: Univoltine, at lower altitudes possibly bivoltine. Usually from late April to July. According to Redondo et al. (2009) until October. In Portugal one generation from April to July, peaking in June (M. Corley pers. comm.). Caterpillars from July or August to April or May, overwintering. Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Cytisus (M. Leipnitz pers. comm.) and Dorycnium pentaphyllum (T. Varenne pers. comm.). Recorded on Genista, Retama (Redondo et al. 2009), Juniperus oxycedrus and Buxus (Ziegler 2019). Reared on Genista (D. Fritsch pers. comm.). Pupation among plant litter on the ground (M. Leipnitz pers. comm., rearing observation). A cocoon observed between leaves of Buxus (T. Varenne pers. comm.).

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Habitat: Xerothermophilous. Open places with scattered trees and bushes. In Spain from 400 up to 1600 m, but apparently most common above 1000 m. In southern France from 300 m.

Similar species: Differing from other Menophra species by its broad silver-grey costal band on the forewing. Genetic data: BIN: BOLD:ADM1765 (n=1 from central Spain). Nearest species: Nearctic Iridopsis perfectaria (McDunnough, 1940) (6.6%), nearest species in Europe Peribatodes perversaria (7.6%), nearest congener M. senilis (Butler, 1878) from Japan (8.8%). Genetic data suggesting this species to be the product of a long, isolated evolutionary lineage.

76. Menophra annegreteae Skou, 2007 Menophra annegreteae Skou, 2007: Nota lepid. 29 (3/4): 138–143 (Spain, Almeria: Cabo de Gata, 10 m). Holotype ♂ (ZMUC).

External characters and abdomen: The smallest European Menophra species: Wingspan 22−30 mm, ♂ a little smaller than ♀. Specimens of the summer generation are smaller, up to 24 mm only. Ground colour brownish to brownish grey with dark mottling. Fringes weakly chequered dark and pale grey. Terminal line blackish. Forewing: Subterminal line greyish with lighter edging, often indistinct. Postmedial line oblique, directed to apex, right-angled near costa. Antemedial line slightly wavy, not reaching costa. Discal spot distinct, black, small. Hindwing with distinct, sligthtly curved postmedial line and indistinct discal spot. On underside discal spots more distinct. Medial line not reaching costa. Abdomen, thorax, frons and collar concolorous with ground colour. Abdomen with two darker, often fused dots on each segment. Antennae bipectinate with long branches in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. 242

Variation: Little. Ground colour, dark mottling and distinctness of lines vary. Male genitalia: Uncus triangular, setose, tip bent. Gnathos arms fused ventrally, upturned, spatulate. Valva small, central part distinctly narrower, apex wider, roundish, densely setose. Ventral margin strongly concave. Base of costa sclerotized with a wide band-shaped projection, concavely ending, in two tips. Juxta shield-shaped, anteriorly rounded, lateral margins bent, posteriorly concave, wavy. Saccus elongated, margins laterally concave. Aedeagus wide, tapered towards apex, ending in spine-shaped process. Vesica membranous, without cornuti. Female genitalia: Papillae anales oval, slightly sclerotized, densely covered with setae. Apophyses posteriores on average 1.35 mm long, slightly bent. Apophyses anteriores shorter, on average 0.83 mm. Lamella antevaginalis sclerotized, alate, sides tapered. Lamella postvaginalis membranous. Ductus bursae wide, weakly sclerotized, narrower than posterior end of corpus bursae. Corpus bursae membranous, elongated, anterior end larger, oval, posterior end narrower. Signum absent. Distribution and abundance: A south-eastern Iberian endemic, so far only known from a very limited distribution area. – Apparently rather local but not rare where occurring and sometimes found in numbers. – Not recorded from outside Europe.

Phenology: The few specimens so far recorded not allowing unambiguous determination of flight period, probably being trivoltine. Scarce records from mid-April to late May and in late July, mid-September and late October. Always present in a number of visits to the species’ best known locality in the Almeria province (P. Skou own data). Larval time and hibernation unknown. Adults active at night, both sexes attracted to light, but males more than females. Biology: Unknown. 243

Habitat: Xerothermophilous. Semideserts. From sea-level up to 400 m. Genetic data: BIN: BOLD:AAG9445 (n=3 from southern Spain). Genetically homogeneous. Apparently the product of a long, isolated evolutionary line, no species at a closer distance than 8%, e.g. an undescribed Menophra from Tanzania (9.6%), in the western Palaearctic no species closer than 10%.

Cleorodes Warren, 1894 Cleorodes Warren, 1894, Novit. zool. 1: 434. Type species: Phalaena lichenaria Hufnagel, 1767. There are no genus synonyms.

Diversity and distribution: Cleorodes is a monotypic genus, after the synonymization of Cleorodes incerta Rungs, 1975 with Peribatodes ilicaria Geyer, 1833 (Leraut 2009). The only species occurs in the western Palaearctic region. External characters, abdomen and genitalia: See species description below. Remarks: Gnophini according to Viidalepp et al. (2007), confirmed by multi-gene analysis (Murillo-Ramos et al. 2019). In the latter Cleorodes being sister to Siona, both of which are sister to Menophra.

77. Cleorodes lichenaria (Hufnagel, 1767) Phalaena lichenaria Hufnagel, 1767: Berlin Mag. 4 (5): 512 (Germany: Berlin region). Syntype(s). Geometra pictaria Thunberg, 1784: Diss. ent. sistens Insecta Suecica: 6 (Sweden: Halland, Upp­ land). Lectotype ♂ (UZIU). Phalaena Geometra dentilinearia Borkhausen, 1794: Natur. eur. Schmett. 5: 231 (Europe). Syntype(s). Phalaena ziczagaria Schrank, 1802: Fauna boica 2 (2): 6 (Germany: Bavaria). Syntype(s). Boarmia lichenaria var. jaspidaria Wagner, 1919: Z. öst. EntVer 4 (2): 11 (Hungary). Holotype ♂. Validated at subspecific rank by Scoble (1999), not mentioned in Leraut (2009). For valid­ity of subspecies rank see remarks. Boarmia leukocyrnea Schawerda, 1929: Z. öst. EntVer 14 (3): 30 (Corsica: Col de Vizzavona). Syntypes. Cleora lichenaria f. obscuraria Schneider, 1941: Ent. Z., Frankf. a. M. 54 (28): 224 (locality not stated). Holotype ♂. Boarmia lichenaria distinctaria Hanson, 1962: Ent. Tidskr. 83 (3–4): 131, fig. 3 (Sweden: Gotland, Sandön, Tallskogen). Holotype ♂ (NHRS). Validated at subspecific rank by Scoble (1999), not mentioned in Leraut (2009). For validity of subspecies rank see remarks. Unavailable names (infrasubspecific): chretieni: Lhomme (1929) (ab.); cineraria: Borkhausen (1794) (ab.).

External characters and abdomen: Wingspan 28–31 mm, second generation on average somewhat smaller. Ground colour whitish grey, more or less mottled with blackish grey dots and streaks. Sometimes with greenish tinge. Post- and antemedial lines black, distinct, on forewing darker towards costa. Postmedial line jagged, near inner margin simply angled, strongly bent inwards in the lower third, distally sometimes associated with brown band. Antemedial line bent, with three arrow marks on veins. Hindwing with jagged postmedial line. Terminal line dotted. Fringes chequered dark and whitish grey. Frons and collar concolorous with ground colour. Antennae bipectinate in male, filiform in female. Hindlegs of both sexes with 2+2 spurs. 244

Variation: Depending on the degree of dark mottling wings can be paler or darker. Terminal and basal area can be brown or reddish brown darkened. Male genitalia: Uncus triangular, double tip rounded, beak-shaped, bent. Gnathos vestigial. Valva simple, narrow, apical third setose. Costa convex, ventral margin concave. Transtilla conspicuous, posteriorly extended, ending in wide triangular, pointed structure. Juxta small, nearly square plate. Saccus wide, slightly rounded. Aedeagus short, thick, vesica with long curved spine, covering 3/4 of total length. Female genitalia: Papillae anales oval, weakly sclerotized, setose. Length of apophyses posteriores 1 mm. Apophyses anteriores shorter, about half as long. Lamella postvaginalis sclerotized, narrow, horizontal, conspicuous. Ductus bursae wide, partly sclerotized. Corpus bursae membranous, striated. Signum small, irregularly formed, sometimes weak. Distribution and abundance: European-West Asiatic. From south-western Russia to the British Isles and from southern Finland and central Sweden to northern Spain, southern France including Corsica, Italy including Sardinia and Sicily and the Balkan peninsula down to southern Greece. – Uncommon, usually occurring as single specimens or very limited numbers. Declined in many parts of its European distribution area, probably due to decline of suitable food-plants caused by air pollution. – Outside Europe in Turkey, Armenia and Georgia.

Phenology: Univoltine or partly bivoltine. In Denmark from early June to mid-August and with a partial second generation from early September to early October. In south-western Germany also from early June to mid-August, but with very few second generation specimens around 1st October (Ebert 2003). In southern Italy at low altitudes in two generations from mid-May to mid-October, further to the north, at higher altitudes and also in Sicily and Sardinia in one generation from June to August (Flamigni et al. 2016). Larva from August to June, hibernating. Adults active at night, attracted to light and sugar bait. 245

Biology: Larva oligophagous on lichens. Found on Ramalina farinacea and R. fastigiata (Wegner 2013). Repeatedly recorded on lichens growing on trees and bushes, rocks, walls etc. (e.g. Freyer 1831–33), recorded on Usnea barbata by Allan (1949). Pupation among lichens on tree trunks (Wegner 2013). Habitat: In a variety of habitat types with trees and bushes such as open forests, scrubland, orchards, parks and gardens. From sea-level up to 850 m in south-western Germany (Ebert 2003) and up to 1600 m in the Pyrenees (Redondo et al. 2009). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC4210 (n=25 from Finland, Denmark, Germany, United Kingdom, France, Italy, Macedonia, Ukraine, Georgia). Genetically heterogeneous, maximum variation 1.9%, Sicilian populations diverging from the main cluster (including ­material from Calabria) by 0.9%. Nearest species: Arichanna consocia (7.9%), Calamodes sub­scudularia (8.4%). Remarks: Wegner (2013) deals in details with the biology and habitat in north-western Germany. The validity of subspecies rank for the taxa jaspidaria (Wagner, 1919) and distinctaria (Hanson, 1962) is questionable and requires further research.

Zernyia Prout, 1929 Zernyia Prout, 1929, Novit. zool. 35: 146. Type species: Gnophos enconistoides Zerny, 1927. There are no genus synonyms.

Diversity and distribution: Seven species in the western Mediterranean area, only one of them in Europe. External characters and abdomen: Medium-sized moths. Proboscis slightly reduced, fovea absent. Forewing veins R1 and R2 arising free from cell, sometimes shortly stalked. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Costa of valva distally with spines. Uncus short, wide. Aedeagus stout with sclerotized tooth. Female genitalia: Papillae anales elongated. Apophyses anteriores shorter than apophyses posteriores. Anterior part of corpus bursae membranous, round, without signum. Lamella antevaginalis, sterigma and ductus bursae strongly sclerotized. Remarks: Zernyia has not been included in a multi-gene analysis so far. Systematic position tentative. In the latest checklist of the Lepidoptera of Spain (Redondo et al. 2015) placed between Calamodes and Sardocyrnia.

78. Zernyia granataria (Staudinger, 1871) Selidosema granataria Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 171 (Spain: Andalusia). Holotype ♀ (ZSM/Herbulot). Gnophos enconistoides Zerny, 1927: Eos, Madr. 3 (4): 428, pl. 10, figs 34, 35 (Spain: Prov. T ­ eruel, Albarracín). Syntypes ♂♀ (ZFMK).

External characters and abdomen: Wingspan 26–33 mm. Ground colour sandy-brown to greyish, strongly irrorated with greyish brown, more in females than in males. Fringes concolorous with ground colour. Terminal line weak, dotted. On forewing postmedial line near246

ly straight, inwardly bent before reaching costa, on hindwing bent, on both wings usually broken into dots. Other transverse lines weak or absent. Terminal area of males with dark blotches, of varying intensity, sometimes forming a band. Terminal area of females darkened. Medial area with discal spots on all wings, on forewing and on underside more distinct. Frons ventrally beige, dorsal half brownish. Vertex, collar, tegula, thorax and abdomen concolorous with ground colour, partly irrorated with darker scales. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variation in ground colour, see above. In males, the row of dark blotches in the terminal area sometimes fused into a band, sometimes absent. The dark terminal area of females occasionally extended to cover the wings nearly all over. Male genitalia: Uncus short, stout, weakly setose, tip and margin sclerotized. Gnathos arms wide, fused ventrally, forming a rounded plate. Valva narrow, membranous, tapering. Costa sclerotized, distally with spines, forming a ridge. Sacculus weakly sclerotized, in the centre with weakly sclerotized, narrow, setose process. Juxta a wide plate, posteriorly ending in two rounded, sclerotized tips, deeply concave between. Saccus wide, rounded. Aedeagus short, slightly bent with one triangular, sclerotized tooth. Female genitalia: Papillae anales elongated, slightly sclerotized, densely covered with setae. Length of apophyses posteriores about 1.95 mm, apophyses anteriores shorter, on average 1.2 mm, both spatulate at tip. Lamella antevaginalis, sterigma and ductus bursae strongly sclerotized, wide. Lamella postvaginalis posteriorly rounded, strongly sclerotized. Posterior part of corpus bursae wide, weakly striated, anterior part of corpus bursae wider, membranous, round, without signum. Distribution and abundance: Endemic in Spain. – Somewhat local, sometimes rather abundant in its localities.

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Phenology: Univoltine. From early August to early October. In southern Spain from late August to early November (P. Hale pers. comm.). Caterpillar in rearings, according to conditions, from March to June and from November to May (M. Leipnitz pers. comm.). Overwintering both as egg and as caterpillar (M. Leipnitz pers. comm.; rearing observation). Adults nocturnal, both females and males readily attracted to light. Biology: Larva monophagous. Recorded on Calluna (Redondo et al. 2009). Reared on Bupleurum falcatum (M. Leipnitz pers. comm.). In rearings, pupation in an open web between plant litter on the ground. Habitat: Xerothermophilous. In open or semi-open and dry places with scattered bushes and trees. From 150 up to 1400 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:ABX1046 (n=2 from Spain: Aragon, Andalusia). Genetically homo­ geneous. Nearest species: Four North African congeners (correct identifications awaiting) with four different BINs at distances of 6.1–7.2%.

Megalycinia Wehrli, 1939 Megalycinia Wehrli, 1939, Mitt. münchn. ent. Ges. 29: 73. Type species: Hemerophila scalaria Christoph, 1893. There are no genus synonyms.

Diversity and distribution: Three species in the western Palaearctic region, in Europe one species only, which is endemic to Italy. One species, which was combined with Megalycinia Wehrli earlier (strictaria (Lederer)), is transferred to Phthonandria Warren, 1894 in this volume. External characters and abdomen (female diagnosis is based on M. serraria only, females of M. scalaria and M. engys are undescribed): Wingspan approximately 35 mm in both sexes. Wings beige to brown-beige, extensively irrorated with brown scales. Postmedial line distinct, blackish, undulate or weakly serrate. Wing termen distinctly undulating in M. serraria, less in other species. Underside paler, slightly shiny, weakly irrorated. Frons protruding. Male antennae bipectinate with long branches, female antennae filiform. Hindlegs of both sexes with 2+2 spurs. Tympanal organs with additional flap medially (Text-figs 137−138). Male genitalia: Uncus long, narrow, curved ventrally. Tegumen elongated. Socii minute, setose. Gnathos fused, upturned medially. Valva narrow, long, apex setose, costa sclerotized. Juxta sub-rectangular, sclerotized. Saccus weakly elongated. Aedeagus apex constricted, with spoon-shaped extension. Vesica weakly enlarged basally, with or without cornuti. Female genitalia (diagnosis is based on M. serraria only, females of M. scalaria and M. engys are undescribed): Papillae anales elongated, densely setose. Lamella antevaginalis narrow, string-like, curved. Ductus bursae short, sclerotized laterally. Corpus bursae very long, anterior end elongated sac. Signum small, weakly stellate. Biology: Larva of M. serraria found on Ribes uva-crispa (Dannehl 1927b), reared on petals of Rosa and later on Rubus and Prunus spinosa (Dannehl 1909; Noack 1939). All species usually found at high altitude, European M. serraria inhabiting almost only hills and mountains in scattered or even shady pine forest with much undergrowth of ferns and Cytisus (A.  Hausmann own data). Adults flying in autumn, typically from August to October and even to November. 248

Text-figs 137−138. Tympanal organs with additional flap medially (at the interior margin facing each other), a diagnostic character of genus Megalycinia Wehrli, 1939. Structure is present in both sexes, and in the closely related Wehrliola Strand, 1932 also (see Kemal & Uçak 2018). Textfig. 137. M. scalaria (Christoph, 1893) male, Turkey: Kars, 1500 m, 11.ix.1993, slide Sihvonen 2258 (coll. Skou). M. scalaria has not been found in Europe so far. Text-fig. 138. M. serraria (Costa, 1882) female, Italy: Calabria, Sila, 1550 m, 31.viii.2000, slide Sihvonen 2275 (coll. ZSM).

Immature stages: Larva of M. serraria grey, darker dorsally, with faint yellowish line laterally (Flamigni et al. 2016, plate 32, fig. 4). Remarks: Megalycinia has been classified in the Boarmiini (Hausmann et al. 2004; 2011a), but the systematic position has not been examined in an analytical context. Few genitalia structures are similar to the Lithinini. The non-European Wehrliola Strand, 1932 is morphologically very similar to Megalycinia (see Kemal & Uçak 2018). Systematic position: Gnophini of uncertain association.

79. Megalycinia serraria (Costa, 1882) Hemerophila serraria Costa, 1882: Atti Acad. Sci. fis. mat. Napoli 9 (6): 41, fig. 13 (Italy: Calabria). Syntype(s). Hemerophila serraria dannehli Noack, 1939: Ent. Rdsch. 56: 266, fig. 7 (Italy: Genzano). Syntypes 18♂♀. Junior synonym (Scoble 1999). Unavailable names (infrasubspecific): paucistrigata: Leraut (2009) (f.); perstrigata: Leraut (2009) (f.).

External characters and abdomen: Wingspan 27−38 mm. Wings pale, straw-coloured, weakly suffused with dark brown. Forewing base and inner part of terminal area darker than rest of wing. Postmedial line distinct, strongly undulating on forewing, curved strongly inwards and weaker before costa. Outer margin of postmedial line bordered with dark brown. Terminal area paler. Terminal line brown. Fringes concolorous with wings. Hindwing termen strongly undulating. Discal spots small, dark brown. Underside markings weaker, almost absent. Frons weakly protruding. Frons, collar, thorax and abdomen concolorous with wings. Male antennae bipectinate with long pectinations, female antennae filiform. Hindtibia of 249

both sexes with 2+2 spurs. Tympanal organs medium-sized, inner margin with additional flap (Text-fig. 138). Other abdominal segments without modifications. Variation: Very little. Forewing base sometimes concolorous with medial area (not dark) and markings sometimes weak. Terminal area often suffused with dark brown but sometimes finely striated. Leraut (2009) mentions specimens with a wingspan of up to 45 mm, such large specimens not seen in our examined material. Male genitalia: Uncus long, narrow, curved ventrally. Tegumen elongated, anterior margin with small, rounded lobes (see Genitalia plate and Remarks). Socii minute, setose. Gnathos fused, medial element smooth, upturned. Transtilla arms separate, knob-shaped. Valva narrow, long, medial element undulating, apex setose, costa sclerotized. Juxta sub-rectangular, sclerotized, posterior margin granulate. Saccus weakly elongated. Aedeagus straight, caecum short, apex weakly constricted. Vesica opens at about 135 degree angle, base weakly enlarged and covered with microcornuti. Base with distinct row of about 15 stout cornuti. Female genitalia: Genitalia very large, maximum length up to 11 mm. Papillae anales elongated, densely setose. Apophyses posteriores about twice length of apophyses anteriores. Lamella antevaginalis narrow, string-like, curved. Lamella postvaginalis sclerotized but undifferentiated. Ductus bursae short, funnel-shaped, sclerotized laterally. Posterior part of corpus bursae weakly sclerotized, striated. Corpus bursae very long, narrow, anterior end elongated sac. Signum small, variable in size and shape, centre solid and margins weakly stellate, or centre broken into small sclerotizations and margins weakly stellate. Distribution and abundance: (South-)European. Endemic in central and southern Italy with a northern distribution limit in higher altitudes of the Bolognese Apennine (Flamigni et al. 2007). – Local but not a rare species, usually found in limited numbers.

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Phenology: Univoltine. From August to late September (Flamigni et al. 2007), in southern Italy from late August and into October (A. Hausmann own data). Also recorded in late July (N. Zahm pers. comm.). Adults active at night, attracted to light. Biology: Larva mono- or oligophagous on Rosaceae and Grossulariaceae. Found on Ribes uva-crispa (Dannehl 1927b), reared on petals of Rosa and later on Rubus and Prunus spinosa (Dannehl 1909; Noack 1939; Flamigni et al. 2007). Recorded on Ribes rubrum (Leraut 2009). Habitat: Silvicolous. Mainly in shady places, such as narrow river valleys, scattered Mediterranean forest with much Quercus, or, at higher elevation in scattered or more dense pine forests with much undergrowth of ferns and Cytisus scoparius (A. Hausmann own observation). Usually from 700 m up to 1690 m, exceptionally down to 200 m in shady habitats (A. Hausmann own observation; Flamigni et al. 2007). Similar species: No similar species in Europe. Wehrliola revocaria (Staudinger, 1892) more uniform in colour, grey with tint of pale yellow, forewing postmedial line less undulating, with dark wedge near costa and not strongly curved inwards on costa, forewing discal spot elongated (in M. serraria wings pale, straw-coloured, postmedial line distinctly undulating, without wedge near costa, strongly curved inwards on costa, and discal spot not elongated). Male and female genitalia similar but diagnostic. Genetic data: BIN: BOLD:AAF3058 (n=5 from Italy). Genetically slightly heterogeneous, maximum variation 1.4%. Nearest species: Wehrliola revocaria (6.1%), Dasypteroma thaumasia (8.8%), Synopsia sociaria (9.2%). Remarks: Male genitalia of M. serraria has paired, small, rounded lobes between valva base and gnathos (see Genitalia plate). Apparently these originate from the anterior margin of the tegumen, and if so, they could potentially be termed peniculi (sing. peniculus) (Klots 1970; Gordh & Headrick 2001). These structures have not been noted in other Megalycinia species. Various appendices in the juxta region and at the base of valvae are common in the Ennominae (see for instance Hylaea Hübner, 1822, Pungeleria Rougemont, 1903 and Stegania Guenée, 1845 in Skou & Sihvonen 2015), but thus far we have not seen similar structures originating from the tegumen. Alternative point of origin of these rounded lobes could be the anellus.

Dasypteroma Staudinger, 1892 Dasypteroma Staudinger, 1892, Dt. ent. Z. Iris 5: 168. Type species: Dasypteroma thaumasia Staudinger, 1892. Paronychora Joannis, 1912, Broteria (Zool.) 10: 19,20. Type species: Enconista oberthuri Vázquez, 1905.

Diversity and distribution: Dasypteroma is a monotypic genus in the West Palaearctic. External characters, abdomen and genitalia: See species description below.

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80. Dasypteroma thaumasia Staudinger, 1892 Dasypteroma thaumasia Staudinger, 1892: Dt. ent. Z. Iris 5: 168, pl. 2, fig. 10 (Spain, Castilia San Ildefonso). Syntypes 3♀. Enconista oberthuri Vázquez, 1905: Boln Soc esp. Hist. nat.: 5 (2): 119, fig. 3a (Spain: Madrid). Holotype ♂. Enconista powelli Oberthür, 1914: Bull. Soc. Ent. Fr. 1914 (12): 387 (Algeria: Sebdou). Syntypes ♂♀. Valid at subspecific rank (Scoble 1999), here confirmed.

External characters and abdomen: Male: Wingspan ♂ 24–27 mm. Forewing ground colour pale grey to dark grey to brownish grey, more or less speckled dark grey. Postmedial line black, usually dotted, subapically bent, then straight. Antemedial line black, dotted, curved. Medial line absent. Subterminal line reduced to a row of diffuse blotches, sometimes missing. Terminal line indistinct, grey. Discal spot distinct. Fringes of ground colour. Hindwing: Ground colour and fringes as in the forewing. Postmedial line indistinct or missing, sometimes also diffuse blotches in the terminal area. Discal spot weaker. Antennae bipectinate. Collar, thorax and abdomen usually of ground colour. Female brachypterous, wing stumps narrow with long hairy scales. Abdomen dark grey to brown. Frons smoothly scaled, medial part elevated as in Megametopon grisolaria. Hindtibia of both sexes with 2+2 spurs. Variation: Males variable in degree of dark speckling. Ground colour, see above. Male genitalia: Uncus with short triangular base, weakly setose; apex very long, beakshaped curved, pointed. Gnathos ventrally fused, forming trapezoid sclerotized plate, lateral margins with some spines of different length. Valva simple, short and wide. Costa and ventral margin slightly wavy (partly concave and convex). Apex and medial area setose. Juxta bilobed. Saccus rounded. Aedeagus short, slightly curved. Apex with sclerotized projection, caecum oblique. Without cornutus. Female genitalia: Papillae anales anteriorly cylindric, posteriorly narrower and rounded, sclerotized, strongly setose. Length of apophyses posteriores about 2.5 mm, apophyses anteriores shorter (about 1.5 mm), both spatulate at tip. Lamella antevaginalis membranous. Lamella postvaginalis inconspicuous. On posterior part of the female genitalia conspicuous, narrow, sclerotized arcuate structure, which can be seen also in genus Wehrliola. Ductus bursae short, posteriorly wider, membranous. Corpus bursae long, narrow, membranous, slightly striated. Without signum. Distribution and abundance: West Mediterranean. In Europe only in the Iberian peninsula, mainly in the northern half. – Somewhat localized, relatively abundant where occurring. – Outside Europe in North Africa: Morocco (subsp. powelli). According to Redondo et al. (2009) in Algeria and according to Rungs (1981) in North Africa including Tunisia, but we have not been able to trace any records from these two countries. Phenology: Univoltine: From late August at higher altitudes and until mid-October at the lowest localities. In southern Andalusia as late as mid-November. Caterpillars in March (G. King pers. comm.), April and May. Males active at night, readily attracted to light. No information on female behaviour in nature traceable, all females in collections apparently reared from larvae. Biology: Larva polyphagous. Found mainly on Thymus lacaitae, secondly on Artemisia campestris (G. King pers. comm.) and on Lepidium subulatum (King & Romera 2004). Reared on Rumex crispus and R. acetosa (G. King pers. comm.), Thymus vulgaris and Vicia 252

(Redondo et al. 2009). Pupation on the bottom of larval container without cocoon (G. King pers. comm.; rearing observation). King & Romera (2004) describe the biology of this species in detail. Habitat: Xerothermophilous. In open localities with scattered bushes and trees, often on gypsum soil. From 100 up to 1400 m above sea-level. Similar species: Males can be similar to males of Isturgia spodiaria (treated in Vol. 5) with non-darkened terminal area. Also similar to males of (the allopatric) Megametopon griso­ laria with more indistinct pattern.

Tribus Boarmiini Duponchel, 1845 Boarmiini Duponchel, 1845 (Boarmites). Type species (of type genus Boarmia): Cymatophora umbrosaria Hübner, 1813. Genus Boarmia Treitschke, 1825 (type species: Geometra roboraria Denis & Schiffermüller, 1775) is currently considered a junior synonym of Hypomecis Hübner, 1821. Amphidasini Duponchel, 1845 (Amphidasites). Type species (of type genus Amphidasis, junior synonym of Biston): Geometra prodromaria Denis & Schiffermüller, 1775, a junior synonym of Phalaena strataria Hufnagel, 1767. Junior subjective synonym (Holloway 1994: 5). Used for concept of Bistonini by Seven (1991). Cleorini Duponchel, 1845 (Cleorites). Type species (of type genus Cleora): Geometra cinctaria Denis & Schiffermüller, 1775. Junior subjective synonym (Holloway 1994: 6), synonymy is supported by molecular results also (Sihvonen et al. 2011; Murillo-Ramos et al. 2019). Fidoniini Duponchel, 1845 (Fidonites). Type species (of type genus Fidonia): Phalaena plummistaria Villers, 1789. Genus Fidonia Treitschke, 1825 is currently considered a junior synonym of Eurranthis Hübner, 1823.

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Hyberniini Duponchel, 1845 (Hibernites). Type species (of type genus Hybernia): Phalaena defoliaria Clerck, 1759. Genus Hybernia Berthold, 1827 is currently considered a junior synonym of Erannis Hübner, 1825. Junior subjective synonym (Holloway 1994: 7). Bistonini Stephens, 1850 (Bistonidi). Type species (of type genus Biston): Geometra prodromaria Denis & Schiffermüller, 1775, a junior synonym of Phalaena strataria Hufnagel, 1767. Junior subjective synonym (Holloway 1994: 13, 167), here confirmed as the inclusion of the Bistonini in the Boarmiini is supported by DNA phylogenies (Wahlberg et al. 2010; Sihvonen et al. 2011; Murillo-Ramos et al. 2019). Viidalepp et al. (2007) presented morphological support for the Bistonini being monophyletic, but noted that this might make Boarmiini sensu stricto paraphyletic. Selidosemini Meyrick, 1892 (Selidosemidae). Type species (of type genus Selidosema): Geo­ metra plumaria Denis & Schiffermüller, 1775. Junior subjective synonym (Holloway 1994: 9), supported by molecular data also (Murillo-Ramos et al. 2019). Ascotini Warren, 1893 (Ascotinae). Type species (of type genus Ascotis): Geometra selenaria Denis & Schiffermüller, 1775. Junior synonym (Sato 1984; Holloway 1994). Eubyjini Warren, 1893 (Eubyjinae). Type species (of type genus Eubyia, junior synonym of Biston): Phalaena betularia Linnaeus, 1758. Genus Eubyja Hübner, 1825 is currently considered a junior synonym of Biston Leach, 1815. Junior subjective synonym (Holloway 1994: 7). Braccini Warren, 1894 (Braccinae). Type species (of type genus Bracca): Phalaena bajularia Clerck, 1764. Junior subjective synonym (Holloway 1994: 167). Erannini Tutt, 1896 (Eranniinae). Type species (of type genus Erannis): Phalaena defoliaria Clerck, 1759. Junior subjective synonym (for instance Holloway 1994: 7), synonymy is supported by molecular results also (Sihvonen et al. 2011; Murillo-Ramos et al. 2019). Melanchroiini Hulst, 1896 (Melanchroiinae). Type species (of type genus Melanchroia): Phalaena chephise Stoll, 1782. The type genus Melanchroia Hübner, 1819 was included in the Braccinae [Braccini] by Warren (1905), and in the Boarmiini by Hodges et al. (1983) and Pitkin (2002). Dalimini Wehrli, 1940 (Daliminae). Type species (of type genus Dalima): Dalima apicata Moore, 1868. Genus Dalima Moore, 1868 was included in the wide concept of Boarmiini by Hollo­ way (1994) but without a tribe synonym. Tribe-level synonymy was proposed in Holloway (1996: 150). Phaseliini Wehrli, 1941 (Phaseliinae). Type species (of type genus Phaselia): Synopsia phaeoleucaria Lederer, 1855, which is a junior subjective synonym of Phaselia serrularia (Eversmann, 1847). Suggested to fall into the wide concept of Boarmiini (Forum Herbulot 2016), supported by molecular data also (Sihvonen et al. 2011; Murillo-Ramos et al. 2019). Melanolophini Forbes, 1948 (Melanolophini). Type species (of type genus Melanolopha): Tephrosia canadaria Guenée, 1858. Tentative junior subjective synonym according to Holloway (1994: 8), treated as a subgroup of the Boarmiini by Pitkin (2002: 131), supported by molecular data also (Sihvonen et al. 2011; Murillo-Ramos et al. 2019). Bupalini Herbulot, 1963 (Bupalini). Type species (of type genus Bupalus): Phalaena piniaria Linnaeus, 1758. Junior subjective synonym (Holloway 1994: 6; 167), synonymy is supported by molecular results also (Sihvonen et al. 2011; Murillo-Ramos et al. 2019). ‡ Milioniini Inoue, 1992 (Milioniini). First mentioned by Inoue (1992) but without description, therefore nomenclaturially unavailable. Junior subjective synonym (Holloway 1994: 8, 302). ‡ Obeidiini Inoue, 1992 (Obeidiini). First mentioned by Inoue (1992) but without description, therefore nomenclaturially unavailable. Junior subjective synonym (Holloway 1994: 8, 302). Unavailable names (incorrect subsequent spellings): Phaselini: Viidalepp (1996); Boarmini: ­Karisch (2005).

The Boarmiini are the most species-rich tribe in the Ennominae, in many areas representing almost half of the subfamily’s fauna. The tribe contains more than 150 genera and 5000 species globally, being particularly diverse in the Palaearctic and Oriental regions.

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The Boarmiini are found in all biogeographical areas, except the Antarctica, but are poorly represented in the northernmost areas in the northern hemisphere. Broad concept of the Boarmiini (Holloway 1994, p. 167) is applied, including the Bistonini but not the Gnophini, as this approach is supported by the DNA phylogenies also (Wahlberg et al. 2010; Sihvonen et al. 2011; Jiang et al. 2017; Murillo-Ramos et al. 2019), making the Boarmiini monophyletic. Rindge (1975) revised the North American Bistonini, and Viidalepp et al. (2007) presented morphological support for the Bistonini being monophyletic, but the latter noted that this might make the Boarmiini sensu stricto paraphyletic. This has been demonstrated to be the case in the above-mentioned DNA studies. External characters and abdomen: Viidalepp et al. (2007) proposed the following potential synapomorphic characters for the Boarmiini (not unique): fovea usually present in male forewings (Text-fig. 139), male abdominal sternite A3 with a transverse comb of setae or hair lock and the male antennae, if pectinate, have the apical segments filiform. Forewing fovea is absent for instance in Biston Leach, 1850 and the related genera (Viidalepp et al. 2007; our observations) and absent or weakly developed in Melanolophia Hulst, 1896 and related genera (Pitkin 2002). Externally the Boarmiini are a variable group and clear defining characters in the facies have not been identified so far. Numerous species have mottled markings of white or cream and brown or grey, which camouflage the moths in their typical daytime resting places against the bark of tree trunks (Pitkin 2002). Biston and related genera are large, with robust abdomen, and some genera have stout setae or even sclerotized structures on abdominal tergites (see Rindge 1975; Nupponen & Sihvonen 2013 for illustrations). At the other end of variation, numerous species are delicately built. Females of several winter flying and montane species have brachypterous wings. Male genitalia: Viidalepp et al. (2007) proposed the following potential synapomorphic characters for the Boarmiini: Uncus often triangular and gnathos usually well sclerotized, provided with a strong, flat medial cochlear. Other characteristic features include (our observations): Cucullus of valva often setose and process of valva in medial position. Juxta often sclerotized plate with long distal (ventral) appendices, whereas basal (dorsal) appendices are absent. Vesica often with various structures, such as sclerotized scobination, plate, or distinct cornuti. Female genitalia: No definitive, synapomorphic characters have been identified. Biston and related genera with very long apophyses, ovipositor telescopic. Signum often stellate, with ‘mushroom form’ centre, being typical shape also for Ennominae (Holloway 1994). Immature stages: Viidalepp et al. (2007) proposed the following potential synapomorphic character for the Boarmiini pupae: cremaster with D1, SD1 and L1 reduced, D2 fused into a stem (Text-figs 141–142). Numerous larvae are twig mimics, resting in erect position. Biology: Numerous Bistonini are arboreal feeders as larvae but generalisations are difficult to make because the group includes both highly polyphagous species and lineages that are more specialised. Biston betularia has been recorded to feed on numerous deciduous trees and bushes, in addition to records also on Filipendula ulmaria, Trifolium, Lythrum and Artemisia (Mikkola et al. 1989). Some species such as C. lichenaria (Hufnagel, 1767) feed on lichens, including Usnea and Alectoria (A. jubatus (Thunberg, 1788)), whereas some lineages feed exclusively on conifers such as mainly New Guinean Milionia Walker, 1854 on Araucaria, Dacrydium and Podocarpus (Holloway 1994). In Biston and related genera the elongated ovipositor is adapted for placing eggs in deep crevices, e.g. in bark, in species where information is available (Holloway 1994). 255

Text-figs 139–142. Selected diagnostic characters (indicated) of the Boarmiini, which are found widely but not in all lineages in the tribe. The reduction of cremaster hooklets to one strong pair is found in the Macariini, Cassymini and Eutoeini also. See text for other diagnostic characters. Text-fig. 139. Forewing base with blister-like fovea, Hypomecis roboraria (Denis & Schiffer­ müller, 1775), Finland: Enonkoski, 8–10.vii.1996, slide Sihvonen 2089 (coll. P. Sihvonen). Text-fig. 140. Valva with process in medial position, Alcis repandata (Linnaeus, 1758), Finland: Savonlinna, 26.vi-1.vii.1995, slide Sihvonen 2150 (coll. P. Sihvonen). Text-fig. 141. Pupa cremaster with setae D2 fused (ventral view), Biston betularia (Linnaeus, 1758), Germany: Berlin, reared by B. Müller in 1985 (coll. ZSM). Text-fig. 142. Pupa cremaster with setae D2 fused (ventral view), Hypomecis roboraria (Denis & Schiffermüller, 1775), Germany: Guben, reared by B. Müller in 1988–1989 (coll. ZSM).

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Remarks: Beljaev (2016: 546) included in the Boarmiini also the Glaucinini Rindge, 1959, but without justification. The Glaucinini are found from central Texas to coastal southern California, to Wyoming and Washington in the USA (Rindge 1976). The adults and genitalia are illustrated in Rindge (1959; 1976), but the facies and structures do not readily associate with any other Eurasian Boarmiini taxa. Glaucinini have not been included in DNA based phylogenetic studies. For these reason we do not include the Glaucinini in the synonymic list of the Boarmiini. Jiang et al. (2017) analyzed phylogeny of the Boarmiini using molecular data. The analysis was in agreement with earlier studies (for instance Sihvonen et al. 2011) and evidence was found to support monophyly of the “Boarmiine” lineage, including the Boarmiini and in a sister-taxon position the cluster of tribes including the Macariini, Cassymini, Eutoeini, Abraxini. Jiang et al. (2017) recovered the Gnophini as the sister-taxon to the “Boarmiine” lineage, but phylogenetic position of the Gnophini was not tested in depth in this study, due to very limited taxon sampling outside Boarmiini. In the analysis of Sihvonen et al. (2011) the Caberini were recovered as the sister to the “Boarmiine” lineage, whereas the Gnophini + “Boarmiine” were revealed as unrelated. The molecular results have been confirmed by muti-gene analysis of Murillo-Ramos et al. (2019).

Microbiston Staudinger, 1882 Microbiston Staudinger, 1882, Stettin. ent. Ztg. 43: 60. Type species: Microbiston tartaricus Staudinger, 1882. There are no genus synonyms.

Diversity and distribution: The genus Microbiston contains three species in the central Palaearctic region. External characters and abdomen: Medium-sized moths. Males with fully developed wings, wingspan 22−28 mm, thorax stout, furry, abdomen covered with shorter hairy scales. Females wingless, thorax and abdomen not furry. Male antennae bipectinate, female antennae filiform. Hindtibia of both sexes with two spurs. Male and female genitalia: European species M. lanaria see below. Biology and immature stages: Unknown.

81. Microbiston lanaria (Eversmann, 1852) Amphidasis lanaria Eversmann, 1852: Bull. Soc. Nat. Moscou 25 (1): 163 (Russia: Indersk region, Ural River). Syntype(s). Microbiston tartaricus Staudinger, 1882: Stettin. ent. Ztg. 43: 60 (Kazakhstan: Oz. Zaysan-­Lepsa). Syntypes 2♂.

External characters and abdomen: Male: Wingspan ♂ 22−25 mm. Forewing: Ground colour pale grey with dense dark speckling on forewings and weak speckling on hindwings. Postmedial, medial and antemedial lines on forewing present, on costa dilated to dark spots. Medial and postmedial lines often approaching at the inner margin. Subterminal line dark, distinctly shadowed whitish grey towards termen. Area between postmedial and subterminal line brownish grey filled. Hindwing with postmedial line often dotted, and with dark and wide subterminal band. All wings with blackish discal spots, on forewing sometimes indistinct. Fringes of ground colour. Terminal line black, dotted. Antennae bipectinate, pale brown, cilia large and dense. Labial palps with long hairy scales, laterally black. Frons, collar, tegula and thorax rough scaled, grey to brownish grey. Abdomen reddish brown, 257

without hairy scales, with double row of black dots. Female: Wingless, frons, collar, tegula and thorax smoothly scaled, grey to brownish grey. Hindtibia of both sexes with two spurs. Variation: Little. Discal spot of males on forewings sometimes indistinct. Subterminal band on the hindwings sometimes narrower. Male genitalia: Uncus triangular, lateral margins concave, with short double tip. Gnathos band-like, ventrally fused, forming a rounded, upturned structure. Dorsal arm of valva sclerotized, apex round, setose, costa slightly concave. Ventral arm of valva wide, apically concave with some short spines, marginal spines at apex somewhat larger, one with double tip. Juxta long, trapeziform. Saccus elongated, rounded. Aedeagus with two straight spines and a small ridge. From the base of the longer spine a third, smaller one arises. This spine is curved and widens the apical half of the aedeagus. Caecum rounded. 8th sternite with conspicuous wide and setose posterior margin. Female genitalia: Papillae anales elongated, slightly sclerotized, setose. Length of apophyses posteriores about 1.8 mm, ending spatulate. Apophyses anteriores shorter, about 1.0 mm. Lamella postvaginalis and lamella antevaginalis fused, forming sclerotized cap. Ductus bursae wide. Corpus bursae wide, membranous, its posterior part containing ridge-like structure. Anterior part of corpus bursae bent. Without signum. Distribution and abundance: Turanian. Very few records from Europe: In the Kiev museum three males from the 1970’s from Kapustin Yar situated about 100 km south-south-east of Volgograd (I. Kostjuk pers. comm.). Further records by the city of Uralsk, on the border between Europe and Asia. According to Zhuravlev (1910), specimens were found by the river Derkul, which is just on the European side of the border. – Rarely observed and poorly known, possibly less rare than indicated by the few records. – Outside Europe the species occurs in western and eastern Kazakhstan (Gorbunov 2011), Uzbekistan and northern Tadjikistan (Viidalepp 1996).

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Phenology: Univoltine. Scarce records from mid- and late March, probably before all snow has melted. A single specimen from western Kazakhstan recorded in late April 2012 where spring temperatures appeared very late (K. Nupponen pers. comm.). Biology: Unknown. Larva perhaps feeding on Salix on riverside meadows (K. Nupponen pers. comm.). Habitat: Halophilous. Found in salty steppe habitats with Artemisia absinthium (Gorbunov, 2011). Recorded on the edge between a large sand dune area and riverside wetland with plenty of Salix bushes, indicating that riverside meadows may be the true habitat for this species (K. Nupponen pers. comm.). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:ABV5725 (n=1 from northernmost Kyrgyzstan). Nearest species: M. phaeothorax Wehrli, 1941 from western Iran (0.9%; conspecific?), M. turanicus ­Staudinger, 1900 from Turkmenistan (3.5%), North American Phigalia titea (Cramer, 1780) (7.0%). Remarks: Close relationship to genus Apocheima was postulated by Leraut (2009) downgrading Microbiston from genus rank to synonymy. DNA barcode data does not support this. Requiring more in-depth analysis.

Apocheima Hübner, 1825 Apocheima Hübner, [1825] 1816, Verz. bekannter Schmett.: 319. Type species: Geometra hispi­ daria Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). ‡ Amphidasis Stephens, 1829, Nom. Br. Insects: Type species: Geometra hispidaria Denis & Schiffermüller, 1775. Junior homonym of Amphidasis Treitschke, 1825 (Geometridae, ­Ennominae), and junior objective synonym of Apocheima Hübner, [1825] 1816.

Diversity and distribution: Two species in the Palaearctic region, one in Europe. External characters and abdomen: Males with fully developed wings, female wingless. Thorax and abdomen stout, covered with hairy scales. Ground colour brownish to grey with dark speckling. Transverse lines black, partly jagged. Male antennae bipectinate, pale reddish brown, female antennae filiform. Male and female genitalia of European species, see A. hispidaria. Remarks: Apocheima and Phigalia are sister taxa, related to Hypomecis and Ematurga in several multi-gene analyses (Wahlberg et al. 2010; Jang et al. 2017; Murillo-Ramos et al. 2019).

82. Apocheima hispidaria (Denis & Schiffermüller, 1775) Geometra hispidaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 99 (Austria: Vienna district). Syntype(s). Phalaena ursularia Donovan, 1808: The natural history of British insects 13: 31, pl. 447 (England?). Syntype(s) ♂♀. Nyssia tauaria Newman, 1833: Ent. Mag. 1: 413 (England: Leominster). Syntype(s). Nyssia hispidaria f. cottei Oberthür, 1913: Études Lépid. comp. 7: 245, pl. 62, fig. 1582 (France: Digne). Syntype(s). Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on clinal transitions of characters and DNA barcodes, and absence of constant and significant differential features.

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Apocheima hispidaria orientis Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.):418, pl. 34: f (Turkey: Malatya). Syntype(s) 4♂. Valid at subspecific rank (Scoble 1999), here confirmed. Apocheima hispidaria popovi Vojnits, 1972: in Vojnits, Mészáros & Popov, Acta zool. hung. 18 (1–2): 233, fig. 1b (Bulgaria: Kostinbrod near Sofia). Holotype ♂. Valid at subspecific rank (Scoble 1999). Herewith downgraded from subspecies rank to synonymy. See remarks. Unavailable names (infrasubspecific): albomarginata: Lempke (1952) (f.); fumosa: Leraut (2009) (f.); fusca: Lempke (1952) (f.).

External characters and abdomen: Male: Wingspan ♂ 28–34 mm. Forewing: Ground colour pale brownish to pale grey with darker speckling. Terminal area usually lighter, divided from the rest of the wing by the distinctly serrate subterminal line. Other transverse lines black, subcostally bent inwards. Postmedial line concave at centre, darker marked on veins. Hindwing pale grey, speckled, postmedial line thick, grey, sometimes indistinct near anal margin. Terminal line indistinct, fringes more or less chequered. Antennae bipectinate, pale reddish brown, cilia large and dense. Frons, collar, tegula and thorax densely covered with hairy scales, dark grey to brown. Abdomen reddish brown, hairy, first segment usually with two black dots. Female wingless, antennae filiform, flagellomeres over half of antennal width with some long setae (length about 3 times of width of flagellum). Frons, collar, tegula and thorax as in males, but hairy scales shorter. Abdomen reddish brown, less hairy than in the male, with double row of black dots, anal segment silver-grey, with spinules on tergites. Hindtibia of both sexes with two apical spurs. Variation: In males the degree of dark speckling as well as the ground colour are variable, i.e. specimens are darker or paler. Melanic forms with uniform blackish grey ground colour and more black veins are known from some localities. Populations from south-eastern France (Alpes-de-Haute-Provence) described as larger, wings more silvery grey (‘f. cottei’). Subsp. orientis from Dagestan, Transcaucasia and Turkey is more dark grey scattered, postmedial line less concave, fringes more chequered (Wehrli 1939–1954). Male genitalia: Uncus triangular, apically bent, rounded. Gnathos forming a ventrally fused ring with upturned triangular apex. Both sclerotized. Costa of valvae wide, sclerotized, apex round, with group of small setae. Sacculus small at base, then wider and bent inwards, forming a median process with 4 to 8 spines at the tip. Number of spines can be different on the right and left valva as well as on valvae of different specimens. Saccus round. Juxta a wide plate with retuse tip. Aedeagus slender, subapically slightly constricted, apex and caecum tapered, without cornutus. Female genitalia: Papillae anales oval, membranous, soft, slightly setose. Apophyses posteriores very long (on average 4.6 mm), apophyses anteriores shorter (on average 2.0 mm), both spatulate at tip. Ductus bursae narrow, hardly sclerotized. Corpus bursae small, membranous, cuticula very thin, slightly striated. Signum absent. Lamella antevaginalis alate, with two small tapered ‘wings’. Lamella postvaginalis absent. Distribution and abundance: European. In Europe from the Urals to Great Britain and northern Portugal and from southern Scandinavia to the Mediterranean. – Rather common, sometimes occurring in numbers. – Outside Europe in the Caucasus region and in Turkey (subsp. orientis). Phenology: Univoltine: In southern Scandinavia from mid-March to mid-April, in south-­ western Germany from mid- or late February to early April (Ebert 2003) and in Italy and 260

Spain from February to April (Flamigni et al. 2007; Redondo et al. 2009). Larvae hatching from April in the south and from May in the north, until June or July. Hibernation as pupa. Males active at night, attracted to light. After emerging, females quickly crawl up the tree trunks, thus seen only occasionally. Biology: Larva oligophagous. Found on Quercus, Salix aurita, Carpinus betulus, Malus domestica, Prunus avium, P. spinosa (Ebert 2003). Recorded on Populus tremula, Castanea, Fagus, Ulmus (Flamigni et al. 2007). At least in northern Europe Quercus appear to be the main food-plant. Pupation in the ground (T. Drechsel pers. comm.) or in fine detritus on the ground (Z. Las˘tu˚ vka pers. comm.). Habitat: Silvicolous. In northern Europe preferring forests with Quercus or places with Quercus scrub, elsewhere in deciduous woodland, gardens and parks. In southern Scandinavia only at low altitudes, in south-western Germany from sea-level up to 800 m (Ebert 2003), In Sicily up to 1300 m (Flamigni et al. 2007) and in Spain up to 1600 m (Redondo et al. 2009). Similar species: Female of Phigalia pilosaria (Denis & Schiffermüller, 1775) with wings reduced to about one mm, antennae without setae, abdomen grey with only a slight tinge of reddish brown, legs scattered blackish and whitish. Genetic data: BIN: BOLD:AAD6044 (n=25 from United Kingdom, Netherlands, Germany, Czech Republic, France, Italy including Sardinia, Slovenia, Croatia, Bulgaria, Turkey). Genetically slightly heterogeneous, maximum variation 2.1%. Nearest species: North ­American Phigalia titea (Cramer, 1780) (6.5%), Phigalia strigataria (Minot, 1869) (6.7%), and central Asian Apocheima cinerarius (Erschoff, 1874) (6.9%). Nearest species in Europe Phigalia pedaria (7.8%).

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Remarks: Vojnits, 1972 described the subsp. popovi mainly because of “four thicker and robuster” spines on the median process of male valves. However, our analyses show high variation in number and size of the spines, different number can be found even on both valvae of same specimen. Four spines being within the normal variation of this species. Even the ‘greyish brown ground colour’ is not suitable for separating popovi from ‘brown A. hispidaria’. Therefore we downgrade the taxon popovi Voinits, 1972 from subspecies rank of A. hispidaria (Vojnits 1972; Scoble 1999) to synonymy based on the absence of diagnostic features.

Phigalia Duponchel, 1829 Phigalia Duponchel, 1829, Hist. nat. Lépid. Papillons Fr. 7 (2): 106, 296. Type species: Geometra pilosaria Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Coniodes Hulst, 1896, Trans. Am. ent. Soc. 23: 321, 353. Type species: Boarmia plumogeraria Hulst, 1888. Metriocera Djakonov, 1949, Ent. Obozr. 30: 377. Type species: Phigalia sinuosaria Leech, 1897. Pterotocera Staudinger, 1882, Stettin. ent. Ztg. 43: 59. Type species: Pterotocera declinata Staudinger, 1882. Rhaphidodemas Hulst, 1896, Trans. Am. ent. Soc. 23: 362. Type species: Phalaena titea Stoll, 1780.

Diversity and distribution: 14 species in the Palaearctic and Nearctic regions. Japanese species revised by Nakajima (1998). External characters and abdomen: Males medium-sized, wingspan 36–54 mm. Female brachypterous. Males of all species with similar pattern, forewing ground colour whitish to pale grey with dark mottling. Hindwing less speckled, lighter. Proboscis short, weak. Labial palpi very small. Head and thorax with hairy scales. Abdominal tergites covered extensively with spine-like setae. Male antennae bipectinate. Female with wings reduced to pads (brachypterous), antennae filiform. Forewing venation with R1 and R2 on common stalk, rarely both anastomosing. Hindwing with long discal cell, M2 present but atrophied (Prout 1915). Male genitalia: Uncus triangular with double tip. Gnathos forming a ventrally fused ring. Valva wider at base, apex rounded, subapically setose. Centrally an oval projection with a group of small spines. Saccus rounded with or without anterior projection. Juxta a triangular plate, lateral margins concave, apex with double tip. Aedeagus tapered at apex. Female genitalia: Papillae anales oval to elongate, membranous, slightly setose. Apophyses posteriores very long, apophyses anteriores shorter, ending spatulate. Ductus bursae as long as wide, sclerotized. Corpus bursae membranous, slightly striated, usually with signum; the latter variable in shape and size, sometimes absent. Shape of signum variable between females of the same species. See Nakajima (1998) and McGuffin (1977). Remarks: Apocheima and Phigalia are sister taxa, related to Hypomecis and Ematurga in several multi-gene analyses (Wahlberg et al. 2010; Jang et al. 2017; Murillo-Ramos et al. 2019).

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83. Phigalia pilosaria (Denis & Schiffermüller, 1775) Geometra pilosaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 100 (Austria: Vienna district). Syntype(s). Phalaena pedaria Fabricius, 1787: Mantissa Insect. 2: 191. Syntype(s). Phalaena Geometra hyemaria Borkhausen, 1794: Natur. eur. Schmett. 5: 193 (Europe). Syntype(s). Amphidasis extinctaria Standfuss, 1847: Stettin. ent. Ztg. 8: 62 (Poland: Szklarska Pore˛ba (Schreiberhau)). Syntype(s) ♂. Phigalia pedaria meridionalis Constantini, 1916: Atti Soc. Nat. Modena (5) 3: 17 (Italy). Syntype(s). Validated at subspecific rank by Scoble (1999), downgraded here from subspecific rank to synonymy (syn. n.), based on genetic homogeneity and absence of differential features, falling within intraspecific variation. Phigalia pilosaria prostae Leraut, 1996: Alexanor 19 (3): 148 (France: Hautes-Alpes, Guillestre). Holotype ♂ (Muséum d’Histoire Naturelle de Dijon). Validated at subspecific rank by Scoble (1999), downgraded here from subspecific rank to synonymy (syn. n.), based on absence of differential features, falling within intraspecific variation. Unavailable names (infrasubspecific): clausa: Lempke (1970) (ab.); destrigaria: Lempke (1952) (ab.); extinctaria: Standfuss (1947) (ab.); fasciaria: Richardson (1952) (ab.); flavescens: Leraut (2009) (f.); grisescens: Leraut (2009) (f.); lichenea: Leraut (2009) (f.); monacharia: Staudinger (1901) (ab.); nervosa: Lempke (1970) (ab.); obscurata: Schawerda (1919) (ab.); pallescens: Leraut (2009) (f.); subnigraria: Uffeln (1914) (ab.); tangens: Lempke (1952) (ab.); uniformata: Lempke (1952) (ab.).

External characters and abdomen: Male broad-winged, wingspan 36–45 mm. Ground colour pale grey. Forewing with dark grey or dark brownish speckling. Transverse lines black, often faint, jagged, strongest on costa and inner margin. Postmedial line and medial lines approximated between CuA2 and A. Hindwing less speckled, therefore lighter. Post­medial line right- or obtuse-angled before reaching the anal margin. Subterminal line more straight, usually diffuse. Terminal line of both wings black, interrupted by veins. Underside of forewing brownish grey, terminal area sometimes paler; hindwing pale grey. Fringes grey, divided by a line parallel to termen. Discal spots present on all wing upper- and undersides. Antennae bipectinate to tip, dark brown, base of flagellomeres usually white, cilia vitreous and dense. Frons usually white, brown in the middle, vertex white. Collar, tegula and thorax densely covered with hairy scales, dark grey to brown. Thorax posterodorsally with two white blotches. Abdomen reddish brown, weakly covered with spine-like setae. Two black dots on each segment form parallel lines. Female brachypterous, wing length hardly more than one millimeter. Wings covered with whitish and brown scales. Antennae filiform, brown with white blotches. Frons half brown, half white. Collar, tegula, thorax and abdomen grey, abdomen with more or less reddish brown scales. Legs chequered grey and white. Hindlegs of both sexes with 2+2 spurs. Variation: Extension of speckling, distinctness of lines and contrast are variable. The pale grey ground colour can have a greenish tinge. A dark olive, reddish beige colouration has been described. The broad range of variation resulted in a relatively big number of names for aberrations. Melanic forms can be found in some regions. They are unicolorous blackish in fresh material or brownish in older collection material with darker veins (“monacharia Staudinger, 1901”). Intermediate forms are known. Both taxa meridionalis and prostae described as being on average larger, the former ‘with distinct discal spots and lines’, the latter ‘with more ash grey hue’, both characters falling within the normal variation and/or with clinal transitions towards neighbouring populations, genetically homogeneous all over the European distribution area (see below under Genetic data). 263

Male genitalia: Uncus triangular, weakly setose, apex with sidewards directed double tip. Gnathos forming a ventrally fused ring with upturned apical part. Both sclerotized. Valva wider at base, apex rounded, subapically setose. Costa and sacculus sclerotized. Centrally an oval projection with a group of small spines of variable number. Saccus rounded with posterior projection. Juxta developed as a triangular plate, lateral margins concave, apex with double tip. Aedeagus short, thick, apex and caecum tapered. Without cornutus. Female genitalia: Papillae anales oval, membranous, soft, slightly setose. Apophyses posteriores very long (on average 2.9 mm), apophyses anteriores shorter (on average 1.8 mm), both ending spatulate. Lamella antevaginalis with two sclerotized oval ‘wings’. Lamella postvaginalis developed as a sclerite with straight lateral margins, base narrower, apex flat, slightly concave. Ductus bursae as long as wide, sclerotized. Corpus bursae membranous, slightly striated, without signum. Distribution and abundance: European-West Asiatic. In Europe from the Urals to Great Britain, Ireland and Portugal and from southern Scandinavia to the Mediterranean. – Relatively common, now and then found in numbers. – Outside Europe in Georgia, Armenia, Azerbaijan and Turkey.

Phenology: Univoltine: In southern Finland from late March to late April when there may still be small patches of snow on the ground (P. Sihvonen own data), in Denmark from mid-February to late March, in south-western Germany from early February to late March (Ebert 2003) and in Italy and Spain in February and March (Flamigni et al. 2007; Redondo et al. 2009). Larval time in southern central Europe from late March to April, in Finland in June and July (Seppänen 1970). Overwintering as pupa. Males active at night, attracted to light. Females rarely seen on tree trunks or branches. Apparently emerging during daytime, then climbing on tree trunks in the evening, remaining rather low on the trunks, up to two 264

metres, until copulating. After that climbing further up the tree. Unmated females can be found at any time of the day on tree trunks. Females emerge within very short periods (approximately one week), sometimes several females can be found in one afternoon. Suitable places for facilitated detection of females are large trees surrounded by grassland where they have no other choice than to climb on the trunks (P. Sihvonen own data). Biology: Larva polyphagous. Found on Populus tremula, P. nigra, Salix viminalis, S. caprea, Carpinus betulus, Corylus avellana, Fagus sylvatica, Castanea sativa, Quercus robur, Crataegus, Pyrus communis, Malus domestica, Sorbus aucuparia, Rosa canina, Prunus dulcis, P. spinosa, Vaccinium myrtillus, Fraxinus excelsior and Artemisia campestris (Ebert 2003). Found also on Acer (north-western Italy: King & González-Estébanez 2015). Recorded on Betula (Skou 1984), Alnus, Ulmus, Prunus armeniaca, P. avius, P. domestica, Acer and Tilia and reared on Picea (Flamigni et al. 2007). Pupation in the ground or in fine detritus on the ground (Z. Las˘tu˚ vka pers. comm.). Habitat: Silvicolous. Due to the large number of food-plants occurring in a large variety of habitats with deciduous bushes and trees and in cultivated areas such as gardens and parks. In northern Europe at low altitudes, in south-western Germany up to 1200 m (Ebert 2003), in Italy up to 1650 m (Flamigni et al. 2007) and in Spain up to 1500 m (Redondo et al. 2009). Similar species: Female of Apocheima hispidaria differs by being completely wingless, antenna half of length with long hairy setae on flagellomeres, abdomen reddish brown, legs brownish grey with only a small number of whitish scales, appearing unicolorous. Genetic data: BIN: BOLD:AAD0877 (n=34 from Finland, United Kingdom, Germany, Austria, Switzerland, Czech Republic, France, Italy, Slovenia). Genetically homogeneous. Nearest species: P. titea (Cramer, 1938) from North America (4.8%).

Hypomecis Hübner, 1821 Hypomecis Hübner, 1821, Index exot. Lepid.: [7]. Type species: Cymatophora umbrosaria H ­ übner, 1813. Boarmia Treitschke, 1825 [October 18], in Ochsenheimer, Schmett. Eur. 5 (2): 433. Type species: Geometra roboraria Denis & Schiffermüller, 1775. Dryocoetis Hübner, [1825] 1816, Verz. bekannter Schmett. (20): 316. Type species: Geometra roboraria Denis & Schiffermüller, 1775. Serraca Moore, [1887] 1884–7, Lepid. Ceylon 3: 416. Type species: Boarmia transcissa Walker, 1860. Narapa Moore, [1887] 1884–7, Lepid. Ceylon 3: 410. Type species: Boarmia adamata Felder & Rogenhofer, 1875. Pseudangerona Moore, [1887] 1884–7, Lepid. Ceylon 3: 413. Type species: Boarmia separata Walker, 1860. ‡ Alcippe Gumppenberg, 1887, Nova Acta Acad. Caesar. Leop. Carol. 49: 335 (key). Type species: Macaria castigataria Bremer, 1864. [Junior homonym of Alcippe Blyth, 1844 (Aves)]. Astacuda Moore, 1888, Descr. new Indian lepid. Insects Colln late Mr W. S. Atkinson: 243. Type species: Astacuda cineracea Moore, 1888. Maidana Swinhoe, 1900, Cat. east. and Aust. Lepid. Heterocera Colln Oxf. Univ. Mus. 2: 280. Type species: Macaria tetragonata Walker, 1863. Pseudoboarmia McDunnough, 1920, Bull. Dep. Agric. Can. ent. Brch 18: 21, pl. 9, fig. 7, pl. 10, fig. 3. Type species: Cymatophora umbrosaria Hübner, 1813. Erobatodes Wehrli, 1943, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 521. Type species: Boarmia eosaria Walker, 1863.

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Diversity and distribution: Genus Hypomecis with 138 plus 24 tentatively assigned species (Scoble 1999). With further research, deletions and additions to the list are likely to be necessary. Distributed in all parts of the world, with focus on the Oriental Region both in lowlands and montane habitats. External characters and abdomen: Large or medium-sized moths, wingspan 22−60 mm. Ground colour usually whitish, grey or brownish, irrorated with darker grey and black. Antennae partially bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs, in male with pencil. Male genitalia: Valvae usually with two bands with short setae at centre of valva, one subcostally and the other as an extension of sacculus. Tegumen dorsally with a pair of slender, curved processes, in some species reduced or absent. Juxta broad at base, then sharply constricted. Aedeagus short, terminating in one or two spined processes. Vesica tubular, more rarely with a lobe or patch with short spines (Holloway 1994; Sato 1984). Female genitalia: Ductus bursae short, rather narrow. Corpus bursae elongate. Anterior part of corpus bursae with signum of two sclerotized teeth linked by a shorter bar, some species without signum (Holloway 1994). Remarks: Several multi-gene analyses revealing close relationship between Hypomecis and Ematurga, both falling in the same cluster as Phigalia and Apocheima (Jiang et al. 2017; Murillo-Ramos et al. 2019).

84. Hypomecis roboraria (Denis & Schiffermüller, 1775) Geometra roboraria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s). Geometra consobrinaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 29, fig. 152 (Europe). Syntype(s). Boarmia menetriesi Staudinger, 1871: in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 165 (Russia, Siberia or.). Syntype(s). Valid at subspecific rank (Scoble 1999). Boarmia displicens Butler, 1878: Ann. Mag. nat. Hist. (5) 1: 395 (Japan: Yokohama). Syntype(s) (NHMUK). Valid at subspecific rank (Scoble 1999). Boarmia arguta Butler, 1879: Ann. Mag. nat. Hist. (5) 4: 372 (Japan: Yokohama). Syntype(s). Boarmia roboraria var. amurensis Staudinger, 1892: Dt. ent. Z. Iris 5: 379 (Kentei, Amur). Syntypes including 1♂1♀. Boarmia roboraria var. kidsicola Gumppenberg, 1892: Nova Acta Acad. Caesar. Leop. Carol. 58 (4): 350 (Russia: Amur). Syntype(s). Alcis molata Warren, 1901: Novit. zool. 8: 32 (Japan: Yokohama). Holotype ♂ (NHMUK). Alcis amplaria Wileman, 1911: Entomologist 44: 345 (Taiwan: Kanshirei). Holotype ♀ (NHMUK). Valid at subspecific rank (Scoble 1999). Boarmia roboraria demonstrata Prout, 1917: Novit. zool. 24: 315 (China: Tibet, Vrianatong). Syntypes 2♂ (NHMUK). Valid at subspecific rank (Scoble 1999). Boarmia isabellae Fernández, 1931: Eos, Madr. 7 (2): 219, pl. 2, fig. 15 (Spain: Salamanca, Béjar). Syntype(s) ♂ (ZFMK). Boarmia danieli Wehrli, 1936: Mitt. münch. Ent. Ges. 22 (2): 34, pl. 1 fig. 4 (Hungary). Syntype(s). Boarmia roboraria nipponica Inoue, 1955: Kontyu 22 (3/4): 75, pl. 7, fig. 24 (Japan: Nagano Prefecture, Nakabusa-onsen, 1400 m). Holotype ♂ (NHMUK). Unavailable names (infrasubspecific): infuscata: Staudinger (1871) (ab.); melaina: Schulze (1913) (ab.); obsoleta: Leraut (2009) (f.); variegata: Leraut (2009) (f.).

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External characters and abdomen: Wingspan ♂ 39−48 mm, ♀ 44−59 mm (largest geometrid moth in Europe). Ground colour of all wings whitish grey, irrorated with dark grey or brownish. Antemedial line curved, near costa bent at right angle, medial line indistinct. Postmedial line serrate, approaching medial line near inner margin without touching it, here the space between both lines often filled black. Wavy line whitish, usually clearly visible. Terminal line dotted. Fringes concolorous with ground colour, uniform or slightly chequered. Discal spots without pale centre. Frons, collar and patagia whitish grey, patagia posteriorly black. Underside of wings yellowish grey. Discal spot, postmedial line and forewing terminal area brownish. Forewing apex with pale blotch (see Text-figs 143–144). Antennae partially bipectinate to about 5/6 of length in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Abdomen concolorous with ground colour and with double row of black dots dorsally, first segment anteriorly silver-grey. Variation: Ground colour and degree of irroration on all wings varying in a wide range. Black and intermediate specimens with conspicuous whitish wavy line on fore and hindwings common mainly in Central Europe, locally being the dominant form. Totally black specimens reported by Malkiewicz (2012). Male genitalia: Uncus tapered, beak-shaped. Gnathos arcuate, arms apically fused to a band-shaped, lobed structure. Valva wide, costa sclerotized, setose, subcostally with a short, rounded spined process. Sacculus with a band-shaped spined extension, shorter and thicker than in H. punctinalis. Saccus rounded. Aedeagus with four ‘cornuti’: a shorter and a longer serrate band, a band spined over half of the length and a nearly triangular, slightly sclerotized structure. Female genitalia: Papillae anales soft, setose. Apophyses posteriores very long, thickened at centre, spatulate at tip. Apophyses anteriores long, thickened at 1/3 from tergum, spatulate at tip. Lamella antevaginalis sclerotized, alate, lamella postvaginalis a circular plate. Ductus bursae wide, short. Corpus bursae elongate, posterior third sclerotized, striated, at centre slightly constricted. Anterior part a membranous concentrically striated sac with signum inside. Signum with two sclerotized teeth linked by a shorter bar. Distribution and abundance: Eurasiatic. In Europe from the Urals to Denmark, southern England, France and Portugal and from central Fennoscandia to northern Spain and Portugal with an isolated occurrence in the Algarve (M. Corley pers. comm.), southern France including Corsica, central Italy and the Balkan peninsula excluding most of Greece. – Rather local and not common, usually found in limited numbers. – Outside Europe in Turkey, Georgia and Armenia (subsp. roboraria) from northern Kazakhstan through southern Siberia and Transbaikalia to Mongolia and northern China to southern Yakutia (subsp. ­isabellaria), and in Far East Russia, southern Kuriles and Japan (subsp. menetriesi and displicens). Phenology: Uni- or bivoltine. In southern Fennoscandia from mid-July to late July, in south-western Germany from mid-May to early August (Ebert 2003), in Italy and in the Iberian peninsula in warmer places in two generations from May to June and from August to September (Flamigni et al. 2016; Redondo et al. 2009). Caterpillars from July to May, hibernating. Adults active at night, attracted to light. Biology: Larva oligophagous. Found on Quercus and Fagus (Ebert 2003); Quercus probably being the preferred food-plant. Recorded also on Ulmus, Betula and Malus (Bergmann 1955). Pupation in a silken cocoon in the ground (Porter 2010).

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Habitat: Silvicolous. Forests with a good population of older Quercus trees, both on dry and more humid soils. In northern Europe from sea-level, in south-western Germany (Ebert 2003) and in the Alps up to 1000 m (Kerschbaum & Pöll 2010), in Italy rarely up to 1200 m (Flamigni et al. 2016). Similar species: H. punctinalis is on average smaller, and its apical area of forewing underside is not different from the rest of the wing (see see Text-figs 143–144). Discal spots are crescent or oval-shaped, on hindwings with pale centre. Male genitalia dorsally with a pair of curved processes on the tegumen. Genetic data: BIN: BOLD:AAC9905 (n=38, from Finland, Sweden, Germany, Austria, Czech Republic, France, Italy, Slovenia, Hungary (four of them pre-identified as H. danieli), Russia, Georgia, Kazakhstan, China, Mongolia). Genetically very homogeneous. Two barcoded specimens from north-eastern China and North Korea at distances of 2.8%, probably not conspecific. Nearest species: unidentified Hypomecis from Bhutan (6.1%), H. philippinensis Sato, 1996 from the Philippines (7.3%). Remarks: We consider H. danieli (Wehrli, 1936) as conspecific with H. roboraria (see genetic data). It was described from Hungary, compared only with H. punctinalis, which is definitely a different species. In a recent Hungarian publication (Gyulai et al. 2010) the authors write that most entomologists in Hungary consider H. danieli as a local ‘variation’ of H. roboraria. Gomboc (pers. comm.) found differences in biology (food-plant Betula) and flight time in Slovenia within the roboraria/danieli populations. This may point to partially univoltine and bivoltine ‘races’ of the same species.

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85. Hypomecis punctinalis (Scopoli, 1763) Phalaena punctinalis Scopoli, 1763: Ent. Carniolica: 217, fig. 537 (Italy: Carnia). Syntype(s). Phalaena urticaria Hufnagel, 1767: Berlin Mag. 4 (5): 508 (Germany: Berlin region). Syntype(s). Phalaena turcaria Fabricius, 1775: Syst. Ent.: 624 (Germany). Syntype(s). Phalaena Geometra griseonigra Goeze, 1781: Ent. Beytr. 3 (3): 426 (France: Paris). Syntype(s). Phalaena bandevillaea Fourcroy, 1785: Entomologia Parisiensis 2: 275 (France: Paris). Syntype(s). Phalaena consortaria Fabricius, 1787: Mantissa Insect. 2: 187 (Austria). Syntype(s) (lost). Boarmia conferenda Butler, 1878: Ann. Mag. nat. Hist. 5 (1): 395 (Japan: Yokohama). Syntype(s) (NHMUK). Valid at subspecific rank (Scoble 1999). Boarmia punctinalis var. grisearia Bastelberger, 1902: Dt. ent. Z. Iris 15: 220 (Russia: southern Urals). Syntype(s). Validated at subspecific rank by Scoble (1999), not mentioned by Leraut (2009), herewith downgraded to synonymy based on DNA barcode patterns, clinal character transitions and absence of constant and significant differential features. Boarmia consortaria var. marginata Herz, 1905: Ezheg. zool. Muz. 9: 372, pl. 1, fig. 6 (Russia: Siberia, S of Minussinsk). Holotype ♂. Valid at subspecific rank (Scoble 1999). Boarmia glos Prout, 1914: Ent. Mitt. 3 (9): 269 (Taiwan: Kosempo). Holotype ♂. Valid at subspecific rank (Scoble 1999). Boarmia punctinalis magyarica Wehrli, 1932: Mitt. münch. ent. Ges. 22 (2): 36, pl. 2, figs 2, 11 (Hungary). Holotype ♂ (NHMUK). Validated at subspecific rank (Scoble 1999), considered as f. by Leraut (2009), synonymy here confirmed. ­ huotsu). Boarmia Boarmia marginata submarginata Bryk, 1949: Ark. Zool. 41A (1): 205 (Korea: S Holotype ♂ (NHRS). Boarmia Boarmia conferenda referendaria Bryk, 1949: Ark. Zool. 41A (1): 205, pl. 6, fig. 14 (Korea: Myokosan). Holotype ♂ (NHRS). Valid at subspecific rank (Scoble 1999). Hypomecis punctinalis kerstinae Fischer & Lewandowski, 2004: Atalanta 35 (1/2): 127, pl. 10b (Cyprus: Prastio). Holotype ♂ (ZSM). Valid at subspecific rank. Unavailable names (infrasubspecific): bicolor: Lempke (1952) (f.); consobrinaria: Borkhausen (1790) (ab.); costamaculata: Leraut (2009) (f.); distincta: Lempke (1952) (f.); ­griseomarginata: Leraut (2009) (f.); grisescens: Leraut (2009) (f.); humperti: Humpert (1898) (ab.); inter­ media: Lempke (1952) (f.); mediomaculata: Leraut (2009) (f.); nigra: Warnecke (1938) (ab.); nigromarginata: Lempke (1952) (f.); obscura: Leraut (2009) (f.); obsoleta: Lempke (1952) (f.); variegata: Lempke (1952) (f.).

External characters and abdomen: Wingspan 37−44 mm, exceptionally up to 49 mm. Ground colour of all wings brownish grey to whitish grey, mottled with dark and light grey streaks. Antemedial line with right-angled bend as also the indistinct medial line. Post­medial line with small spots on veins, approaching medial line near inner margin without touching it. Wavy line, marginal line and fringes as in H. roboraria. Underside of forewings grey, apical area concolorous. All discal spots crescent or oval shaped, usually pale centred, especially on hindwings (see H. roboraria). Frons and collar whitish grey, patagia whitish grey, dorsally black. Antennae partially bipectinate to about 3/4 of length in male, filiform in female. Hindtibia and abdomen as in H. roboraria. Male with a sparse row of stout setae on third abdominal segment. Variation: Generally as in H. roboraria. Specimens from southern Urals (f. grisearia) very small, ground colour dark grey without whitish irroration, blackish pattern indistinct, underside without darker scales (Bastelberger 1902). Specimens from Hungary (f. magyarica) described as more contrasted than in nominotypical subspecies, medial lines deeper black, underside paler with stronger black irroration.

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The non European subsp. kerstinae constantly smaller (wingspan 34−40 mm), wing colour pale ash grey, wing pattern more diffuse. Antemedial, medial and postmedial lines grey, often less visible. Postmedial line on veins slightly thicker. Submarginal line on hindwings present. All discal spots weak. Subsp. kerstinae is less variable than the nominotypical subspecies. Male genitalia: Uncus tapered, beak-shaped. Top of tegumen with a pair of long narrow processes, ending in a hooked spine (lacking in H. roboraria). Gnathos rounded, lobed. ­Valva wide, apically becoming broader. Costa sclerotized, setose, with elongated spined process of one third of valva length. Sacculus with elongated harpe, at costa with more than 20 short spines of different length. Saccus rounded. Aedeagus stout, tapered; vesica with two spined bands, one shorter than the other. Vesica opens at approximately 45 degree a ­ ngle, membranous, wide at base with spinose band. Apex of vesica tapered, partly granular. Female genitalia: Papillae anales soft, setose, elongate. Apophyses posteriores very long, apo­physes anteriores shorter, both spatulate at tip. Lamella antevaginalis with two sclerotized ‘wings’, posterior half striated, lamella postvaginalis a triangular plate. Ductus bursae wide, short. Corpus bursae membranous, elongate, posterior third sclerotized, striated. Signum minute, pointed. Distribution and abundance: Eurasiatic. In Europe from southern Urals to Denmark and south-eastern England, and from southern Scandinavia to northern Spain and Portugal (including Algarve, M. Corley pers. comm.), southern France including Corsica, Italy including a single record in Sardinia, and the Balkan peninsula, excluding most of Greece. Erroneously recorded for Norway by Leraut (2009), in southern Spain only unverified literature records from Sierra Nevada (Redondo et al. 2009), awaiting confirmation. – Common or rather common, now and then found in numbers. – Outside Europe subsp. kerstinae described from Cyprus, across Turkey to Georgia, Armenia and Azerbaijan, nomino­typical subspecies in the east to northern Kazakhstan, subsp. marginata in southern Siberia, Dahu­ria, Far East Russia and Korea, subsp. conferenda in China, southern Kurile islands, ­Sakhalin and Japan.

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Text-figs 143–144. Selected diagnostic characters (indicated) of European Hypomecis species, wing underside. Text-fig. 143. H. punctinalis (Germany: Brandenburg, Oberhavel, 30.iii.1987 e.o.); Text-fig. 144. H. roboraria (Germany: Brandenburg, Spree-Neisse, Guben, 07.v.1989 e.o.) with different pattern of the apex.

Phenology: Uni or bivoltine. In Denmark from mid-May to mid-July, in south-western Germany from early May to late July (Ebert 2003) and in Italy at lower altitudes in two generations from late April to late June and from July to August or September, at higher altitudes only one generation (Flamigni et al. 2016). Caterpillars from May or June to September or October, depending on latitude and altitude. Overwintering as pupa. Adults active at night, readily attracted to light. Biology: Larva polyphagous. Found on Larix decidua, Salix, Carpinus betulus, Alnus glutinosa, Quercus, Berberis vulgaris, Sorbus aucuparia, Rubus idaeus, R. fruticosus, Prunus spinosa, Acer, Frangula alnus, Cornus sanguinea and Vaccinium myrtillus (Ebert 2003). Pupation on or in the soil surface. Habitat: Silvicolous. In a wide variety of habitat types with deciduous trees and bushes including forests with Larix. From sea-level up to 1600 m in the Alps and Italy. Similar species: Ascotis selenaria with male antennae ciliate-fasciculate, larger thorax and more conspicuous and larger discal spots. H. roboraria on average larger, apical area of forewing underside with bright blotch (see Text-figs 143–144), discal spots elongate, without pale centre. Male genitalia dorsally without pair of curved processes on tegumen. In females apophyses anteriores and apophyses posteriores with widening at half way. Genetic data: Species diverging into four deeply separated genetic clusters (BINs): BIN: BOLD:AAB1058 (n=66 from 15 countries from Finland to northernmost Italy and from United Kingdom to central and Far East China), genetically comparatively homogeneous, rarely sympatric with specimens of the other BINs; BIN: BOLD:AAB1059 (n=13 from northern (one) and southern Italy, Bulgaria, Cyprus, the last as “subsp. kerstinae”), genetically somewhat heterogeneous, maximum variation 1.6% (between populations of southern Italy and Cyprus); BIN: BOLD:ACA2461 (n=4 from Austria and Russia: Primorye), genetically homogeneous; BIN: BOLD:AAB1057 (n=1 from Israel). The four different BINs diverging from each other by 6.4–7.6%. Nearest species: three unidentified specimens, possibly 271

­ elonging to three different species from China (Hainan) and at distances of 3.8–4.4% from b BOLD:ACA2461. Requiring more research.

Ematurga Lederer, 1853 Ematurga Lederer, 1853, Verh. zool.-bot. Ver. Wien 3 (Abh.): 179, 227, 231. Type species: Phalaena atomaria Linnaeus, 1758. There are no genus synonyms.

Diversity and distribution: The genus Ematurga includes two sister species, one each in the Palaearctic and Nearctic regions. External characters, abdomen and genitalia: See species description below. Remarks: Several multi-gene analyses revealing close relationship between Hypomecis and Ematurga, both falling in the same cluster as Phigalia and Apocheima (Jiang et al. 2017; Murillo-Ramos et al. 2019).

86. Ematurga atomaria (Linnaeus, 1758) Phalaena (Geometra) atomaria Linnaeus, 1758: Syst. Nat. (Ed. 10) 1: 521 (Europe). Syntype(s) (LSL). Phalaena isoscelata Scopoli, 1763: Ent. Carniolica: 225, fig. 558 (Slowenja: Carnia). Syntype(s). According to Leraut (2009) valid at subspecific rank. Herewith downgraded to synonymy based on absence of constant and significant differential features, see under ‘Variation’. Phalaena pennata Scopoli, 1763: Ent. Carniolica: 228, fig. 569 (Slowenja: Carnia). Syntype(s). Phalaena aceraria Hufnagel, 1767: Berlin Mag. 4 (5): 520 (Germany: Berlin region). Syntype(s). Phalaena artemisiaria Herbst, 1782: in Fuessly, Arch. Insectengesch. 2: pl. 4, figs 1–5 (Europe). Syntype(s). Phalaena microcosma Fourcroy, 1785: Entomologia Parisiensis 2: 279 (France: Paris). Syntype(s). Phalaena picta Fourcroy, 1785: Entomologia Parisiensis 2: 275 (France: Paris). Syntype(s). Geometra glarearia Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae 1: pl. 25, fig. 131 (­Europe). Syntype(s). Fidonia obsoletaria Zetterstedt, 1839: Insecta Lapponica 4: 957 (Sweden). Syntype(s) ♂. Ematurga atomaria var. orientaria Staudinger, 1861: in Staudinger & Wocke, Cat. Lepid. Eur. und angrenzenden Länder 1: 74 (Europe). Syntype(s). Validated at subspecific rank (Scoble 1999), downgraded to synonomy of ‘subsp.’ isoscelata by Leraut (2009). Ematurga atomaria var. iliaria Alphéraky, 1883: Horae Soc. ent. ross. 17 (3/4): 181 (Russ. Armenia: Ili Valley). Syntypes ♂♀. Valid at subspecific rank (Scoble 1999). Bupalus atomarius Meyrick, 1895: A handbook of British Lepidoptera: 265. Emendation of atomaria L. Ematurga atomaria var. krassnojarscensis Fuchs, 1899: Jb. nassau. Ver. naturk. 52: 152 (Russia: Siberia, Krassnojarsk region). Syntype(s) ♂. Valid at subspecific rank (Scoble 1999). Ematurga atomaria var. transalpinaria Fringes, 1909: Societas ent. 24 (10): 77 (Italy: Gardasee, Gardone; Gargnano). Syntypes ♂♀. In Scoble (1999) as synonym of E. a. orientaria. Ematurga atomaria var. meinhardi Krulikovski, 1909: Ent. Obozr. 8 (3/4): 273 (Central Asia: Semiretshje). Syntype(s). Valid at subspecific rank (Scoble 1999). Ematurga atomaria f. alpicolaria Vorbrodt & Müller-Rutz, 1917: Mitt. schweiz. ent. Ges. 12 (9/10): 490 (Switzerland: Fusio). Syntype(s). Valid at subspecific rank (Scoble 1999). Herewith downgraded to synonymy, based on absence of constant and significant differential features, see under ‘Variation’. Ematurga atomaria minuta Heydemann, 1925: Int. ent. Z. 18 (44): 269 (Germany). Syntypes ♂♀. Valid at subspecific rank (Scoble 1999). Herewith downgraded to synonymy, based on absence of constant and significant differential features, see under ‘Variation’.

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Ematurga arenaria Candèze, 1926: Bull. Annls Soc. roy. ent. Belg. 66 (5): 252 (Belgium: Campine). Syntypes ♂♀. Valid at subspecific rank (Scoble 1999). Herewith downgraded to synonymy, based on absence of constant and significant differential features, see under ‘Variation’. Ematurga atomaria zetterstedtaria Heydemann, 1930: Int. ent. Z. 24 (12): 149, pl. 2, fig. 39. Original description with a damaged letter “d”, but clearly as “d” recognizable. In Scoble (1999) synonymized, here accepted. Ematurga atomaria ngana Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 649, pl. 52: b (China: south Kansu, Ngan-shan, Tschöng, 2000 m; south-east Kansu, San-tscha, 1800 m; Tschili Prov., Kalgan). Syntypes 5♂1♀. Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): bistrigaria: Heydemann (1930) (ab.); coloraria: Stauder (1920) (ab.); dentaria: Stauder (1920) (ab.); fallax: Stauder (1920) (ab.); fasciata: Heqvist (1950) (ab.); felicis: Krausse (1915) (ab.); gynaekoides: Heinrich (1923) (ab.); kindervateri: Schawerda (1929) (ab.); nigrescens: Lempke (1952) (f.); ochrearia: Rebel (1910) (ab.); ophthalmaria: Stauder (1920) (ab.); pseudoclathrata: Heydemann (1930) (ab.); pseudoglarearia: Stauder (1920) (ab.); tenuilineata: Lempke (1952) (f.); unicoloraria: Staudinger (1871) (ab.); unimarginata: Cornelsen (1923) (ab.); ustaria: Fuchs (1901) (ab.); virilis: Stauder (1915) (ab.).

External characters and abdomen: Wingspan 22–31 mm. Male: Ground colour yellow-ochre irrorated with brown. Transversal lines, veins and often terminal area darker brown. All transversal lines diffuse, curved, medial line thickened on veins. Wavy line reduced to row of dots of ground colour, discal spot larger, often paler. Female with ground colour white or yellowish white, irrorated with greyish brown. Medial lines and veins as in males, but greyish brown, creating contrasted appearance. Fringes of ground colour, chequered brown. Discal spots usually well developed, sometimes indistinct. Underside because of reduced darker parts more contrasted. Frons roughly scaled in male, smoothly scaled in female. ­Patagium, thorax and abdomen brownish grey. Antennae partially bipectinate (about four flagellomeres free of branches) in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Highly variable. In males of the nominotypical subspecies the ratio of brownish yellow and greyish brown parts varies considerably. Extreme forms are nearly unicolorous brownish yellow or greyish brown. In females the same ratio ranges between whitish and greyish brown. Females bred from whitish forms under warm room conditions often show a reddish brown ground colour as is typical for southern forms. At higher altitudes of the Alps with darker pattern and smaller (‘alpicolaria’). In north-western Europe smaller, wingspan ♂ 19–24.5 mm, ♀ 19–20 mm, male ground colour whitish to yellowish white, hindwing slightly yellower than forewing, female white (‘minuta’). In Mediterranian habitats with ground colour brownish orange, lines distinct, combined with brown lines in both sexes (‘transalpinaria’). In northern and north-eastern Europe similar to ‘minuta’ but larger (by 2–3 mm on average), male ground colour yellowish grey to dull ochreous, hindwing slightly yellower than forewing, discal spot on hindwing distinct, females white with brownish black lines (‘zetterstedtaria’ cf. Heydemann 1930). All forms are highly influenced by climatic conditions: Lower temperatures, higher altitudes and northern latitudes result in smaller, more sombre specimens with more white in males and females and darker transverse lines. Specimens from warm climates and lower altitudes are more yellowish orange in males and females. Intermediate forms are observed. Thus, many subspecies and aberrations of doubtful value have been described from Europe (see Heydemann 1930; Wehrli 1939–1954; Leraut 2009). Male genitalia: Uncus triangular, laterally concave, tip bent, slightly setose. Gnathos arms long, narrow, fused ventrally, forming large quadrate or trapezoid plate. Socii with scle 273

rotized small plates. Valva membranous, weakly setose, wide at base, apex narrower. Costa concave, setose subapically. Sacculus sclerotized with ridge. Central part of valva with narrow spinose harpe. Juxta sclerite of variable shape, often heart-shaped, anteriorly with small tip. Saccus round with ventrally bent projection. Aedeagus cylindrical, apex tapered with fan-shaped group of about ten cornuti of different length of spines at margin. Female genitalia: Papillae anales very long, slender, sclerotized. Apophyses posteriores very long, on average 4.4 mm, slightly bent, ending in small spatula. Apophyses anteriores shorter, on average 1.8 mm, ending in wide spatula. Lamella antevaginalis inconspicuous. Lamella postvaginalis a sclerotized, unevenly rounded plate. Corpus bursae anteriorly sacciform, posteriorly wider, sclerotized, with longitudinal folds. Signum small, inconspicuous, nearly membranous. Ductus bursae narrow, ostium bursae wider. Distribution and abundance: Eurasiatic. In Europe from the Urals to the British Isles and from northern Fennoscandia to the northern Iberian peninsula, southern France including Cor­sica, Italy and the Balkan peninsula including Greece. Occurrence in southern Iberian peninsula misleadingly suggested by Leraut (2009). – In northern Europe local, but usually found in numbers in its localities. In central and southern Europe more widespread, here less often in numbers. – Outside Europe widely distributed from Turkey, Caucasus and Transcausus through central Asia and Siberia to Mongolia, Far East Russia, Sakhalin and Kamchatka.

Phenology: Uni- or bivoltine. In Denmark from late April to late July, in south-western Germany from mid-April to late July (Ebert 2003). North of the Alps late specimens probably belong, at least to a certain degree, to a partial second generation. In the Alps two generations at low and one at higher altitudes, generally from early April to late August (Kerschbaum & Pöll 2010). In Italy bivoltine from April to June and from July to August; in one generation at higher altitudes (Flamigni et al. 2016). Caterpillars from May, June or 274

July to late September (Ebert 2003). Hibernation as pupa. Adults flying by day, very easily flushed up from vegetation during day time. In Denmark numerous males observed flying slowly around just above Calluna vulgaris vegetation in sunshine in the morning between 8 and 9 o’clock (P. Skou pers. observation). Adults occasionally attracted to light, but it is not clear if this is due to active flight or to disturbance. Biology: Larva polyphagous. In northern Europe above all on Calluna vulgaris, in south-western Germany also on Rumex acetosella, Prunus spinosa, Cytisus scoparius, Genista germanica, G. tinctoria, G. sagittalis, Medicago sativa, Lotus corniculatus, Hypericum perforatum, Helianthemum nummularium, Vaccinium myrtillus, Achillea millefolium and Artemisia vulgaris (Ebert 2003). Pupation in the ground. Habitat: In northern Europe mainly occurring in places with Calluna vulgaris such as heathland and heather moors, further to the south in a wide variety of open habitat types. In England rather common on chalk grassland (M. Corley pers. comm.). From sea-level up to about 2000 m. Similar species: Whitish females of E. atomaria somewhat similar to Macaria carbonaria (Clerck, 1759), but the latter considerably smaller and forewing medial and postmedial lines not fused on inner margin (Skou & Sihvonen 2015). Genetic data: BIN: BOLD:AAC1877 (n=56 from 15 countries from Finland to southern Italy, from United Kingdom and Portugal to Turkey, Georgia and central Asia). Genetically slightly heterogeneous, maximum variation 2.2%. A singleton from Bavaria more strongly diverging (minimum distance 2.1%), probably due to the sequencing of a pseudogene. Nearest species: North American sister species E. amitaria (Guenée, 1858) BIN-sharing (minimum distance: 1.5%).

Lycia Hübner, 1825 Lycia Hübner, [1825] 1816, Verz. bekannter Schmett.: 319. Type species: [Phalaena] hirtaria Clerck, 1759. Genus synonyms after Scoble (1999). Nyssia Duponchel, 1829, in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 7 (2): 283. Type species: Geometra zonaria Denis & Schiffermüller, 1775. Amorphogynia Warren, 1894, Novit. zool. 1: 429. Type species: Boarmia necessaria Zeller, 1849. Poecilopsis Harrison, 1910, Entomologist 43: 197. Type species: Phalaena pomonaria Hübner, 1790. ‡ Melanocoma Harrison, 1910, Entomologist 43: 197. Type species: Phalaena alpina Sulzer, 1776. [Junior homonym of Melanocoma Wollaston, 1867 (Coleoptera)]. Hypapocheima Wehrli, 1938, Ent. Rdsch. 55: 355. Type species: Apocheima suleimania Wehrli, 1938. Harrisonodes Leraut, 2009, Moths of Europe, vol. 2: 110. Type species: Phalaena alpina Sulzer, 1776. For synonymy see genetic data under L. zonaria.

Diversity and distribution: The genus Lycia contains 16 species in the Palaearctic and Nearctic regions. External characters and abdomen: Males fully-winged, thorax thick, furry. Wings sparsely scaled, grey tones predominating. Postmedial, medial and antemedial lines usually blackish, more or less distinct. Females brachypterous with a few exceptions: in Europe L. hirtaria fully winged, L. necessaria half-winged. Wing stumps of ‘wingless’ species with long hairy 275

scales. Antennae bipectinate in ♂, filiform in ♀. Hindtibia of both sexes with two apical spurs. Abdominal tergites covered densily with bifurcate setae (Nupponen & Sihvonen 2013). Male genitalia: Uncus wide, triangular, apically with hooked tip. Gnathos forming a ventrally fused small ring. Valva simple, elongated, in the centre slightly constricted, base wider, slightly setose. Apex rounded, with a group of setae. Saccus rounded, in some species with inwardly directed projection. Aedeagus short, thick, straight or slightly bent. Apically two projections, called “arms” by Leraut (2009). Vesica with inconspicuous, small, spinose sclerite. Female genitalia: Papillae anales elongate, striated, retractile, posteriorly setose. ­Apophyses posteriores very long, 6−12.5 mm long, straight or slightly bent. Apophyses anteriores shorter, 2.7−4.5 mm long, both usually ending spatulate. Apophyses variable, even in length, sometimes asymmetrical in the same specimen. Corpus bursae without sclerotization, striated, in European Lycia with distinct signum except for L. zonaria. Remarks: Lycia associates in several multi-gene analyses with non-European genera (Jiang et al. 2017; Murillo-Ramos et al. 2019). The most extensive hybridization experiments were carried out by Harrison. In a number of publications he created (unavailable) names for primary and secondary hybrids. They are listed by Prout (1912−1916) and Wehrli (1939−54). Not rarely, hybridization apparently also occurs in nature. As a consequence, the transfer of mitochondrial DNA from one taxon to another (‘genetic introgression’) frequently leads to shared DNA barcodes between ‘good’ species and to overlapping ‘mosaic’ haplotype patterns.

87. Lycia hirtaria (Clerck, 1759) Phalaena hirtaria Clerck, 1759: Icones Insectorum Rariorum 1: pl. 7, fig. 1 (Sweden). Lectotype ♂ (NHRS). Phalaena eremita Scopoli, 1763: Ent. Carniolica: 201, fig. 501 (Carnia). Syntype(s). Phalaena atomaria Hufnagel, 1767: Berlin Mag. 4 (5): 510 (Germany: Berlin region). Syntype(s). Phalaena Geometra contiguaria Borkhausen, 1794: Natur. eur. Schmett. 5: 187 (Europe). Syntype(s). Geometra congeneraria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2): pl. 33, fig. 174 (Europe). Syntype(s) ♂. Biston hirtarius var. hanoviensis Heymons, 1891: Berl. ent. Z. 34 (2): 446, pl. 15, figs 9, 9a (north-western Germany). Syntypes. Validated at subspecific rank by Scoble (1999), synonym according to Leraut (2009), confirmed here, see Remarks. Lectotype ♂ (Naturkundemuseum Hanau, Seidler 1939). Biston hirtaria var./ab. duplicaria Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (Edn 3) 1: 336 (Mauretania; Germany). Syntype(s). Biston hirtaria f. istriana Galvagni, 1901: in Rebel, Verh. zool.-bot. Ges. Wien 51 (8): 625 (Istria: Monte Spaccato; Padrich; Baowitza; Italy: Triest). Syntypes. Validated at subspecific rank by Scoble (1999) and Flamigni (2007), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Biston hirtaria var. uralaria Krulikovski, 1909: Mater. Pozn. Fauny Flory SSSR 9: 163 (Russia: Vyatka Province). Syntypes including ♀. Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Lycia hirtaria f. diniensis Oberthür, 1913: Études Lépid. comp. 7: 250, pl. 163, fig. 1590 (Basses-­ Alpes; Abruzzes). Syntype(s). Validated at subspecific rank by Scoble (1999) and Leraut

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(2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Lycia hirtaria sibirica Djakonov, 1926: Ezheg. gosud. Muz. N. M. Martyanova 4 (1): 65 (Russia: mountains E of Minusinsk). Syntypes 1♂3♀. Biston hirtarius Stephens, 1929: Nom. Br. Insects: 43. Emendation (gender agreement) of hirtaria Clerck. Lycia hirtaria parallelaria Inoue, 1958: Tinea 4 (2): 254, pl. 34, figs 28, 29 (Japan: Hokkaido, Sorachi-gun, Yamabe-musa). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999). Lycia hirtaria pusztae Vojnits, 1971: Acta zool. hung. 17: 455, fig. 1b (Hungary: Inundation Woods, Lakitelek). Holotype ♂ (TMB). Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Varga et al. (2010) suggest synonymy with taxon hanoviensis. Lycia hirtaria cataloniae Vojnits & Mészáros, 1973: Acta zool. hung. 19 (1/2): 212, fig. 1b (Spain: Prov. Teruel, Albarracín). Holotype ♂ (TMB). Validated at subspecific rank by Scoble (1999) and Leraut (2009), downgraded to synonymy by Redondo et al. (2009). Lycia bulgariensis Vojnits & Mészáros, 1973: Acta zool. hung. 19 (1/2): 212, fig. 1a (Bulgaria: Kostinbrod). Holotype ♂ (MNHN). Validated at species rank by Scoble (1999), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Unavailable names (hybrids): arethusa: Harrison (hirtaria ♂ X isabellae ♀); buloveci: Harrison (1916) (hirtaria ♂ X graecarius ♀); denhami: Harrison (1910) (hirtaria ♂ X zonaria ♀); pilzii: Standfuss (1891) (hirtaria ♂ X pomonaria ♀); recognita: Harrison (hirtaria ♂ X alpina ♀); wallacei: (Harrison) (hirtaria ♂ X lapponaria ♀). – (infrasubspecific): albescens: L ­ empke (1952) (f.); aurogutta: Leraut (2009) (f.); congeneraria: Hübner (1799) (ab.); contrasta: Lempke (1970) (f.); extrema: Lempke (1970) (f.); flavescens: Kitt (1914) (ab.); flavida: Lempke (1970) (f.); fumaria: Haworth (1809) (ab.); marginata: Lempke (1970) (f.); terroraria: Krulikovski (1908) (ab.); variegata: Lempke (1952) (f.).

External characters and abdomen: The only European Lycia species with fully developed wings in females. Wingspan ♂ 34–45 mm. Ground colour whitish grey to yellowish grey, irrorated with dark scales. Male forewing: Antemedial, medial and postmedial lines black, on veins thickened, usually wider near costa. Medial line closer to postmedial line, sometimes touching it. Subterminal line weak, often indistinct, outwardly edged with a diffuse whitish, slightly bent band. Terminal line distinct, dotted. Fringes of ground colour, chequered with black spots. Discal spot usually distinct, integrated into the medial line or slightly outwards. Hindwing more homogeneous, antemedial line absent. ♂ with slightly convex forewing costa. All transverse lines weaker in females than in males. Veins blackened. Discal spots more or less distinct. Frons with long hairy scales, sandy coloured, laterally dark brown. Vertex and thorax furry, mixed with sandy and brown hairy scales. Darker specimens with reduced number of sandy scales. Abdomen greyish brown, with hairy scales. Antennae bipectinate with noticeably long branches and speckled shaft in ♂, filiform, black with white dots in ♀. Hindtibia of both sexes with two apical spurs. Variation: Highly variable species. Ground colour can be darker or lighter. Extreme male forms are uniformly blackish. All transverse lines or some of them are indistinct or distinct, black or grey. The whitish band can be present or absent. The area between postmedial line and subterminal line is sometimes darkened. Because of the geographical variation numerous subspecies have been described. Most of them lie within the range of normal variation and can be observed in many localities in different percentages. Specimens from Istria (‘f. ­istriana’) described as large, light coloured, ground colour yellowish or white, pat 277

tern with high contrast. In southern France (‘f. diniensis’) often with ash-grey tinge. In Hungary (‘f. pusztae’) usually with ground colour uniformly brown, whitish band conspicuous, hindwing brownish yellow, thorax and abdomen covered with yellowish brown hairy scales. Male genitalia: Uncus wide, lateral margins straight, apically triangular with hooked tip. Gnathos forming ventrally fused ring with upturned rounded apex. Both weakly sclerotized. Valva simple, slender, weakly setose, in centre slightly narrower, apex round, with group of setae. Saccus rounded, with upwardly directed projection. Juxta broad at base, centre sometimes constricted. Aedeagus with two ‘arms’, narrow, caecum tapered. Vesica with small sclerite with dentate margin. Female genitalia: Papillae anales elongate, striated, retractile, posteriorly setose. Apophyses posteriores very long (on average 11 mm), straight, apophyses anteriores shorter (on average 3.7 mm), both spatulate at tip. Lamella postvaginalis sclerotized rectangular plate, anteriorly double-pointed. Lamella antevaginalis and posterior part of ductus bursae weakly sclerotized. Ductus bursae funnel-shaped, membranous, striated. Corpus bursae long, membranous, striated. Its posterior part wider than ductus bursae. Between posterior and anterior part of corpus bursae a distinct constriction (in L. necessaria only weakly constricted), anteriorly expanded, bearing an irregularly shaped signum with marginal spines. Distribution and abundance: Palaearctic. In Europe from Ural mountains to British Isles and Portugal and from northern Fennoscandia to the Mediteranean region including Corsica and Sardinia. Not mentioned for Corsica according to Fauna Europaea (Hausmann et al. 2004; 2011a), but three specimens from northern Corsica in coll. Skou. – Rather common, now and then occurring in numbers. – Outside Europe in Georgia, Armenia and Kazakhstan, and through Siberia to Far East Russia, Sakhalin, Kamchatka (subsp. sibirica) and Japan (subsp. parallelaria).

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Phenology: Univoltine: In southern Scandinavia from mid-April to late May, in south-western Germany from late March to early May (Ebert 2003), in Italy from February or March to May (Flamigni et al. 2007) and in Spain from March to early June (Redondo et al. 2009). Larvae from May to early July (Ebert 2003). Hibernation as pupa (Z. Las˘tu˚ vka pers. comm.). Adults active at night, males readily attracted to light, females rarely. Most females apparently found by day with both sexes sitting on tree trunks. Biology: Larva polyphagous. Found on Populus tremula, Salix caprea, Carpinus betulus, Corylus avellana, Betula pendula, Alnus glutinosa, Quercus, Ulmus, Malus domestica, Rosa canina, Prunus spinosa, Frangula alnus, Tilia platyphyllos, Lonicera xylosteum and Fraxinus excelsior (Ebert 2003). It has further been recorded on Robinia and Hippophae (Flamigni et al. 2007). Pupation in the ground (T. Drechsel pers. comm.) or among fine detritus on the ground (Z. Las˘tu˚ vka pers. comm.). Habitat: Silvicolous. In a large variety of habitats, such as deciduous forests, gardens, parks and other places with deciduous trees and bushes. In Scandinavia at low altitudes, in south-western Germany mainly between 100 and 500 m (Ebert 2003), in the Appenines up to 1300 m (Flamigni et al. 2007) and up to 1600 m in Spain (Redondo et al. 2009). Similar species: Males of L. necessaria (see there). Genetic data: BIN: BOLD:ACF3346 (n=61 from 14 countries from Spain and United Kingdom to Turkey and Kazakhstan, from Finland to Sardinia and Calabria; including the holotype of catalonica). Genetically homogeneous, but populations from Tunisia (n=4; BIN: BOLD:ACF5636) diverging by 1.2%. Typical forms of ‘hanoviensis’ and all material from Bulgaria (‘bulgariensis’) exactly barcode-sharing, supporting the synonymy of these taxa. Similarly, so far, almost all Lycia from Kazakhstan revealed to be exactly barcode-sharing, possibly pointing to synonymy of the taxon Lycia degtjarevae Viidalepp, 1986, described from Kazakhstan. Nearest species: North American L. ursaria (Walker, 1860) (1.0%), possibly requiring downgrading to subspecies rank. Nearest species in Europe: L. necessaria (exactly barcode-sharing, see below) and L. isabellae (2.3%). Remarks: Scoble (1999) validated the taxa hanoviensis Heymons, 1891 and uralaria Kruli­ kovski, 1909 at subspecific level. Several authors recently consider hanoviensis as a small, more yellowish aberration of hirtaria (for instance Leraut 2009; Malkiewicz 2012). Since hanoviensis coexists together with other forms in Europe (Wehrli in Seitz 1941) and DNA barcodes are exactly the same we follow Malkiewicz (2012) and consider this taxon as a form of the variable hirtaria (see also genetic data). The taxon uralaria has not been validated at subspecies rank in Russia by Viidalepp (1996) and Olshwang et al. (2004).

88. Lycia necessaria (Zeller, 1849) Boarmia necessaria Zeller, 1849: Stettin. ent. Ztg. 10 (7): 206 (Turkey: Ephesus). Syntype(s). Biston inversarius Rebel, 1903: Annln. naturh. Mus. Wien. 18 (2–3): 265, pl.3, fig. 8 (Bulgaria: Slivno). Holotype ♀. Orig. as “?hybrid. n.”. The name would potentially have nomenclatural priority over the following taxon if validated at subspecies level. Amorphogynia necessaria grupchei Carnelutti, 1986: Fragm. balcan. 13 (6): 61, fig. 1: 3, 4; fig. 2: 2 (Macedonia: Titov Veles-Babuna). Holotype ♂. Validated at subspecific rank (Scoble 1999), paratype DNA barcoded (see below). Herewith downgraded to synonymy based on absence of constant and significant differential features.

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External characters and abdomen: Wingspan ♂ 30−40 mm, ♀ about 20 mm. Male: Ground colour whitish grey, heavily irrorated with grey. Forewing: Antemedial, medial and postmedial lines dark grey, on veins thickened (weaker than in hirtaria), at costa usually wider. Medial line close to postmedial line, undulating, usually faint. Subterminal line weak, often indistinct, outwardly edged with diffuse light grey, slightly bent band. Terminal line dotted, sometimes indistinct. Fringes of ground colour, chequered with grey spots. Discal spot oval, black, usually on medial line or nearby. Hindwing more homogeneous, less irrorated, ante­ medial line absent. Discal spot diffuse, light grey. Frons and vertex rough scaled. Vertex black or dark brown, frons paler. Thorax with a mixture of silver and black long hairy scales. Abdomen greyish brown. Antennae partially bipectinate (about 9–11 apical flagellomeres free of branches). Female: Wings reduced, apex pointed, heavily irrorated with dark grey. Frons, vertex, thorax and abdomen with long black and white scales. Antennae filiform. Variation: Males little varying. Transverse lines more or less distinct. Macedonian population (‘grupchei’) described as follows (cf. Carnelutti 1986): On forewing postmedial line rarely double, medial line weak, antemedial line distinct. All transverse lines on the hindwing weak. We are of the opinion that all these features are variable, and can be found throughout the whole distribution area in different percentages. Further investigations are necessary to confirm the validity of the subspecies. Male genitalia: Generally as in L. hirtaria. No diagnostic difference in uncus and gnathos. Valva simple, slender. Costa and sacculus concave, weakly setose, apex with more setae. Sacculus at 1/4 from base distinctly curved, in L. hirtaria weakly bent. Juxta wider than in L. hirtaria. Saccus rounded. Aedeagus apically with two wider ‘arms’, caecum tapered with distinct process. Vesica with elongated, smoothly bordered plate. Female genitalia: Generally as in L. hirtaria. Papillae anales elongate, striated, retractile, posteriorly setose. Apophyses posteriores very long (the longest in European Lycia), length on average 12 mm, straight. Apophyses anteriores shorter, on average 4.7 mm, all apophyses ending spatulate. Lamella postvaginalis sclerotized, slightly trapezoid plate, anteriorly straight. Lamella antevaginalis and posterior part of ductus bursae weakly sclerotized. Ductus bursae funnel-shaped, short, striated. Corpus bursae long, membranous, striated. Distal half narrow, anteriorly sac-shaped, bearing an irregularly shaped signum with marginal spines. Between posterior and anterior part of corpus bursae less constricted than in L. hirtaria. Distribution and abundance: (East-)European-West Asiatic. In Europe only in Macedonia and Bulgaria (type locality of inversarius). According to Leraut (2009) and Fauna Europaea (Hausmann et al. 2004; 2011a) in Greece, but we have not been able to trace any verified records from there. In Fauna Europaea (Hausmann et al. 2004; 2011a) not recorded from Macedonia. – Local and rare, usually found as single specimens. – Outside Europe only in Turkey, western Caucasus, Georgia and Azerbaijan. Phenology: Univoltine: Scarce European records from early to late April, in Turkey also from late March (C. Tautel pers. comm.). In Turkey, caterpillars found in June (T. Drechsel pers. comm.). Hibernation as pupa. Biology: Larva polyphagous. Found on Paliurus spina-christi (Georgia: B. Müller own data, J. Gelbrecht pers. comm.) and on Elaeagnus angustifolia (Kappadokia, Turkey: T. Drechsel pers. comm.). Recorded on Quercus petraea, Carpinus betulus and Sorbus torminalis (Mol 1977). Larva reared on Prunus spinosa (B. Müller own data). Pupation in the ground. 280

Habitat: Xerothermophilous. Recorded on south-facing slopes. In Europe at lower altitudes, in Caucasus at 700 m and in Turkey at 1000 m (J. Gelbrecht pers. comm.). Similar species: Males very similar to lighter, faintly patterned forms of males of L. hirtaria. Genetic data: BIN: BOLD:ACF3346 (n=3 from Macedonia and Turkey, including a paratype of grupchei and a short-winged female from eastern Turkey). In Europe – including the paratype of grupchei – exactly sharing the barcode of Lycia hirtaria. The short-winged female from eastern Turkey diverging by 0.9%. Genetic patterns deserving further study but synonymization of Amorphogynia with Lycia at genus level well confirmed by DNA data (necessaria was often combined with Amorphogynia in previous literature). Remarks: Downgrading to synonymy of L. hirtaria disputable, but female winglessness of L. necessaria correlated with slight differential features in genitalia and wing pattern pointing to different species despite DNA barcode sharing with L. hirtaria. Barcode sharing may be the consequence of occasional hybridization (genetic introgression) as it often occurs in Lycia.

89. Lycia lapponaria (Boisduval, 1840) Nyssia lapponaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 195 (Lapponia). Syntype(s). Poecilopsis lapponaria scotica Harrison, 1916: Naturalist, Hull No. 716: 278 (Scotland). Syntype(s). Valid at subspecific rank (Scoble 1999). Poecilopsis lapponaria livonica Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 420, pl. 34: h ([Latvia]: Kurland). Syntypes including 1♂1♀ (ZFMK). Poecilopsis lapponaria sajanensis Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 420, pl. 34: i (Russia: Sajan Mountains, Chulugaischa). Syntype(s) ♂ (ZFMK). Poecilopsis lapponaria scotiae Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 420, pl. 34: h (Scotland: Perth, Rannoch). Syntype(s) ♂ (ZFMK).

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Unavailable names (hybrids): lappograecaria: Bretschneider (1939) (lapponaria ♂ X graecaria ♀); leesi: Harrison (lapponaria ♂ X hirtaria ♀); mirabilis: Harrison (lapponaria ♂ X isabellae ♀); smallmani: Harrison (1910) (lapponaria ♂ X zonaria ♀); tarda: Harrison (lapponaria ♂ X alpina ♀).

External characters and abdomen: Male: Wingspan 27−34 mm (Leraut 2009). Wings weakly scaled, therefore hyaline. Ground colour light grey with diffuse lines. Veins black. Terminally directed cell margin conspicuously black. On forewing postmedial, medial and antemedial lines dark grey, usually visible. Medial line approaching postmedial line near ­inner margin, sometimes with grey filling between. Subterminal line often a grey, diffuse band. Whitish wavy line indistinct. Costal area reddish brown, irrorated with black, interrupted by thickened dark grey lines. Fringes brownish grey, not chequered. Medial line of hindwing indistinct. Frons rough scaled, black, vertex usually reddish brown. Collar black, thorax strongly covered with mixed black and white hairy scales, laterally dark. Abdomen black with central stripe of reddish brown blotches. Antennae bipectinate with 5–6 apical flagellomeres free of branches. Hindtibia with 2 spurs. Female: Wings reduced, tapered, covered with single reddish scales and long grey hairy scales with whitish tips. Thorax and abdomen with black and reddish scales and pale hairy scales. Reddish scales are concentrated in the centre forming a stripe as in males. Scales are narrow with three or four tips. Variation: Males varying little, ground colour from lighter to darker grey, lines more or less distinct. Medial line of hindwing indistinct. Medial and postmedial lines of forewing sometimes only marked by a blotch near inner margin. Subsp. scotica is an isolated Scottish race with male wingspan 31−34 mm (Skinner 2009). Male genitalia: Generally as in L. hirtaria and L. isabellae. Gnathos forming a ventrally fused ring with upturned rounded apex, fused part and ring wider than in L. hirtaria. Valva simple, elongated, not constricted at centre, length on average 2.2 mm, shorter than valva of L. hirtaria and L. isabellae, at base wider. Costa concave. Saccus rounded, upwardly directed projection short. Juxta at base and apex wide, in the centre constricted. Aedeagus with arms, shorter than in L. hirtaria, vesica with a small sclerite with dentate margin. Female genitalia: Generally as in L. isabellae. Papillae anales elongate, striated, retractile, posteriorly setose. Apophyses posteriores very long (on average 9 mm), straight, apophyses anteriores shorter (on average 3.7 mm), both ending in the form of a small spatula. Lamella antevaginalis alate, membranous. Lamella postvaginalis developed as an inconspicuous, weakly sclerotized plate. Ductus bursae funnel-shaped, membranous, striated, anteriorly elongate, here with the origin of ductus seminalis. Corpus bursae long, membranous, striated. Anteriorly expanded, bearing semicircular signum with marginal spines. Distribution and abundance: (North) European-(West) Siberian. In Europe from northern Ural mountains to Scandinavia and Scotland (subsp. scotica) and from the Arctic Sea to central Fennoscandia and northern Balticum. – Local and uncommon, now and then occurring in numbers. – Outside Europe only recorded from Sayan Mountains, and possibly also from south-western Siberia (Mironov et al. 2008). Phenology: Univoltine: In Scotland in April (Skinner 2009). In southern Fennoscandia only within a short period, often only a few days, usually in late April or early May. Further north with more uncertain flight period, scarce records from late May to mid-June (Skou 1984). In Finland emerging when there are still small patches of snow on the ground (P. Sihvonen own data). Larva in Scotland from mid-May to early August (Porter 2010), in Finland from 282

May to July (Seppänen 1970). Hibernating as pupa, in rearings overlying for up to four winters (Skinner 2009). Males active at sunset. Most females in Finland found on Pinus trunks growing on moors/wetlands, not being a food-plant but the bark facilitating their detection. Females also found on trunks of Betula nana, but there they are even more difficult to find (P. Sihvonen own data). Biology: Larva polyphagous. In Fennoscandia recorded on Betula nana (Skou 1984), in Scotland on Myrica gale, Salix, Calluna vulgaris and Erica tetralix (Skinner 2009). In rearings accepting Betula (Skinner 2009). Pupation in the ground (Porter 2010). Habitat: Hygrophilous, boreo-montane. In southern Fennoscandia in moors and in the north in mountain heaths from the treeline and somewhat up, from about 500 up to 800 m altitude. Similar species: L. lapponaria, L. isabellae, L. pomonaria and the North American L. rache­ lae (Hulst, 1896) form a group of very similar species with brachypterous females (see External characters). Genetic data: BIN: BOLD:AAB1160 (n=3 from Finland), BIN-sharing with L. pomonaria. Diverging into two intraspecific clusters at 1.4% distance, in one of them exactly barcode sharing with L. pomonaria probably due to genetic introgression. The haplotype of this shared cluster exactly matches that of the North American L. rachelae (Hulst, 1896).

90. Lycia isabellae (Harrison, 1914) Poecilopsis isabellae Harrison, 1914: Entomologist 47: 93–94 (Austria: Tyrol, Innsbruck). Syntypes 1♂1♀. Poecilopsis isabellae var. silesiaca Harrison, 1917: J. Genet. 6 (4): 270 ([Poland]: Silesia). Syntype(s).

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Lycia isabellae strobinoi Dujardin, 1965: Entomops 1: 3, figured (France: Esteng, Cayolle region, towards Alpes Maritimes, 2000 m). Holotype ♂. Valid at subspecific rank (Scoble 1999). Herewith synonymized with nominotypical subspecies based on the absence of significant and constant differential features. Unavailable names (hybrids): oberthuri: Harrison (isabellae ♂ X zonaria ♀); omissa: Harrison (isabellae ♂ X lapponaria ♀); proxima: Harrison (1916) ({isabellae ♂ X pomonaria ♀} ♂ X {pomonaria ♂ X isabellae ♀} ♀); robsoni: Harrison (1916) (isabellae ♂ X pomonaria ♀); soffneri Harrison (isabellae ♂ X hirtaria ♀). – (incorrect subsequent spelling): strobioni: Scoble (1999).

External characters and abdomen: Generally as L. lapponaria, see above, but with following differences. Male forewing: Pattern on average more reduced, lines usually faint. Terminal area grey, only divided by the weak wavy line. Hindwing without or with very weak postmedial line. Fringes darker. Collar narrowly white, thorax laterally paler. Abdomen black, with long white hairy scales and beige (not reddish brown) scales. Antennae with 4−8 flagellomeres free of branches. Female: Wing stumps mainly covered with reddish and white scales and long pale grey hairy scales, whitish ending. Thorax and abdomen with a mixture of black, white and reddish scales, without reddish median stripe. Scales are narrow with three or four tips as in L. lapponaria (see L. pomonaria). Variation: Very little, ground colour varies from lighter to darker grey, terminal area sometimes darker than ground colour. Male genitalia: Generally as in L. hirtaria and L. lapponaria. Gnathos forming a ventrally fused ring with upturned rounded apex, fused part and ring wider than in L. hirtaria. Valva simple, elongated, in the centre weakly constricted, length on average 2.6 mm, longer than in L. lapponaria, at base wider. Costa concave. Saccus and juxta as in L. lapponaria. Aedea­ gus with ‘arms’, longer than in the latter. Vesica with a small plate with dentate margin. Female genitalia: Papillae anales elongate, striated, retractile, posteriorly setose. Apophyses posteriores very long (6.5−9 mm), straight, apophyses anteriores shorter (2.7−3.9 mm), both ending in the form of a small spatula. Lamella antevaginalis alate, membranous. Lamella postvaginalis developed as an inconspicuous, weakly sclerotized plate. Ductus bursae funnel-shaped, membranous, striated, anteriorly elongate, here with the origin of ductus seminalis. Posterior part of corpus bursae slender, striated. Anterior part expanded, bearing a triangular signum with three tips and marginal spines. Distribution and abundance: European. Limited distribution from southernmost Poland and southern East Germany through the Czech Republic, western Slovakia and most of Austria to the southern Alps in Italy, Switzerland and south-eastern France. Recently, the species is expanding in Saxonia and Brandenburg in Germany. – Rather local, now and then found in numbers. Abundance fluctuating distinctly between years and over longer time periods (Z. Las˘tu˚ vka pers. comm.). Phenology: Univoltine: From late February to mid-April, at higher altitudes until late May, often emerging simultaneously with the flowering of Primula in its habitats (N. Pöll pers. comm.) and occurring immediately after the snow has melted around the tree trunks (M. ­Leipnitz pers. comm.). Caterpillar from May (Fajčik 2003) until at least early July (D. Fritsch pers. comm.). Hibernation as pupa. Males active at night, but mainly observed sitting on tree trunks by daytime, females found the same way (J. Gelbrecht, M. ­Leipnitz, Z. Las˘tu˚ vka pers. comm.). Both sexes best searched for before noon when being in ­copulation (N. Pöll pers. comm.). Adults usually emerging in the afternoon and then sitting on tree trunks about 1.5 m above the ground (Z. Las˘tu˚ vka pers. comm.). 284

Biology: Larva monophagous. Recorded on Larix decidua (Robineau et al. 2007). Pupation in the ground (J. Gelbrecht, M. Leipnitz pers. comm.) or in fine detritus on the ground (Z. Las˘tu˚ vka pers. comm.). Habitat: Silvicolous, montane to subalpine. In forests with Larix decidua. In Valais in Switzerland in old, open Larix forests and also where younger and more scattered trees are growing (D. Fritsch pers. comm.), in semi-old and old Larix forests (M. Leipnitz pers. comm.), but not in dense forest (N. Pöll pers. comm.). Vertical distribution from 1100 m up to 2000 m (D. Fritsch pers. comm.). Similar species: See remarks under L. lapponaria. Genetic data: No BIN assigned (n=2 from Austria, 407bp). Genetically homogeneous. Nearest species: L. pomonaria (1.0%), L. lapponaria (2.0%), L. hirtaria (2.3%). North American L. rachelae (Hulst, 1896) (1.5%).

91. Lycia pomonaria (Hübner, 1790) Phalaena Geometra pomonaria Hübner, 1790: Beitr. Gesch. Schmett. 2 (3): 73, pl. (3) 4, figs U, 1–3 (Ukraine: Niemirow). Syntypes ♂♀. Bombyx grisea Thunberg, 1784: Diss. ent. sistens Insecta Suecica 1: 1 (Sweden: Uppland). Syntype(s). Psyche stigmatella Zetterstedt, 1840: Insecta Lapponica: 928 (Lapponia). Syntypes ♂. Nyssia vertumnaria Lefèbvre, 1835: Annls. Soc. ent. Fr. 1835 (1): 102 (Germany; France). Syntypes ♂♀. Poecilopsis borealis Rangnow, 1935: Ent. Rdsch. 53 (2): 22, pl. 4, fig. 12 (Northern Lappland). Syntype(s).

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Lycia pomonaria nigricans Beljaev, 1996: Chteniya Pamyati Alekseya Ivanovicha Kurentsova 6: 67, figs 27, 28 (Russia: Primorskii Krai, Bikin river, Verkhnii Pereval). Holotype ♂ (ZISP). Valid at subspecific rank (Scoble 1999). Unavailable names (hybrids): brooksi: Harrison (1913) (pomonaria ♂ X {hirtaria ♂ X pomonaria ♀} ♀); christianae: Harrison (1916) (pomonaria ♂ X isabellae ♀); confusa: Harrison (1916) ({pomonaria ♂ X lapponaria ♀} ♂ X {pomonaria X isabellae ♀} ♀); helenae: Harrison (1910) (pomonaria ♂ X zonaria ♀); heslopi: Harrison (1916) (pomonaria ♂ X lapponaria ♀); hulli: Harrison (1916) ({pomonaria ♂ X hirtaria ♀} ♂ X hirtaria ♀); hünii: Oberthür (1897) (pomonaria ♂ X hirtaria ♀); connexa: Harrison (1916) ({pomonaria ♂ X isabellae ♀} ♂ X zonaria ♀); felkeli: Harrison (1916) ({pomonaria ♂ X isabellae ♀} ♂ X {isabellae ♂ X pomonaria ♀} ♀); inversa: Harrison (1916) ({pomonaria ♂ X isabellae ♀} ♂ X {isabellae ♂ X pomonaria ♀} ♀); laricis: Harrison (1916) ({pomonaria ♂ X isabellae ♀} ♂ X isabellae ♀); mixta: Harrison (1916) ({pomonaria ♂ X lapponaria ♀} ♂ X zonaria ♀); haeneli: E. Müller (1920) ({pomonaria ♂ X hirtaria ♀} ♂ X pomonaria ♀); – (infrasubspecific): pallidata: Leraut (2009) (f.).

External characters and abdomen: Generally as L. lapponaria and L. isabellae, see above, but with the following differences. Male: larger, wingspan 27−33 mm, forewing paler, coarser scaled, postmedial, medial and antemedial lines deeper in colour, distinct. Terminal area pale grey, speckled with dark scales. Fringes chequered pale grey and black. Collar broadly white, broader than in L. isabellae. Thorax paler, especially both tegulae. Abdomen with less reddish brown hairy scales. Antennae with 7−10 flagellomeres free of branches. Female: Wing stumps shorter, covered with white and black scales and pale grey hairy scales, whitish ending. Thorax and abdomen as in L. isabellae. Scales much broader, ending in up to eight small tips (main difference). Variation: Very little, ground colour varies from lighter to darker grey, lines sometimes less distinct. Male genitalia: Uncus wide with oval base, apically triangular with hooked tip. Gnathos forming ventrally fused ring. Fused part angled, rounded. Valva simple, weakly setose, in centre less constricted, base wider. Apex round, with group of setae. Saccus rounded, with inwardly directed projection. Juxta heart-shaped, apically with projection of double length of the heart-shaped base. Aedeagus slightly bent with ‘arms’. Vesica with bent and laterally spinose sclerite. Female genitalia: Generally as in L. isabellae and L. lapponaria, but with following differen­ ces: Funnel-shaped ductus bursae much smaller. Distal, narrow part of corpus bursae longer. Signum forms an irregular triangle with spines on margins and tips. Distribution and abundance: Eurasiatic. In Europe from the Urals to Scandinavia, central ­Europe and France and from northernmost Fennoscandia to central Europe, in the southeast the distribution reaching the northern Balkan peninsula. – In northern Fennoscandia common, further to the south more local. Usually found in limited numbers. – Outside ­Europe through southern Siberia to Far East Russia and Kamchatka (subsp. nigricans). Phenology: Univoltine: In Fennoscandia from mid- or late April to late May (Skou 1984), in south-western Germany from mid-March to mid-April (Ebert 2003). In Finland the species emerges when there are still small patches of snow on the ground. Larva in Fennoscandia from late June to early August (Skou 1984), in south-western Germany from May to late June (Ebert 2003). Hibernation as pupa. Males in Fennoscandia flying in the evening and

286

in warm nights, attracted to light. Females found on Betula trunks during day time (Finland: P. Sihvonen own data). After hatching the females quickly climb up the tree trunks and usually they do not stay below at the base of the trunks. Biology: Larva oligophagous. Found on Alnus glutinosa, Quercus, Pyrus communis and Malus domestica (Ebert 2003). In Fennoscandia recorded on Betula pubescens, Salix and Vaccinium (Skou 1984). Pupation in an open web between moss and soil on the ground (Skou 1984). Habitat: Silvicolous. In northern Fennoscandia in forests, especially in mountain Betula forests. In central Europe in deciduous forests and gardens (Ebert 2003) and forests with many Tilia trees (J. Gelbrecht, B. Tóth, Z. Las˘tu˚ vka pers. comm.). In northern Bavaria in Germany in forests with many Carpinus betulus trees (F. Rosenbauer pers. comm.). In Fennoscandia up to the treeline, in southern Germany from 100 up to 500 m altitude. Similar species: See remarks under L. lapponaria. Genetic data: BIN: BOLD:AAB1160 (n=6 from Finland, Germany), BIN-sharing with L. lapponaria. One barcode from Germany diverging by only 0.15% (1bp) from that of Finnish populations. In Finland diverging into two intraspecific clusters at 1.9% distance, in one of them exactly barcode sharing with L. pomonaria probably due to genetic introgression. The haplotype of this shared cluster exactly matches that of the North American L. rachelae (Hulst, 1896).

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92. Lycia zonaria (Denis & Schiffermüller, 1775) Geometra zonaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 100 (Austria: Vienna district). Syntype(s). Bombyx zona Fabricius, 1787: Mantissa Insect. 2: 131. Emendation of zonaria (Denis & Schiffer­ müller, 1775). Ithysia zonaria var. britannica Harrison, 1912: Entomologist 45: 316 (United Kingdom). Syntype(s). Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on homogeneity of DNA barcodes and absence of constant and significant differential features. Ithysia zonaria var. rossica Harrison, 1912: Entomologist 45: 316 (Russia: Ural Mountains). Syntype(s). Valid at subspecific rank (Scoble 1999). Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features. Nyssia atlantica Harrison, 1938: Proc. Univ. Durham phil. Soc. 10 (1): 21 (United Kingdom: Coll; Canna; Sanday; Barra; Mingulay). Syntypes. Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features. Unavailable names (hybrids): langei: Harrison (1910) (zonaria ♂ X pomonaria ♀); merana: Burrows (1906) (zonaria ♂ X lapponaria ♀); harrisoni: Harrison (1910) (zonaria ♂ X hirtaria ♀); – (infrasubspecific): albicans: Nitsche (1925) (ab.); obscura: Harrison (1912) (ab.).

External characters and abdomen: Males: Wingspan 28−35 mm. Ground colour of wings greyish white. Forewing costa slightly concave. Terminal area and fringes brownish grey, divided by white line, proximally bordered by oblique white band. Postmedial and antemedial lines diffuse, latter on forewing angled near costa, on hindwing absent. Medial area often, basal area sometimes dark grey, such forms without contrasted transverse lines. Discal spots distinct, elongate linear. All veins strongly marked blackish. Frons with group of long, dark brown and sandy coloured hairy scales, vertex dark brown with group of sandy hairy scales on both sides. Thorax blackish with two pale longitudinal bands. Abdomen black, with hairy scales, yellowish ringed. Antennae partially bipectinate (about 5−6 flagellomeres free of branches). Female with black wing stumps, covered with long pale hairy scales. Frons and vertex black with whitish or brownish hairy scales, frons also with pale scales. Abdomen black with whitish or brownish hairy scales, yellowish ringed, easily distinguished from other wingless females by their abdomen. Antennae filiform.

Variation: Males variable in wing span and extension of dark markings. Extreme forms are nearly black. The white subterminal band can be developed as narrow line or wider. Females less variable, hairy scales whitish or brownish. Specimens from the British Isles and Ireland (‘f. britannica’) with males having ‘much greyer markings’ (Harrison 1912), which are usually reduced proximally of the postmedial line. Populations of Isle of Baleshare and other Hebridean islands (‘f. atlantica’) described as smaller than ‘britannica’, male wingspan 24−29 mm, according to Skinner (2009) this is the only tangible character. Specimens from Ural (‘f. rossica’) mountains described as small, with strong markings and a line-shaped subterminal band (Harrison 1912). Male genitalia: Uncus wide, triangular, apically with hooked tip. Gnathos forming ventrally fused small ring, fused part conspicuous, angled, sclerotized, rounded. Valva simple, elongated, in centre slightly constricted, base wider, length on average 2.0 mm, apex round, with group of setae. Saccus rounded. Juxta inconspicuous, developed as weakly sclerotized plate. Aedeagus short, thick, straight. Apically with short ‘arms’, vesica with inconspicuous small spinose plate. 288

Female genitalia: Generally as in L. hirtaria. Papillae anales elongate, striated, retractile, posteriorly setose. Apophyses posteriores very long (on average 9 mm), apophyses anteriores shorter (on average 3.7 mm), both spatulate at tip. Lamella postvaginalis sclerotized, slightly trapezoid plate, anteriorly straight. Lamella antevaginalis circular, wide. Ductus bursae funnel-shaped, long, narrow, striated, posterior part sclerotized. Corpus bursae soft, membranous. Without signum. Distribution and abundance: Eurasiatic. In Europe from eastern European Russia to the British Isles including Ireland and from southernmost Scandinavia to northern Spain, the French Alps, the northern Alps and northernmost Balkan peninsula. – Local and generally uncommon, usually found in limited numbers. However, with records of mass occurrences and in some cases being a pest on cultivated plants, mainly in former times. A large outbreak by the Danish town Billund recorded in 1996 with thousands of males attracted to the light at night at a gas station; after searching for a suitable locality, males and females were found by day sitting in the vegetation, on fence posts etc. in large numbers. – Outside Europe in Turkey and subsp. rossica from northern Kazakhstan through southern Siberia to Trans-Baikal.

Phenology: Univoltine: In Denmark from early April to mid-May. In south-western Germany from early March to mid-April (Ebert 2003). Larvae from early June to mid-July. Overwintering as pupa, sometimes up to eight times (T. Drechsel pers. comm.; rearing observation). Males active at night, readily attracted to light, usually rather late. Both sexes found during daytime sitting in the vegetation, on fence posts etc. Biology: Larva oligophagous. Found on Medicago falcata, Lotus corniculatus, Onobrychis viciifolia, Daucus carota, Heracleum sphondylium, Galium boreale, Solidago gigantea, Achillea millefolium, Salvia pratensis, Artemisia campestris and Centaurea scabiosa (Ebert 289

2003), in Crimea on Hedysarum tauricum (V. Savchuk pers. comm.). In mass occurrences sometimes being pest on cultivated plants; for example, several fields with Daucus carota were destroyed in southern Denmark in 1943 (Hoffmeyer 1966). In other mass ­occurrences, a variety of food-plants have been used (J. Gelbrecht pers. comm.), especially Salvia nemorosa and Medicago sativa (Z. Las˘tu˚ vka pers. comm.). Pupation in or on the ground in a loose cocoon (J. Gelbrecht, T. Varenne pers. comm.). See Lévêque (2017) for an important and comprehensive overview on the biology, distribution and decline in France with special attention on the Massif Central. Habitat: Mesophilous. In sandy ground and dry meadows, mainly in places with low vegetation. Also in ruderal areas and calcareous low-nutrient meadows (J. Gelbrecht pers. comm.). In northern Germany in heaths and edges of bogs (F. Rosenbauer pers. comm.). In Hungary in dry, open grassland, on limestone and sand (B. Tóth pers. comm.). In Crimea in steppe slopes (V. Savchuk pers. comm.). In south-eastern Turkey on flower rich high mountains steppe (J. Gelbrecht pers. comm.). In Europe mainly in lowlands, in south-­ western Germany from 250 up to 400 m (M. Leipnitz pers. comm.), in the Czech Republic and in Slovakia from 110 up to 350 m, in southern Moravia up to 500 m (Z. Las˘tu˚ vka pers. comm.) and in the French Alps from 1400 up to 2000 m (T. Varenne pers. comm.). In Crimea from sea-level up to 250 m (V. Savchuk pers. comm.). In Armenia at 1860 m, in Turkey up to 2400 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAD9557 (n=13 from United Kingdom, France, Germany, Austria, Hungary, Turkey). Genetically homogeneous, exactly barcode-sharing with L. alpina and L. graecarius. DNA barcode sharing is an indication of comparatively recent hybridization and thus a strong argument against the validity of the genus name Harrrisonodes Leraut, 2009 as a separate genus which would separate L. zonaria and the other two aforementioned species in two different genera.

93. Lycia alpina (Sulzer, 1776) Phalaena Bombyx alpina Sulzer, 1776: Geschichte Insecten 1: 159 (Switzerland: Alps). Syntype(s). Phalaena alpinaria Römer, 1789: Genera Insectorum Linnaei et Fabricii iconibus illustrata: 75, pl. 21, figs 5, 6. Emendation of alpina Sulzer. Nyssia bombycaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 195 (Valesia; Lombardia). Syntype(s). Ithysia italica var. carniolica Harrison, 1912: Entomologist 45: 319 ([north-eastern] Italy). Syntype(s). Valid at subspecific rank (Flamigni et al. 2007). Unavailable names (hybrids): difficilis: Harrison (alpina ♂ X lapponaria ♀); incerta: Harrison (alpina ♂ X hirtaria ♀). – Unavailable names (infrasubspecific): extincta: Harrison (1910) (ab.); tenebraria: Rebel (1910) (ab.).

External characters and abdomen: Wingspan ♂ 33−37 mm. Males: Ground colour of wings slightly glossy white, sometimes weakly speckled with brown. On forewing postmedial, ­medial and antemedial lines oblique, distinctly thickened on veins, two former between anal vein and inner margin slightly oblique. Medial line extended at costa, closer to postmedial line than to antemedial line, sometimes medial line anastomosing with postmedial line. Antemedial line angled on costa. Terminal area with whitish wavy line proximally with grey shadow, and with mostly band-shaped subterminal line. On hindwing antemedial line 290

­ bsent, medial line often diffuse, postmedial line often distinct. Discal spot on hindwing disa tinct. Fringes of ground colour, terminal line interrupted. Frons rough scaled, covered with brown hairy scales. Thorax (vertex, patagia, tegulae) densely covered with white fur-shaped hairy scales above, brownish below. Abdomen dark brown with white-grey hairy scales. Antennae partially bipectinate (about 6–8 apical flagellomeres free of branches). Females with white hairy scales on frons. Subsp. carniolica very similar to L. florentina. Usually with whitish ground colour with distinct lines. Specimens with brownish ground colour are more similar to L. graecarius but can be distinguished by the postmedial line (Flamigni et al. 2007). Male genitalia: Generally as in L. florentina and L. graecarius. Uncus curved, apex with hooked tip like in L. florentina. Gnathos a ventrally fused band, fusion part spatulated. Valva simple, costa bent, in the middle slightly constricted, base wider. Length on average 2.9 mm. Apex round, with group of setae. Saccus rounded. Juxta like a compressed “8”. Aedeagus short (length on average 2 mm), thick, bent. Vesica with some weakly spinose ridges. Female genitalia: Papillae anales elongate, striated, retractile. Posteriorly setose. Apophyses posteriores very long, up to 10.5 mm, straight or slightly bent. Apophyses anteriores shorter, up to 4.5 mm long, spatulate at tip. Apophyses variable, even in length, sometimes asymmetrical in same specimen. Wing-shaped lamella antevaginalis and posterior part of ductus bursae sclerotized. Lamella postvaginalis developed as plate, posteriorly sclerotized, weakly rounded. Ductus bursae long, narrow, striated. Corpus bursae membranous, striated, without signum. Distribution and abundance: European. Restricted to the Alps with occurrence in France, Italy, Switzerland, Austria, Germany. In north-easternmost Italy and Slovenia replaced by subsp. carniolica. – Usually in limited numbers. – Unknown from outside Europe.

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Text-figs 145–147. Diagnostic characters (indicated) of selected European Lycia species (from above): Text-fig. 145. Lycia alpina (Sulzer, 1776), Switzerland, Visperterminen, 30.iv.1995 (B.  Müller). Text-fig. 146. L. graecarius (Staudinger, 1861), Croatia, Krk, Punat, 27.iii.2003 (B.  Müller). Text-fig. 147. L. florentina (Stefanelli, 1882), Italy, Emilia Romagna, Forli, 02.iii.1992 (P. Skou).

Phenology: Univoltine: In the Italian Alps from shortly after the snow melting in April or May to July depending on altitude. Larva from June to August, sometimes as early as May and as late as September, depending on altitude (M. Leipnitz pers. comm.). Fourth instar larva on 21st July at 2000 m altitude (D. Fritsch pers. comm.). Half grown larva in late July and mid-August, last instar larvae in late August (N. Pöll pers. comm.). Hibernation as pupa. Males active at night, attracted to light. Females found by day in sunshine when sitting or crawling around, relatively high up, in the vegetation (M. Leipnitz pers. comm.). Females also found early in the morning in copula with males, often 20 to 30 cm up in the vegetation (N. Pöll pers. comm.). 292

Biology: Larva polyphagous. Found on Hypericum, Knautia, Crepis (N. Pöll pers. comm.), Salix, Alnus, Dryas octopetala, Alchemilla, Vaccinium myrtillus and Carduus (M. Leipnitz pers. comm.). Recorded on Rumex, Ranunculus, Sisymbrium, Trifolium repens, Lotus corniculatus, Lithospermum, Scabiosa lucida, Bellis perennis, Achillea millefolium, Adenostyles alpina, Centaurea and Chaerophyllum (Flamigni et al. 2007). Pupation in the ground (M. Leipnitz pers. comm.). Habitat: Alpine. In open places at higher altitudes, below as well as above treeline. From 1300 up to 2500 m altitude. Similar species: L. graecarius and L. florentina (see no. 94, 95, and Text-figs 145–147). Genetic data: BIN: BOLD:AAD9557 (n=11 from Germany, Austria, Italy). Genetically homo­geneous, exactly barcode-sharing with L. zonaria, L. graecarius. Remarks: Leraut (2009) combines this and the following two species in his genus Harrisonodes, but from a molecular view this is not justified (see generic data under L. zonaria).

94. Lycia graecarius (Staudinger, 1861) Biston graecarius Staudinger, 1861: in Staudinger & Wocke, Cat. Lepid. Eur. und angrenzenden Länder 1: 70. Biston graecarius var./ab. istrianus Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (Edn 3) 1: 335 (Istria; southern Carniola). Syntype(s). Validated at subspecific rank (Scoble 1999), herewith synonymized with the nominotypical subspecies based on identical DNA barcodes and the absence of significant and constant differential features. Biston graecarius var. odrinarius Buresch, 1915: Spis. bulg. akad. nauk 12 (Nat. Sci. & Maths) (5): 48 (Turkey: Edirne, Fort Aiwas Baba). Holotype ♂. Validated at subspecific rank (Scoble 1999), herewith synonymized with the nominotypical subspecies based on the absence of significant and constant differential features.

External characters and abdomen: Wingspan ♂ 31−35 mm. Generally as in L. alpina. Males: Forewings silvery grey, sometimes weakly speckled with brown scales. On forewing postmedial, medial and antemedial lines oblique, on veins less thickened than in L. alpina, the first two between anal vein and inner margin slightly oblique, shorter than in L. florentina. Medial line straight, inwardly angled on vein M1. All transverse lines thickened at costa. Pattern in terminal area similar to that of L. alpina but less contrasting. Hindwing often more brownish, giving a bicolorous impression. Postmedial line angled on vein M1 (bent in L. florentina). Discal spot on hindwing faint, sometimes distinct. Fringes silvery. Frons covered with long reddish brown fur-shaped hairy scales. Thorax, vertex, patagia and abdomen like in L. alpina. Antennae partially bipectinate (about 5–6 apical flagellomeres free of branches). Females see genus description, differing from L. alpina in their almost or entirely black frons and vertex. Females often indistinguishable from those of L. florentina (Flamigni et al. 2007). Variation: Ground colour of ♂ varying from silvery grey to brown, also geographically. All transverse lines more or less distinct. Specimens from Istrian limestone areas (‘istrianus’) in north-western Croatia lighter with faint pattern. Pattern bicolorous, silvery grey on forewing and more brownish on hindwing, in fresh specimens (Flamigni et al. 2007). Frons apparently darker brown than in Greek specimens with reddish brown frons. Intermediate forms in southern Croatia, Bosnia-Herzegovina and Macedonia. Validity of taxon istrianus 293

questioned also by Flamigni et al. (2007). Ground colour of specimens from Thrace dark grey, lines weak or nearly absent, but wavy line distinct, bright (Buresch & Iltschew 1915). Male genitalia: Generally as in L. alpina and L. florentina. Uncus reversed heart-shaped with small hooked tip, not forming a folded structure. Gnathos a ventrally fused band, fused part spatulate. Valva simple, usually elongated, weakly setose. Costa bent, often slightly constricted at centre, base wider, length 2.4−2.5 mm. Apex rounded, setose. Shape of ­valva geographically variable in size (Flamigni et al. 2007). Aedeagus generally as in L. florentina, vesica with some weakly spinose ridges, without more strongly sclerotized, large ridge. Female genitalia: Generally as in L. alpina and L. florentina. Apophyses posteriores very long, up to 11 mm, straight or slightly bent. Apophyses anteriores shorter, length up to 4.5 mm, spatulate at tip. Lamella antevaginalis alate, wider than in L. alpina. Lamella antevaginalis and posterior part of ductus bursae sclerotized. Lamella postvaginalis inconspicuous, weakly sclerotized. Ductus bursae long, less narrow than in L. alpina, striated. Corpus bursae a membranous, striated sac, without signum. Distribution and abundance: (South-east) European. In Europe from north-easternmost Italy through the Balkan peninsula to Greece. – Rather locally distributed, now and then occurring in numbers. – Outside Europe in a small area of north-western Turkey.

Phenology: Univoltine: In Italy and Slovenia from early March to early May. In Croatia from early March to mid-May, in Bulgaria from late February to early April and in Greece from early March to late April. Last instar larvae found in early June in eastern Bulgaria (J. ­Gelbrecht pers. comm.). Hibernation as pupa. Males active at night, readily attracted to light. Females found by day when sitting on top of stones, on fence posts etc. Pupation in the ground (J. Gelbrecht pers. comm.). 294

Biology: Larva polyphagous. Found on Convolvulus (J. Gelbrecht pers. comm.) and Spartium junceum and Centaurea dichroantha (Lepiforum 2018) and recorded on Achillea millefolium, Cytisus scoparius, Taraxacum, Rumex, Trifolium, Medicago sativa, Lotus, Artemisia, Centaurea diffusa and Salix (Flamigni et al. 2007). Habitat: Xerothermophilous. Open meadows or meadows with scattered bushes and trees. In Italy up to 500 m, in Greece up to at least 1000 m altitude. Similar species: L. alpina and L. florentina. Males of L. alpina and L. graecarius with postmedial and medial lines between vein A and inner margin shorter than in L. florentina, less oblique (see Text-figs 145–147). Lines in L. alpina usually more distinct and black. Females with frons covered with white hairy scales, in L. florentina and L. graecarius frons entirely or almost black (Flamigni et al. 2007). Genetic data: BIN: BOLD:AAD9557 (n=5 from Bulgaria, north-easternmost Italy: Friuli). Genetically homogeneous, exactly barcode-sharing with L. zonaria, L. alpina.

95. Lycia florentina (Stefanelli, 1882) Biston graecarius florentina Stefanelli, 1882: Boll. Soc. ent. ital. 4 (2): 222 (Italy: Fiesolana). Syntypes ♂♀. Ithysia italica Harrison, 1912: Entomologist 45: 317 (Italy). Syntype(s). Valid at subspecific rank (Scoble 1999), downgraded to synonymy by Flamigni et al. (2007).

External characters and abdomen: Wingspan ♂ 30−35 mm. Similar to L. alpina and L. graecarius. Males: Forewings light grey, usually speckled with brown scales. On forewing postmedial, medial and antemedial lines oblique, less thickened on veins than in L. alpina. Terminal line interrupted, grey, often less distinct. Postmedial line on anal vein abruptly angled, between anal vein and inner margin very oblique, longer than in L. alpina and L. graecarius. Other male features as in L. graecarius. Females see above. They can be separated from L. alpina by their almost or entirely black frons. Females often cannot be distinguished from L. graecarius without males (Flamigni et al. 2007). Variation: Ground colour of ♂ varying from silvery grey to brown, also geographically. All medial lines more or less distinct. Male genitalia: Generally as in L. alpina and L. graecarius. Structure of uncus as in L. alpina. Gnathos a ventrally fused band, spatulate. Valva simple, costa bent, in the middle usually slightly constricted, base wider. Length on average 2.9 mm, but apex rounded, with group of setae. Valvae geographically variable in size (Flamigni et al. 2007). Aedeagus thick, bent, larger than L. alpina, length on average 2.7 mm. Vesica with some weakly spinose ridges, one ridge wider and more sclerotized. Female genitalia: Generally as in L. alpina and L. graecarius. Apophyses posteriores very long, up to 13 mm, on average longer than in the other species of this group. Apophyses anteriores shorter, up to 5 mm, spatulate at tip. Lamella antevaginalis alate, wider than in L. alpina. Ductus bursae long, less narrow than in L. alpina, striated. Corpus bursae a membranous, striated sac, without signum.

295

Distribution and abundance: Central Mediterranean. Only known from Italy throughout most of the country and from south-easternmost France, close to the border with Italy. Not recorded for France in Fauna Europaea (Hausmann et al. 2004; 2011a). – Males sometimes found in numbers (T. Varenne pers. comm.).

Phenology: Univoltine: From February or March to April, sometimes even into May (Flamigni et al. 2007). Larva from April to June (K. Takáts pers. comm.). Overwintering as pupa. Males active at night, readily attracted to light. Females found on tree trunks (T. Varenne pers. comm.; A. Hausmann own data). Biology: Larva polyphagous. Recorded on Genista tinctoria, Spartium, Medicago sativa, Trifolium, Hedysarum coronarium and Onobrychis viciifolia (Flamigni et al. 2007). Reared on Medicago falcata, M. sativa, Achillea millefolium and Salvia pratensis (M. Leipnitz pers. comm.). In earlier times recorded as pest on Medicago sativa (Flamigni et al. 2007). Pupa­ tion in the ground (M. Leipnitz, K. Takáts pers. comm.), in a loose cocoon of plant litter (T. Varenne pers. comm.). Habitat: Xerothermophilous. Open areas, meadows. From sea-level up to 1200 or 1300 m (Flamigni et al. 2007). Similar species: L. alpina and L. graecarius. Males of L. alpina and L. graecarius with postmedial and medial lines between vein A and inner margin shorter than in L. florentina, less oblique (see Text-figs 145–147). Lines in L. alpina usually more distinct and black. Females of L. alpina with frons covered with white hairy scales, in L. florentina and L. graecarius frons entirely or almost black (Flamigni et al. 2007). Genetic data: BIN: BOLD:AAF4970 (n=3 from southern Italy). Genetically homogeneous. Nearest species: L. zonaria, L. alpina, L. graecarius (all 3.1%). 296

Biston Leach, 1815 Biston Leach, [1815] 1830, Brewster’s Edinburgh Encyclopaedia 9: 134. Type species: Geo­ metra prodromaria Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Dasyphara Billberg, 1820, Enumeratio Insect. Mus. G. J. Billberg: 89. Type species: Geometra prodromaria Denis & Schiffermüller, 1775. Pachys Hübner, 1822, Syst.-alphab. Verz.: 38–44, 46,47,49,50, 52. Type species: Geometra prodromaria Denis & Schiffermüller, 1775. [Junior objective synonym of Biston Leach]. Amphidasis Treitschke, 1825, in Ochsenheimer, Schmett. Eur. 5 (2): 434. Type species: Geo­metra prodromaria Denis & Schiffermüller, 1775. [Junior objective synonym of Biston Leach]. Eubyja Hübner, [1825] 1816, Verz. bekannter Schmett.: 318. Type species: Phalaena betularia Linnaeus, 1758. Amphidasys Sodoffsky, 1837, Bull. Soc. imp. Nat. Moscou 10: 90. [Emendation of Amphidasis Treitschke]. Amphidasea Unger, 1856, Arch. Ver. Freunde Naturg.-Mecklenb. 10: 61. [Emendation of Amphi­ dasis Treitschke]. Buzura Walker, [1863] 1862, List Specimens lepid. Insects Colln Br. Mus. 26: 1531. Type species: Buzura multipunctaria Walker, [1863] 1862. Culcula Moore, 1888, Descr. new Indian lepid. Insects Colln late Mr W. S. Atkinson: 266. Type species Culcula exanthemata Moore, 1888. Eubyjodonta Warren, 1893, Proc. zool. Soc. Lond. 1893: 416. Type species: Eubyjodonta falcata Warren, 1893. Blepharoctenia Warren, 1894, Novit. zool. 1: 428. Type species: Amphidasys bengaliaria Guenée, 1858. Epamraica Matsumura, 1910, Thousand Insects Japan (Suppl.) 2: 130. Type species: Epamraica bilineata Matsumura, 1910.

Diversity and distribution: The genus Biston contains approximately 50 species, the distribution area covers the Palaearctic and Oriental regions as well as Africa. B. betularia also in North America. The Chinese species are revised by Jiang et al. (2011). External characters and abdomen: Large, robust species. Wingspan 30–80 mm. Females tend to be larger than males. Abdomen, thorax, tegula furry-looking. Fovea absent from male forewings. Wings white, pale yellow or greyish brown, transverse lines black, brown or white. On forewing antemedial, medial and postmedial lines usually wavy or dentate, on hindwing medial line indistinct, postmedial line wavy or dentate. Discal spot on forewing dot-shaped or elliptic, on hindwing less conspicuous or absent. Antennae bipectinate in male, filiform in female. Hindtibia with 2 x 2 spurs, without pencil. Venation: On forewing Sc free, R1 and R2 usually stalked. R2 sometimes connected with R3–R4 (R5) by a transverse vein. R3–R5 not stalked with M1. On hindwing Sc+R1 close to cell, M1 from anterior angle of cell. CuA1 before or from posterior angle of cell (Jiang et al. 2011). Male genitalia: Uncus short, broad, bifid. Gnathos generally present, arms connected medially, rugose. Socii usually absent. Valva simple, without distinct sclerotized structures, bearing long setae from centre to apex. Juxta elongate, acute. Aedeagus apically sclerotized, fluted, vesica simple, reflexed, scobinate (Holloway 1994; Jiang et al. 2011). Female genitalia: Papillae anales setose. Apophyses posteriores very long. Apophyses anteriores shorter. Lamella postvaginalis sometimes present, triangular or oval. Ostium bursae occasionally weakly sclerotized. Ductus bursae longitudinally striated. Corpus bursae often long, membranous, anteriorly expanded, bearing a signum. Signum elliptic, bar-shaped or, irregularly shaped, often with marginal spines (Jiang et al. 2011). 297

Remarks: In multi-gene analyses Biston is sister to Erannis (Jiang et al. 2017; Murillo-Ramos et al. 2019).

96. Biston strataria (Hufnagel, 1767) Phalaena strataria Hufnagel, 1767: Berlin Mag. 4 (5): 514 (Germany: Berlin region). Syntype(s). Phalaena (Geometra) prodromaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 99 (Austria: Vienna district). Syntype(s). Phalaena (Geometra) marmorata Sulzer, 1776: Geschichte Insecten 1: 161; ibidem 2: pl. 23, fig. 1 (Switzerland). Syntype(s). Phalaena vernalis Fourcroy, 1785: Entomologica Parisiensis 2: 264 (France: Paris). Syntype(s). Amphidasys prodromaria meridionalis Oberthür, 1913: Études Lépid. comp. 7: 251, pl. 163, fig. 1591 (France: Digne). Holotype ♂. Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Amphidasys prodromaria nigricans Oberthür, 1913: Études Lépid. comp. 7: 251, pl. 163, fig. 1592 (Poland; Czech Republik: Silesia). Holotype ♀. Biston strataria hasegawai Inoue, 1955: Tinea 2 (1/2): 86 (Japan: Hokkaido, Teshio, Kamikawa-­ gun, Asashi-mura). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): albata: Tautel (1993) (f.); albescens: Lempke (1970) (f.); crassestrigata: Lempke (1970) (f.); extrema: Lempke (1970) (f.); iaspidea: Leraut (2009) (f.); latefasciata: Lempke (1970) (f.); melanaria: Koch (1949) (f.); robinaria: Fringes (1898) (ab.); terraria: Weymer (1878) (ab.).

External characters and abdomen: Wingspan 38–50 mm. Forewing: Ante- and postmedial lines black, dentate, medial line more straight, often reduced to a black costal spot. Basal, medial and apical area whitish, densely speckled with black. Antemedial line proximally, postmedial line distally accompanied by a reddish brown band. Hindwing: whitish grey, with postmedial line black, slightly waved, accompanied by a reddish brown band, medial line thicker, sometimes lacking. Pattern of female more contrasted. Discal spot of forewing a short black line, on hindwings usually absent, sometimes very inconspicuous. Fringes of ground colour, chequered with black spots. Frons white, collar white with some black scales, posteriorly with black ring. Tegula whitish, speckled with black, posteriorly a black line, thorax darker. Antennae bipectinate with noticeable reddish brown branches and speckled shaft in ♂, filiform, black with white dots in ♀. Hindtibia with 2 x 2 spurs. Abdomen brownish to grey, speckled with darker scales. Variation: Variable species. Ground colour of forewings can be lighter or darker, extreme (non-melanic) forms monochrome reddish brown with black lines, speckled with darker scales. Melanic forms occurring locally, with unicolorous black wings and body except white frons, usually with inconspicuous lighter shadows on the terminal area of both wings. Taxon ‘meridionalis’ described from southern France (Digne) and mentioned from several localities in the Mediterranean, variable in external appearance with wings usually paler, white sprinkled, with light brown bands, showing clinal character transitions towards nomino­ typical populations. Male genitalia: Uncus triangular, weakly setose, wide at base, margin slightly concave, apex usually with rounded double tip, about four-fifths as long as basal width. Gnathos arms fused ventrally, folded, margin serrate. Juxta long, triangular, proximal third slightly constricted, in centre wider, often significantly, then rapidly narrowing. Apex forming a hook (see Biston rosenbaueri and B. achyra). Valva broad, simple, membranous, approximately 298

showing same width basally and apically. Valva bears long setae from centre to apex. Sacculus nearly straight. Saccus wide, rounded, medially with short, often bent trapezoid projection. Aedeagus narrow, slightly bent, apical 1.2–1.45 mm fluted, without cornutus. Length of aedeagus: 2.8–3,1 mm (on average 3.0 mm). For differential features from B. rosen­ baueri and B. achyra see table 5. Table 5: Differential features in male genitalia of B. strataria, B. rosenbaueri and B. achyra.  

 

B. strataria

B. rosenbaueri

B. achyra

Juxta      

posteriorly above centre lateral margins base width (mm)

beak-shaped, pointed thickened concave 0.5 to 0.63

slightly bent, not pointed not thickened nearly straight 0.5 to 0.6

slightly bent, not pointed not thickened nearly straight 0.4 to 0.5

Saccus

anteriorly

medially with short, trapezoid projection

medially with short, ­trapezoid projection

rounded

  Aedeagus  

  total length (mm) striated part (mm)

2.77 to 3.14 1.23 to 1.45

3.05 to 3.2 1.35 to 1.56

  2.56 to 2.69 0.96 to 1.11

Female genitalia: Papillae anales membranous, slightly setose. Apophyses posteriores very long, on average 7 mm, apophyses anteriores shorter (half length), both ending swollen, bent, anteriorly usually membranous. Lamella antevaginalis membranous, triangular, lamella postvaginalis oval, sclerotized. Ostium bursae and colliculum wide. Corpus bursae often long, membranous, posteriorly an irregularly sized sclerotized structure with small teeth, anteriorly expanded. Signum developed as sclerotized, narrow plate. Distribution and abundance: West Palaearctic. In Europe widely distributed from the Ural mountains to Ireland and Portugal and from southern central Fennoscandia to the Mediterranean including Corsica, Sardinia, Sicily and Crete. – Common or rather common, usually found in limited numbers in the north, but sometines quite abundant in southern Europe. – Outside Europe in North Africa (Morocco, Algeria and Tunisia), in the east from Turkey, across Transcaucasus, south-western Siberia to the Altai region. Also in Japan (subsp. hasegawai) (Inoue et al. 1982). Phenology: Univoltine: In Denmark from late March to early May. In south-western Germany from early March to mid- or late April (Ebert 2003). In Italy from March to April or May, depending on altitude (Flamigni et al. 2007). Caterpillars in southern Scandinavia from late May to July, further to the south slightly earlier. Overwintering as pupa. Adults active at night, males readily attracted to light, females rarely (restricted to few females very early in the evening). Females also found on Quercus trunks during daytime (P. Sihvonen own data). Biology: Larva polyphagous. Found on Populus tremula, Salix alba, S. purpurea, S. ­caprea, Quercus robur, Q. rubra, Ribes uva-crispa, R. nigrum, Rubus idaeus, Prunus spinosa, Acer campestre and Lonicera xylosteum (Germany: Ebert 2003), on Prunus spinosa (north-­ western Italy: King & González-Estébanez 2015), on Ulmus, Corylus avellana and Alnus (U.K.: Skinner 2009) and on Pyrus spinosa and Platanus orientalis (Samos: Fritsch et al.

299

2014). In southern Scandinavia the main food-plant appears to be Quercus. Recorded also on Tilia (Redondo et al. 2009). Habitat: Silvicolous. In a wide variety of deciduous forests on both dry and more wet soil, parks and gardens. In southern Scandinavia at low altitudes, in south-western Germany usually up to 700 m (Ebert 2003), in the Alps up to 1500 m (Forster & Wohlfahrt 1981) and in Sicily to 1560 m (Flamigni et al. 2007). Similar species: Biston rosenbaueri, B. achyra. Genetic data: BIN: BOLD:AAB4693 (n=59 from Finland, United Kingdom, Netherlands, Germany, Austria, Switzerland, France, Spain, Italy, Greece, including Crete, Morocco, Tuni­sia, north-eastern Turkey, Russia (Urals), Turkmenistan). Genetically homogeneous, but North African populations slightly diverging (0.6%), one barcoded specimen from Turkmeni­stan diverging by 1.0%. Nearest species: B. achyra from Israel (3.3%).

97. Biston rosenbaueri Müller, 2018 Biston rosenbaueri Müller, 2018: Nota lepid. 41 (2): 207–213, figs 1–2, 5–6 (Eastern Greece, Thrace, Avantas). Holotype ♂ ZSM; gen.prp. 1033 B. Müller).

External characters and abdomen: Wingspan 40–45 mm. Abdomen, thorax, tegula appearing furry. Forewing: Ground colour white, densely irrorated with light and dark brown, sometimes also black scales, together giving a pale grey or brownish impression. Veins brown or black. Post- and antemedial lines distinct, black, on veins thickened, forming small, medially directed tips. Postmedial line undulating, angled on veins M3 and M2. Antemedial line bent, sharp or more diffuse, sometimes angled on M3 and A. Medial line indistinct. Postmedial line on outer margin, antemedial line on inner margin developed as a broad pale brown band, in darker specimens indistinct. Terminal area with a diffuse whitish line, sometimes 300

absent. Discal spot elongate, distinct, slightly blurry, between postmedial and medial line. Hindwing less irrorated, therefore paler. Postmedial line of hindwing nearly straight, medial line diffuse, straight. Discal spot a distinct dot. Both wings with fringes weakly chequered brownish and grey. Labial palpi with furry scaling, medium brown, frons roughly scaled, whitish. Patagia anteriorly whitish, posteriorly black. Tegulae whitish, irrorated with more or less black scales. Thorax greyish brown, posteriorly concolorous with tegulae. Abdomen beige and grey scaled. Antennae of males bipectinate with reddish brown, ciliated branches and speckled shaft. Hindtibia with 2 + 2 spurs. Female unknown. Variation: Known males vary in degree of dark speckling. Male genitalia: Similar to those of Biston strataria and B. achyra. Differing from B. strataria in the shape of the juxta and from B. achyra in the larger aedeagus. Valva as in B. strataria. Juxta tapering evenly, without distinct constriction, not wider at centre (see above, table 5). Tip of juxta pointed, sometimes very little bent, not developed as distinct hook. Aedeagus as in B. strataria. For differential features from B. strataria and B. achyra see table 5. Distribution and abundance: (South-east) European. Endemic to Balkan peninsula, so far known from northern Croatia and northern Greece. – Apparently local and rare.

Phenology: Problably univoltine, scarce data from late February to mid-April. Biology: Unknown. Habitat: Scarce data available. In north-western Greece in open Quercus forest, mainly with Q. cerris, on a south facing slope (F. Rosenbauer pers. comm.) and mosaic of deciduous xerothermic Quercus woodland and dry grassland on limestone (Tóth et al. 2013). Vertical distribution from about 200 up to 1000 m. 301

Similar species: Biston strataria, B. achyra. Separation of males by external markings from B. achyra is uncertain. B. achyra usually with brownish grey patagia, tegulae and thorax and without white blotches. For differences in genitalia see table 5. Strongly diverging in the DNA barcode (see below). Genetic data: BIN: BOLD:ADA6153 (n=6 from Croatia: Krk, Greece: Epirus). Genetically homogeneous. Nearest species: B. achyra (4.1%).

98. Biston achyra Wehrli, 1936 Biston achyra Wehrli, 1936: Mitt. münchn. ent. Ges. 26: 35, pl. 2, fig. 3 (Turkey: Achyr Dağh [Maras]; Yüksek Dağh, Amanus). Syntypes 3♂. Biston achyra engonia Wehrli, 1936: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 430, pl. 35: h (Turkey: Amanus Mts., Yüksek Dagh). Syntype(s) ♂.

External characters and abdomen: Generally as in B. rosenbaueri. However, patagia, tegulae and thorax brownish grey and without white blotches. Separation of males by external markings from B. rosenbaueri is uncertain. Female unknown. Variation: Males vary in degree of dark speckling. Male genitalia: Similar to those of Biston rosenbaueri and B. strataria. Differing from both species in the length of the aedeagus. Valva as in B. strataria. Juxta as in B. rosenbaueri, i.e. evenly tapering, without distinct constriction, not wider at centre. Tip of juxta pointed, sometimes very little bent, not developed as a distinct hook (see B. strataria). Structure of aedeagus as in B. strataria and B. rosenbaueri but shorter. Length only 2.56–2.69 (on aver­ age 2.63) mm. For differential features from B. rosenbaueri and B. strataria see table 5. Female genitalia: Unknown. Distribution and abundance: Mediterranean-Turanian. In Europe only known from the Greek island Samos (Fritsch et al. 2014). Earlier reported from Crimea (Viidalepp 1996), but according to Kostjuk (2004), the specimen was misidentified. – Poorly known species, apparently rare. – Outside Europe known from Turkey, northern Caucasus, Georgia, Azerbaijan, Kopet Dagh in Turkmenistan (Viidalepp 1996), Israel and Jordan. Phenology: Univoltine: From early March to late April, depending on latitude and altitude, in Samos recorded from early to late March and in early April (D. Fritsch pers. comm.). Perhaps even later at the highest altitudes. Larval time unknown, but two caterpillars found in Turkey in mid-May (M. Leipnitz pers. comm.). Early flight period suggesting hibernation as pupa. Males active at night, attracted to light. Female rate extremely low; so far, no females found in Europe. Biology: Larva possibly monophagous. Found on Quercus, probably Quercus coccifera (M. Leipnitz pers. comm.). Habitat: Probably silvicolous. In Samos often in cultivated areas, but surrounded by rocks and extremely rich vegetation. In Turkey in open landscapes with shrubbery of Quercus, Rhamnus and Crataegus (M. Leipnitz pers. comm.). From sea-level up to 500 m in Samos (D. Fritsch pers. comm.), in Turkey up to 2200 m. Similar species: Biston strataria, B. rosenbaueri.

302

Genetic data: BIN: BOLD:ABZ6431 (n=7 from Greece (Samos), Turkey, Israel, Lebanon). Genetically heterogeneous, Israeli populations diverging from the nominotypical populations from Samos and southern Turkey by 2.2%. Nearest species: B. rosenbaueri near achyra (4.1%).

99. Biston betularia (Linnaeus, 1758) Phalaena (Geometra) betularia Linnaeus, 1758: Syst. Nat. (Ed. 10) 1: 521. Syntype(s) (LSL). Phalaena (Noctua) p-graecum Poda, 1761: Insecta Musei Graecensis: 90. Syntype(s). Phalaena marmoraria Sepp, 1792: Nederlandsche Insecten 2: pl. 10, 11 (Netherlands). Syntype(s). Phalaena (Geometra) ulmaria Borkhausen, 1794: Natur. eur. Schmett. 5: 181 (Europe). Syntype(s). Amphidasis cognataria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 208 (North America). Holotype ♀ (USNM). Valid at subspecific rank (Scoble 1999), see Remarks. Amphidasys betularia var. doubledayaria Millière, 1870: Iconogr. Descr. Chenilles Lépid. inédits 3 (25): 117, pl. 111, fig. 1 (England). Syntypes, including 3♂4♀. Biston betularius var. fumosarius Alphéraky, 1897: in Romanoff, Mém. Lépid. 9: 140 (Transcaucasia; Tachien-lu [Kangding]). Syntype(s). Valid at subspecific rank (Scoble 1999). Biston robustum var. parva Leech, 1897: Ann. Mag. nat. Hist. (6) 19: 323 (China: Pu-tsu-fong; Che-tou; Tachien-lu [Kangding]). Syntypes 1♂2♀ (NHMUK). Valid at subspecific rank (Scoble 1999). Amphidasis cognataria fortitaria Barnes & McDunnough, 1917: Contr. nat. Hist. N. Am. 3 (4): 246, pl. 26, fig. 2 (U.S.A.: Colorado, Garfield County, Glenwood Springs). Lectotype ♂ (USNM), see Remarks.

303

Amphidasis cognataria f. swettaria Barnes & McDunnough, 1917: Contr. nat. Hist. N. Am. 3 (4): 246, pl. 26, fig. 6 (U.S.A.: Pennsylvania, Beaver County, New Brighton). Lectotype ♂ (USNM), see Remarks. Amphidasis cognataria contrasta Barnes & Benjamin, 1923: Lepidopterist 4 (2): 12 (U.S.A.: Utah, Eureka). Holotype ♂ (USNM). Valid at subspecific rank (Scoble 1999), see Remarks. Lycia cognataria var. mesle Forbes, 1928: in Leonard, Mem. Cornell Univ. agric. Exp. Stn 101: 603 (U.S.A.: New York, Queens County, Glendale). Syntype(s), see Remarks. Biston cognataria alexandrina Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 432, pl. 36: a ([Kyrgyzstan]: Alexander Mts.). Syntypes 4♂ (ZFMK). Raised from synonymy (Scoble 1999) back to subspecific rank, based on divergence of COI gene, see Genetic data. Biston cognataria coreae Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 432, pl. 36: a (Korea: Seishin-Olto; Poktussan, 2500 m). Syntypes 10♂. Valid at subspecific rank (Scoble 1999), see Remarks. Biston cognataria sinitibetica Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 433, pl. 36: a (West China: Tachien-lu [Kangding]). Syntypes ♂♀. Synonym of subsp. parva (Scoble 1999). For potential validity of taxon cognataria at species rank see Remarks. Biston betularia nepalensis Inoue, 1982: Bull. Fac. domest. Sci. Otsuma Wom. Univ. 18: 175, figs 40a, b (Nepal: Tukcha, near Daulagiri). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999), confirmed by Jiang (2011). Eurbyjodonta concinna Warren, 1899: Novit. zool., 6: 50. Holotype ♂ (Kazakhstan?: Ili district). (NHMUK) (Synonymized by Wehrli (1941), treated as a synonym of Biston huberaria by Scoble (1999), synonymized with B. betularia by Jiang et al. (2011). Amphidasis huberaria Ballion, 1866: Horae Soc. ent. ross., 4: 29, pl. 1, fig. 1. Syntype(s), Russia: Western Siberia, near Omsk. (ZISP). Synonymized by Vijdalepp (1979) and Jiang et al. (2011). Biston (Eubyjodonta) huberaria tienschana Wehrli, 1941: in Seitz, Gross-Schmett. Erde, 4 (Suppl.): 435, pl. 36: d, g. Holotype ♂, China: Xinjiang, Ürümqi, Tian-shan. (ZFMK). Treated as a synonym of Biston huberaria by Scoble (1999), synonymized with B. betularia by Jiang et al. (2011). Unavailable names (infrasubspecific): benesignata: Lempke (1952) (f.); brunnea: Lempke (1970) ­ empke (f.); carbonaria: Jordan (1869) (ab.); centristrigata: Lempke (1970) (f.); destrigata: L (1952) (f.); fasciata: Prout (1915) (ab.); funebraria: Lambillion (1905) (ab.); insularia: Thierry-­ Mieg, (1886) (ab.); nigromarginata: Lempke (1952) (f.); ochrearia: Mansbridge (1908) (ab.); variegata: Lempke (1970) (f.).

External characters and abdomen: Wingspan ♂ 40–51 mm, exceptionally only 36 mm; ♀ 38–55 mm. Ground colour whitish to brownish grey, densely speckled with black scales. Forewing transverse lines black, forming distinct streaks on costa. Postmedial line distinct, slightly wavy. Medial and basal lines thicker at costa, posteriorly sometimes inconspicuous or fading. Antemedial line bent, distinct to inconspicuous, serrate. Hindwing with post­medial and medial line. Postmedial line protruding outwards on vein M2 on both wings. Terminal line absent. Discal spots on both wings elongate. Fringes of ground colour, chequered with black spots. Frons white, collar, tegula, thorax, abdomen concolorous with wings. Collar posteriorly black. Antennae bipectinate and speckled shaft in ♂, filiform, black with white dots in ♀. Hindtibia with 2 x 2 spurs.

Variation: Highly variable species in relation to degree and extent of black speckling. ­Melanic specimens are monochromous black with darker veins on wings, black thorax and abdomen (ab. carbonaria), wings often irrorated with scattered white scales. The intermediate form (ab. insularia) is genetically not related to ‘carbonaria’ (Skinner 2009). The frons of all forms remains white. Often the wing roots are white, too. See Remarks below.

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Male genitalia: Uncus trapezoid, wide at base, margin concave, with rounded double tip. Gnathos arms fused ventrally, forming a wide, slightly rounded serrate plate. Juxta wide, nearly rectangular plate, anteriorly less narrow, posterior end flat. Valva evenly curved, wide at base, apex setose. Medial ridge weakly sclerotized, widely spined over half of valva. Costa narrow, sclerotized. Saccus wide, rounded. Aedeagus long, stout, vesica with two groups of cornuti, one arranged in a band, other triangularly. Female genitalia: Papillae anales elongate, membranous, slightly setose. Apophyses posteriores very long, on average 5.3 mm. Apophyses anteriores little less than half length of apophyses posteriores (on average 2.5 mm), both ending swollen, bent, tip usually membranous. Ductus bursae funnel-shaped, sclerotized, striated, mid-section and anterior end constricted. Corpus bursae membranous, constricted between posterior and anterior part. Posterior part funnel-shaped with many striae (partially scrambled). Anterior part pearshaped. Signum arcuate, both sides with diffuse margins. Distribution and abundance: Palaearctic. In Europe from the Urals to Ireland and Portugal and from central Fennoscandia to the northern half of the Iberian peninsula, France incl. Corsica, Italy incl. parts of Sicily and the Balkan peninsula except most of Greece. – Common, found nearly everywhere, usually in limited numbers. – Outside Europe in Turkey, Georgia, Armenia, Azerbaijan (subsp. fumosarius) and central Asia (subsp. alexandrina) across southern Siberia (subsp. sibiricus) to China, Far East Russia, Korea, Sakhalin (subsp. coreae) and Japan (subsp. parvus). Recorded also in Nepal (subsp. nepalensis) and North America (subsp. cognataria), for classification of the latter see Remarks.

Phenology: Univoltine, occasionally bivoltine: In northern and central Europe from midor late May to early August. In Italy according to Flamigni et al. (2007) in two generations from May to July and from August to September. In the Iberian peninsula from late April 305

to mid-August and regarded as bivoltine (Redondo et al. 2009). Flight period very long, requiring further study to examine if really referring to two generations in Italy and Spain. In Portugal from May to September, either univoltine or with partial second generation, peaking in June and especially in July, with much less specimens in August and very few in September (M. Corley pers. comm.). Caterpillars from July to early October. Hibernation as pupa. Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Populus tremula, P. x canadensis, Salix fragilis, S. viminalis, S. cinerea, S. caprea, Juglans regia, Carpinus betulus, Corylus avellana, Betula pendula, B. pubescens, Alnus glutinosa, Fagus sylvatica, Quercus robur, Urtica dioica, Platanus, Bergenia crassifolia, Sorbus aucuparia, Rubus idaeus, R. fruticosus, Cotoneaster, Crataegus monogyna, Rosa, Prunus domestica insititia, P. spinosa, Sanguisorba officinalis, Cytisus scoparius, Medicago sativa, Robinia pseudoacacia, Acer, Rhamnus, Frangula alnus, Tamarix gallica, Cornus sanguinea, Hippophae rhamnoides, Buddleja davidii, Fraxinus, Arte­misia vulgaris (Germany: Ebert 2003) and on Pyrus communis (north-western Italy: King & González-Estébanez 2015). Habitat: Silvicolous. In most European habitat types as long as deciduous trees and bushes are present, thus absent from open habitats and from the highest mountains. Absent also from hot habitats close to sea-level in the Mediterranean area. In central Europe from sea-level up to 1300 m (Ebert 2003), in the Alps up to 1600 m (Kerschbaum & Pöll 2010), in Spain up to 2000 m (Redondo et al. 2009), in central Asia up to 2200 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:ABY9367 (n=96 from 14 countries from Finland to southern Italy and from Portugal and United Kingdom to Russia (Altai) and Mongolia in the east, further n=255 from Asia and North America). Genetically comparatively homogeneous in Europe, but with heterogeneous patterns in Asia and North America (partly BIN-sharing), requiring further study. The majority of North American populations (subsp. cognataria) BIN-sharing but slightly diverging from European populations (1.3%). One DNA barcode from Kyrgyz­ stan (subsp. alexandrina) diverging by 2.1%. Nearest species in Europe: B. strataria (7.2%). Remarks: Melanic forms were first observed in industrial areas in England in the middle of the 19th century, later also elsewhere (‘industrial melanism’). The ratio of melanic and non-melanic forms can vary considerably. There are many publications about this phenomenon. History and distribution see Wehrli (1939–1954), selective forces and dominance see Mikkola (1984a; 1984b). The taxon cognataria, described from North America, was subordinated at subspecies rank under B. betularia in Rindge (1975), Covell (1999), and Handfield (1999). Pohl at al. (2010: 383) in their fauna of Alberta conclude, based on mtDNA analysis, that American and European populations belong to different species, Albertan populations belonging to ‘more than one species’. However, in the most recent checklists (Pohl et al. 2016; 2018) the taxon was again subordinated under B. betularia at subspecies rank. The European BIN (BOLD: ABY9367) is shared by the majority of North American populations. Former entomologists (e.g. Wehrli 1941) reported ‘B. cognataria’ at species rank from Asia. Recent publications do not mention this taxon for Asia (Viidalepp 1996; Kim et al. 2001; 2017; Nakajima 2011). In a recent revision of the genus Biston (Jiang et al. 2011), cognataria is not listed in the synonymy of betularia but in the eastern Palaearctic region no differential features in genitalia were found which would justify the discrimination of two different species (Jiang pers. comm.). 306

Erannis Hübner, 1825 Erannis Hübner, [1825] 1816, Verz. bekannter Schmett.: 320. Type species: [Phalaena] ­defoliaria Clerck, 1759. Genus synonyms after Scoble (1999). Hybernia Berthold, 1827, in Latreille, Nat. Familien des Thierreichs: 485. Type species: [Phalaena] defoliaria Clerck, 1759. Lampetia Stephens, 1827, in Anonymous, Retrospective Review (2) 1: 242. Type species: [Phalaena] defoliaria Clerck, 1759. Chimadia Speyer, 1839, Isis, Leipzig. 1839: 109. Type species: [Phalaena] defoliaria Clerck, 1759. ‡ Thriptera Gistl, 1848, Naturg. Thierreichs: xi. Type species: [Phalaena] defoliaria Clerck, 1759. [Junior homonym of Thriptera Solier, 1836 (Coleoptera), and junior objective synonym of Erranis Hübner].

Diversity and distribution: The genus Erannis includes eleven species in the Palaearctic and in the Nearctic regions. External characters and abdomen: ♂ fully winged, wingspan 22–47 mm. Male forewing grey to sandy or reddish brown more or less speckled with dark scales. Postmedial and antemedial lines usually distinct, rarely reduced or absent. Hindwing lighter. Females with short wing stumps, abdomen grey or yellowish brown with rows of double dots or lines. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Most species are variable. Male genitalia: Uncus and gnathos well developed, gnathos ventrally fused. Valva with short ventral arm, apex rounded, setose. Sacculus curved, setose. Costa with medial process (missing in E. declinans) bearing a group of spines. Saccus rounded, sometimes pointed. Aedeagus variable in size between species. Female genitalia: Apophyses posteriores long, apophyses anteriores shorter, both spatulate at tip. Ductus bursae wide, sclerotized. Corpus bursae long, membranous, slender, slightly striated. Details of European species see below. Remarks: In multi-gene analyses Erannis is sister to Biston (Jiang et al. 2017; Murillo-Ramos et al. 2019).

100. Erannis defoliaria (Clerck, 1759) Phalaena defoliaria Clerck, 1759: Icones Insectorum Rariorum 1: pl. 7, fig. 4 (Europe, Sweden). Lectotype ♂ (NHRS). ‡ Phalaena pulveraria Hufnagel, 1767: Berlin Mag. 4 (5): 518 (Germany: Berlin region). Syntype(s). Junior primary homonym of Phalaena pulveraria Linnaeus, 1758. Phalaena Geometra discolor Ström, 1783: Nye Saml. K. dansk. Vid. Selsk. Skr. 2: 82 (Europe). Syntype(s). Phalaena bistrigaria Giorna, 1791: Calendario Entomologico: 131 (Torino?). Syntype(s). Hybernia defoliaria suffusa Cockerell, 1889: Entomologist 22: 55. Syntype(s). Hybernia defoliaria teriolensis Dannehl, 1933: Ent. Z., Frankf. a. M. 47 (17): 139 (Austria: Tyrol). Syntype(s). Validated at subspecific rank by Scoble (1999), downgraded to synonymy in Leraut (2009), here confirmed based on DNA barcode patterns, clinal character transitions and absence of constant and significant differential features. Unavailable names (infrasubspecific): albescens: Schawerda (1914) (ab.); brunnescens: Rebel (1910) (ab.); costafumata: Leraut (2009) (f.); crocinea: Leraut (2009) (f.); decorata: Leraut (2009) (f.); depuncta: Lempke (1952) (f.); destrigata: Gornik (1932) (ab.); deumbrata: Lempke

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(1952) (f.); eutaeniaria: Schawerda (1916) (ab.); fasciata: Lempke (1952) (f.); fumosa: Lempke (1970) (f.); fuscosignata: Heinrich (1916) (ab.); holmgreni: Lampa (1885) (ab.); lutescens: Leraut (2009) (f.); nigra: Bandermann (1928) (ab.); obscura: Helfer (1896) (ab.); obscurata: Staudinger (1901) (ab.); obscurata-progressiva: Haverkampf (1906) (ab.); rufescens: L ­ eraut (2009) (f.); suffusa: Cockerell (1886) (ab.); tangens: Cockayne (1942) (ab.); transitoria: L ­ empke (1952) (f.); trifasciata: Schawerda (1914) (ab.); unistrigata: Lempke (1952) (f.); ustulata: Leraut (2009) (f.).

External characters and abdomen: Male: Wingspan 29–42 mm. Forewing of nomino­typical form with pale sandy ground colour, often with yellow to reddish brown tinge, speckled with more or less fine dark scales. Wavy line weak, post- and antemedial lines black, usually distinct. Postmedial line angled on M2 and between CuA2 and A, antemedial line only between CuA2 and A. Medial area constricted. Postmedial line shadowed by broad brown band. Terminal area often with row of dark blotches towards costa. Fringes chequered brownish and pale sandy. Terminal line absent. Discal spot dark, conspicuous. Hindwing pale grey, with more or less dark freckling, fringes weakly chequered or unicolorous, discal spot sometimes less conspicuous. Frons, vertex, thorax and tegula with long hairy scales, pale sandy to reddish brown, collar reddish brown ring. Abdomen concolorous with thorax, with dark freckling. Antennae bipectinate, branches becoming shorter towards tip. Female: Nearly wingless, wing stumps very short (less than 1 mm), with black and white hairy scales. Frons blackish brown, vertex whitish. Collar and thorax whitish with blackish brown blotches. Abdomen yellow-grey, dorsally with double row of black dots, sometimes forming double line. Laterally also with dark blotches, often fused into a line. Antennae filiform, black and white ringed as legs are. Hindtibia of both sexes with 2+2 spurs. Variation: Males extremely variable. In addition to the nominotypical form described above, there are rusty brown forms with conspicuous forewing lines. The transverse lines can also be reduced, inconspicuous or absent. Forms with extended dark freckling are often observed. Discal spot on hindwing sometimes weak or absent. Because of the high variability a large number of individual forms are described and many unavailable infrasubspecific names exist. Male genitalia: Uncus triangular, lateral margins convex, apex with rounded tip. Gnathos ventrally fused, triangular, apex rounded. Uncus and gnathos sclerotized. Valva wide and short, setose. Costa with sclerotized curled process, bearing large comb of spines. Apex rounded, setose. Ventral arm weakly sclerotized, submarginally with long setae. Juxta wide plate with rounded apex, lateral margins concave. Saccus rounded. Aedeagus broad, apex tapered. Vesica with 3–4 spines. Female genitalia: Papillae anales oval, membranous, soft, slightly setose. Apophyses posteriores long (on average 2.5 mm), apophyses anteriores shorter (on average 1.2 mm), both spatulate at tip. Ductus bursae as long as wide, sclerotized. Ostium bursae with proximally directed flap. Corpus bursae long, membranous, slender, slightly striated, without signum. Lamella antevaginalis sclerotized, alate, laterally partially serrate, round at tip. Lamella postvaginalis a triangular small plate with rounded corners, both fused. Distribution and abundance: West Palaearctic. In Europe from the Urals to westernmost Europe and from central Fennoscandia to the Balkan peninsula (including Crete but absent from most of Greece), Italy including Sardinia and Sicily, France including Corsica and the northern half of the Iberian peninsula with isolated occurrence further south. In Iceland one single specimen on the south coast. – Common or very common, often occurring in numbers. 308

Sometimes mass occurrences with millions of caterpillars defoliate large numbers of trees. – Outside Europe in Georgia, Armenia, Azerbaijan, Kazakhstan, northern Iran and Turkey. Also in North Africa in Morocco and Algeria. Phenology: Univoltine: In southern Scandinavia from early October to mid- or late November, in south-western Germany from early October to early December (Ebert 2003), in Italy from October to December (Flamigni et al. 2007) and in Spain from October to February (Redondo et al. 2009). Caterpillars from late April to early or mid-June. Hibernation as egg or as adult. Males active at night, readily attracted to light. Females found at night on tree trunks or branches. Biology: Larva polyphagous. Found on Populus tremula, P. nigra, P. trichocarpa, Salix alba, S. viminalis, S. cinerea, S. caprea, Juglans regia, Carpinus betulus, Corylus avellana, Betula pendula, Alnus, Fagus sylvatica, Quercus, Ulmus, Pyrus communis, Malus domestica, Sorbus aucuparia, Crataegus, Rubus idaeus, Filipendula ulmaria, Rosa canina, R. vosagiaca, Prunus cerasifera, P. padus, P. avium, P. spinosa, Acer pseudoplatanus, A. campestre, Tilia platyphyllos, Cornus sanguinea, Vaccinium myrtillus, Ligustrum vulgare and Sambucus ebulus (Germany: Ebert 2003). Found on Crataegus also in north-western Italy (King & González-Estébanez 2015). Pupation in the ground. Habitat: Silvicolous. Due to the wide variety of food-plants the species occurs in a wide variety of open habitats as well as deciduous forests, gardens and parks. From sea-level, in south-western Germany up to 800 m (Ebert 2003). In southern Europe up to 1600 m. Similar species: Females of E. ankeraria with different structure of genitalia. Genetic data: BINs: BOLD:AAB4418 (n=69 from Finland, Netherlands, United Kingdom, Germany, Austria, Czech Republic, Hungary, France, Italy, Bulgaria, Greece, Tunisia, Turkey); BOLD:ABZ2920 (n=1 from Tunisia), at a distance of 2.2% (pseudogene?). ­Genetically 309

somewhat heterogeneous in the main cluster, maximum variation 1.4%. Nearest species: Erannis kashmirensis László, 2003 from Pakistan (2.1%), E. jacobsoni (Djakonov, 1926) from Altai Mountains (2.2%).

101. Erannis ankeraria (Staudinger, 1861) Hibernia ankeraria Staudinger, 1861: Stettin. ent. Ztg. 22: 292 (Hungary: near Pest). Syntypes 2♂. Erannis ankeraria syriaca Wehrli, 1934: Mitt. münch. ent. Ges. 24 (2): 41 (Syria: SE Taurus [Toros Dağları], Akbés [Maydan Ikbis]; Turkey: Malatia [Malatya]). Syntypes 2♂. Valid at subspecific rank (Scoble 1999), here confirmed. Erannis ankeraria bervaensis Jablonkay, 1965: Folia ent. hung. (N.S.) 18: 522 (Hungary: Berva valley). Holotype (DIBE). Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features.

External characters and abdomen: Male: Wingspan 30–37 mm. Forewing light beige, more or less speckled with beige scales, which are a little darker than the ground colour. Post- and antemedial lines usually weak, pale brown. Postmedial line angled on M1 and Cu1. Antemedial line curved. Medial, subterminal and terminal lines absent. Discal spot weak. Fringes of ground colour. Hindwing creamy white, without lines. Frons, collar, thorax and abdomen of ground colour. Female: Nearly wingless, wing stumps very short, length about 1 mm with long scales. Frons black, vertex white, patagium, thorax and abdomen white with black blotches. Antennae filiform. Hindtibia of both sexes with 2+2 spurs. Variation: Little. In Bükk mts. in Hungary described (‘f. bervaensis’) as smaller, male forewings with more brownish scales, hindwings pure white. Male genitalia: Uncus wide, sclerotized, bilobed with short, rounded process at centre of apex. Gnathos ventrally fused. Dorsal part of valva apically widened, apex strongly setose, ball-shaped. Centre of valva with arcuate, long and pointed harpe. Juxta tapered, apex rounded. Saccus anteriorly compressed. Aedeagus cylindrical, apex pointed, caecum slightly bent, rounded, without cornutus. Female genitalia: Papillae anales membranous, soft, slightly setose. Apophyses posteriores long (on average 3.9 mm), apophyses anteriores shorter (on average 2.2 mm), both ending spatula-like and longer than in E. defoliaria. Ductus bursae as long as wide, compared with the latter weakly sclerotized. Ostium bursae wide. Corpus bursae long, membranous, slender, slightly striated, without signum. Lamella antevaginalis sclerotized, alate, with smaller ‘wings’ than in E. defoliaria. Lamella postvaginalis a triangular small plate with rounded corners, both lamellae fused. Distribution and abundance: South-east European-Anatolian. In Europe only in the southeast with a scattered occurrence in Italy, Slovenia, Hungary, Romania, Croatia, Serbia, Macedonia and Greece. Erroneously recorded for Austria by Leraut (2009) (N. Pöll pers. comm.). – Rare and local, usually found as single specimens. – Outside Europe in Turkey, northern Iran, Armenia (subsp. syriaca) and Turkmenistan. Phenology: Univoltine: From late February to late March (B. Tóth pers. comm.), in Bulgaria from mid-February to mid-March (S. Beshkov pers. comm.). Larva in May and June (Fajčik 2003). Overwintering as pupa. Males active at night, attracted to light. Females found on tree trunks or branches. 310

Biology: Monophagous. Larva recorded on Quercus (Fajčik 2003). Pupation in the ground, usually close to the tree where the caterpillars have been feeding and it seems to be important that there is grass around the tree, and not just dead Quercus leaves (S. Gomboc pers. comm.). Habitat: Xerothermophilous. Preferring xerothermic, scattered Quercus forests in karst pastures, usually the so-called Illyrian neutrocline sessile oak-hornbeam forests. Requiring very open oak forest, with grassland around the trees (S. Gomboc pers. comm.). Occurring at relatively low altitudes from 130 m up to 700 m. Similar species: Agriopis aurantiaria. Male forewings of E. ankeraria narrower than in A. auran­ tiaria, hindwing without lines and discal spot. Females with wing stumps of max. 1 mm length, in aurantiaria about 2.5 mm. Genetic data: BIN: BOLD:AAF3006 (n=6 from Hungary, Bulgaria). Genetically homogeneous. Nearest species: Erannis kashmirensis László, 2003 from Pakistan (4.1%), E. defoliaria (5.5%), Remarks: Leraut (2009) – followed by Nahirnić & Beshkov (2016) – provisionally placed E. ankeraria in the Central Asian genus Desertobia Viidalepp, 1989, suggesting similarity in male genitalia. We confirm differences in the main structure of the male genitalia between E. ankeraria and other Erannis species but COI barcodes do not reflect such a relationship (see above). Until final (hopefully integrative) revision clarifies the matter we leave the taxon ankeraria in genus Erannis.

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102. Erannis declinans (Staudinger, 1879) Hybernia declinans Staudinger, 1879: Horae Soc. ent. ross. 14: 448 (Turkey: Amasia; Taurus). Syntypes 14♂4♀. One examined female (probably the other three as well) from the syntype series of Staudinger found to belong to Agriopis bajaria (BM). Erannis declinans tauriphila Wehrli, 1940: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 416, pl. 33: g (Turkey: Taurus [Toros Dağları], Marasch [Maras]). Syntype(s).

External characters and abdomen: Male: Wingspan 33–37 mm. Forewing: Ground colour pale to dark grey, speckled with more or less fine dark scales. Black postmedial line double, thickened at costa, lower 2/3 of length straight, slightly wavy towards costa. Basal line, if any, is reduced to a costal blotch. Medial and antemedial lines diffuse, often weak or absent. Costal margin with small black streaks. Wavy line whitish, often distinct, in darker specimens weak. Terminal line black, broken into black dots between veins. Fringes of ground colour with a dark longitudinal line. Hindwing pale grey with distinct dark grey discal spot. Antennae bipectinate. Female: Nearly wingless, wing stumps about 1 mm long. Hindtibia of both sexes with 2+2 spurs, frons smoothly scaled. Antennae filiform, whitish. Legs, frons, collar and thorax whitish, abdomen with indistinct black dots (see female of Agriopis bajaria). Variation: Male wing pattern is variable. Most specimens are poor in contrast, dark grey with weak lines. Contrasted forms can be colourful, reddish brown between postmedial and wavy line, medial area pale grey. Male genitalia: Uncus triangular, lateral margins weakly setose, apex slightly rounded. Gnathos ventrally fused, wide, tip spatulate. Valva at base wide, narrower at centre. Costa slightly setose, apex claviform, sclerotized, strongly setose. Valva partly membranous. Saccus unusually long, anteriorly curved. Juxta inconspicuous plate. Aedeagus very long, narrow. One part of apex pointed, other hook-shaped. No differences in male genitalia between Bulgarian, northern and eastern Turkish populations. Female genitalia: Papillae anales relatively small, membranous, soft, slightly setose. Apophyses posteriores long (on average 2.15 mm), apophyses anteriores shorter (on average 1.1 mm), spatulate at tip. Lamella postvaginalis a plate with two pointed, anteriorly directed projections. Antrum (posterior part of ductus bursae) large, funnel-shaped, sclerotized, anterior part of ductus bursae short, membranous. Corpus bursae posteriorly elongated, striated, partly sclerotized, anteriorly membranous, partly striated, round. Corpus bursae constricted between posterior and anterior part, with short diverticulum. Distribution and abundance: South-east European-Anatolian. In Europe only in Romania, Bulgaria and Greece. – Rather local, now and then abundant in its localities. – Outside ­Europe in Turkey, Armenia, Chechenya, Dagestan and northern Iran. Phenology: Univoltine: From early November to early December. In Bulgaria from October to January (S. Beshkov pers. comm.). Caterpillars from late April to mid-June (Ziegler 2019). Overwintering as egg. Males active at night, readily attracted to light, also found at midnight, resting in low vegetation (K. Nupponen pers. comm.). No records of females in nature traced. Biology: Larva oligophagous. Found on Paliurus spina-christi in northern Greece (Ziegler 2019) and on Cytisus in north-eastern Turkey (J. Gelbrecht pers. comm.). Pupation just below the surface of the ground.

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Habitat: Xerothermophilous. In northern Greece open mountain slopes with scattered bushes. Preferring lowlands, hills and promontories, from 160 m up to 740 m. Similar species: Female very similar to female of Agriopis bajaria, the latter differing by black collar (patagium), legs and antennae greyish. Double row of dots on abdomen of Erannis declinans usually reduced or lacking. Males without similar species in Europe. Genetic data: BINs: BOLD:AAL5951 (n=6 from Bulgaria), BOLD:AAF0208 (n=7 from eastern Turkey, Iran). Both BINs diverging by 2.5%, possibly suggesting status as different subspecies despite the lack of differences in male genitalia. In the COI gene closer to Erannis than to Agriopis species, e.g. Erannis ankeraria (5.5%), E. defoliaria (6.5%).

Kresnaia Leraut, 2009 Kresnaia Leraut, 2009, Moths of Europe: 122. Type species: Agriopis beschkovi Ganev, 1989.

Diversity and distribution: Kresnaia is a monotypic genus, the only species occurs in Europe and south-western Asia. External characters, abdomen and Genitalia: See species description below. Remarks: Differing from Agriopis and Erannis by its male genitalia structure with simple, apically rounded valva, long uncus and two groups of microcornuti in the vesica. Kresnaia has not been included in a multi-gene analysis so far. COI data suggest an isolated phylogenetic lineage for this genus (see under genetic data).

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103. Kresnaia beschkovi (Ganev, 1987) Agriopis beschkovi Ganev, 1987: Entomofauna 8 (18): 273, figured (south-western Bulgaria: Kressna gorge). Holotype ♂.

External characters and abdomen: Male: Wingspan 26−31 mm. Forewing narrow, ground colour grey, more or less speckled with dark grey scales. Post- and antemedial lines dark grey, usually distinct, sometimes weak, jagged. Postmedial line with light grey edging. ­Medial line diffuse, dark grey, usually straight. Terminal area with a straight, dark grey subterminal line, running to tornus, accompanied by a light grey edging. Hindwing paler with indistinct lines near costa. Terminal line on both wings dark grey. Antennae bipectinate. Female: Nearly wingless, wing stumps about 1 mm long. Antennae filiform, chequered brown and white. Both sexes with frons and vertex rough-scaled, these and thorax covered with a mixture of white and blackish grey scales. Abdomen brownish grey with indistinct blackish dots. Hindtibia of both sexes with 2 spurs. Variation: Males: little variation in distinctness of transverse lines. The pale grey edging on the postmedial and subterminal lines may be missing. Male genitalia: Uncus at base wide, long, apically slightly bent, tip round. Gnathos forming two ventrally non-fused bands. Valva simple, slender, weakly setose, at base wider, in centre slightly narrower, apex flattened, membranous with group of setae. Costa sclerotized, apically pointed. Saccus rounded. Juxta bifurcate, base trapezoid. Aedeagus narrow, curved. Vesica with two groups of small cornuti. Female genitalia: Papillae anales oval, membranous, soft, slightly setose. Length of apophyses posteriores about 3.5 mm, apophyses anteriores shorter (about 2.2 mm), both spatulate at tip. Ductus bursae short, posteriorly wider, membranous. Lamella antevaginalis strongly sclerotized, lunate. Lamella postvaginalis inconspicuous. Corpus bursae long, membranous. Posterior part narrow, long, striated. Anterior part oval with very small sclerotized triangular signum. Distribution and abundance: South-east European-Anatolian. Very limited distribution in south-western Bulgaria and in northern Greece. – In Europe an extremely local species, but in Kresna Gorge in Bulgaria relatively abundant with up to 20 specimens per night (Beshkov pers. comm.). – Outside Europe only known from western Iran where it can occur as a pest species defoliating the food-plants (Petschenka et al. 2006). Phenology: Univoltine: From mid-November to early December (Beshkov pers. comm.). In western Iran the larval time begins between early March and early April and lasts until midMay (Petschenka et al. 2006). Hibernation partly as adult or as egg. Males active at night and readily attracted to light. Females on branches of Quercus bushes up to 1.5 m above ground (Beshkov pers. comm.). Biology: Larva monophagous on Quercus. In western Iran found on Quercus brantii and Q. infectoria (Petschenka et al. 2006). Habitat: Xerothermophilous. On slopes with silicious rocks and stones where Juniperus excelsa, J. oxycedrus, Pistacia terebinthus, Paliurus spina-christi, Quercus pubescens, Q. virgiliana and Fraxinus ornus are growing. In western Iran in subtropical Quercus forests (­Petschenka et al. 2006). In Europe from 200 up to 585 m (Beshkov pers. comm.). In western Iran from 1100 up to 1750 m (Petschenka et al. 2006).

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Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAL6808 (n=5 from Bulgaria and Iran). Genetically homogeneous, exactly barcode-sharing between European and Iranian populations. Nearest species: North American Leucobrephos brephoides (Walker, 1857) (Archiearinae) (7.4%), genetic data not suggesting any closer relationship with any Palaearctic ennomine genus, hence the classification as a separate genus (Kresnaia) seems justified from a molecular view.

Agriopis Hübner, 1825 Agriopis Hübner, [1825] 1816, Verz. bekannter Schmett.: 320. Type species: Geometra leucophaearia Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Anisopteryx Stephens, 1827, in Anonymous, Retrospective Review (2) 1: 245. Type species: Geometra leucophaearia Denis & Schiffermüller, 1775. ‡ Amalthea Gumppenberg, 1887, Nova Acta Acad. Caesar. Leop. Carol. 49: 326. Type species: Geometra bajaria Denis & Schiffermüller, 1775. [Junior homonym of Amalthea Rafinesque, 1815 (Hymenoptera)]. Cryopega Dumont, 1925, Encycl. ent. (B, 3, Lepid.) 1 (1): 29. Type species: Cryopega legrasi Dumont, 1925.

Diversity and distribution: Six species in the Palaearctic region. External characters and abdomen: Males full-winged, females with strongly reduced wings. Male: medium-sized, wingspan approximately 22−38 mm. Wings grey, beige, yellowish, brown or mixture of these. Postmedial and antemedial lines usually distinct. Hindwing paler than forewing. Thorax and abdomen slender compared with size of wings. Female: Wing stumps present, short, except in A. marginata. Antennae partially bipectinate in males, filiform in females.

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Male genitalia: European species: Uncus and gnathos well developed, gnathos ventrally fused. Valvae simple, apex often with spines. Subventrally with spined or setose ridge. Juxta bifurcate or apically indented. Aedeagus with various shape, containing a spine-shaped cornutus (except in A. bajaria). Structure of valvae of sister genus Erannis more complex. Female genitalia: Papillae anales long, retractible, membranous, soft, slightly setose. Apo­ physes posteriores extremely long, apophyses anteriores shorter. Ductus bursae different between species, but simple, suitable for determination of the species. Corpus bursae membranous, usually ovate, containing a small blotch-shaped signum, missing in A. leuco­ phaearia. Remarks: Agriopis is a heterogeneous genus. Therefore the division of this genus into several subgenera was made or discussed in the past. According to Leraut (2009), of the European species only A. leucophaearia belongs to Agriopis (as type species), shifting A. bajaria to Cryopega Dumont, 1925, A. marginaria and A. aurantiaria to Phigaliohybernia Inoue, 1942. We consider this premature until a complete revision including all known species is carried out and it involves genetic data. In muti-gene analyses Agriopis is sister to a large group including for instance Arichanna and numerous non-European genera (Jiang et al. 2017), whereas in more extensive analysis it is sister to Calamodes (Murillo-Ramos et al. 2019).

104. Agriopis leucophaearia (Denis & Schiffermüller, 1775) Geometra leucophaearia Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wiener­ gegend: 101. Syntype(s) lost. Phalaena (Geometra) brumaria Borkhausen, 1794: Natur. eur. Schmett. 5: 214 (Europe). Syntype(s). Ph[alaena] (Geom[etra]) marmorinaria Esper, 1801: Die Schmett. 5 (8): 205, pl. 37, figs 1,2 (Europe). Syntypes ♂♀. Geometra nigricaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2), pl. 35, fig. 181 (Europe). Syntype(s). ­ empke Unavailable names (infrasubspecific): approximata: Lempke (1970) (f.); brunneoalbata: L (1952) (f.); brunneomarmorata: Lempke (1952) (f.); brunneostrigata: Lempke (1952) (f.); brunnescens: Lempke (1952) (f.); crassestrigata: Lempke (1952) (f.); confusa: Leraut (2009) (f.); confusaria: Preisecker (1922) (ab.); desparsata: Heydemann (1938) (ab.); effusa: Lempke (1952) (f.); flavescens: Lempke (1952) (f.); flavilinea: Leraut (2009) (f.); fuscata: Haverkampf (1906) (ab.); fuscomarmorata: Lempke (1952) (f.); grisescens: Lempke (1952) (f.); intermedia: Lempke (1952) (f.); medioobscuraria: Uffeln (1905) (ab.); merularia: Weymer (1878) (ab.); nigricaria: Hübner (1799) (ab.); obsoleta: Lempke (1952) (ab.); pallidalinea: Lempke (1952) (f.); rufomarmorata: Lempke (1952) (f.); uniformata: Lempke (1952) (f.); uniformata-obsoleta: Lempke (1952) (f.); trilinearia: Sibille (1905) (ab.).

External characters and abdomen: Male: Wingspan 22–29 mm. Forewing ground colour pale grey, more or less speckled dark grey. Postmedial line distinct, black, wavy, with black crescent-shaped blotch between M3 and CuA1, approaching medial line on inner margin. Antemedial line black, evenly bent. Medial line indistinct, wavy, sometimes absent. Basal and terminal areas brownish grey, the latter with small, interrupted whitish band. Medial area often slightly paler. Hindwing pale grey, lines indistinct, discal spot usually present, sometimes missing. Fringes weakly chequered or unicolorous. Terminal line black, delicate. Male antennae partially bipectinate (about six apical flagellomeres free of branches). Basal flagellomeres with ciliae only, then with branches of different length, more distally with symmetrical branches. Frons, collar and thorax and abdomen of ground colour, abdomen 316

brownish grey, with double row of black dots. Female: Nearly wingless, wing stumps very short. Antennae filiform. Hindtibia of both sexes with 2+2 spurs. Variation: Extremely variable species. Dark grey speckling more or less developed. Males occur in three forms almost without intermediates: 1. Nominotypical form as described above. Ground colour darker or lighter, lines more or less distinct. Dark grey speckling varies, too. 2. Whitish form on forewing with basal and terminal area black to dark blackish brown (“f. nigricaria”). 3. Melanic form, all wings black, in older collection specimens brownish, unicolorous, lines hardly visible. Veins darker (“f. merularia”). Species with a lot of described infrasubspecific (unavailable) names, see above. Male genitalia: Uncus and gnathos well developed. Uncus double-pointed, lateral margins slightly concave, gnathos ventrally fused, tip rounded. Valva partly membranous, apex with spine, base wider, costa on apical third with spined ridge and medial valva process with thorn on tip. Juxta constricted at centre, apically ending in two pointed forks. Aedeagus cylindrical, apex round, vesica with one spine. Spine base corrugated. Female genitalia: Papillae anales oval, membranous, soft, slightly setose. Apophyses posteriores very long (on average 3.8 mm). Apophyses anteriores shorter (on average 1.9 mm), ending spatulate. Ductus bursae short, wide, sclerotized. Lamella antevaginalis sclerotized, alate. Lamella postvaginalis inconspicuous. Corpus bursae nearly globular, anteriorly narrower. Central part weakly sclerotized, without signum. Distribution and abundance: West Palaearctic. In Europe from the Urals through the southern half of Russia and central Europe to the British Isles and from southern Scandinavia to the northern half of the Iberian peninsula, southern France, Italy including Sardinia and Sicily and the Balkan peninsula. – Rather common, usually occurring in large numbers. – Outside Europe in southern Kazakhstan, Armenia, Georgia and Turkey.

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Phenology: Univoltine: In southern Scandinavia from early March to mid-April, in south-­ western Germany from early February to late March (Ebert 2003) and in Italy from February to March (Flamigni et al. 2007). In England in recent years as early as December and January (M. Corley pers. comm.). In the mountains in northern Greece until late April. Larvae in May and June. Overwintering as pupa. Males active at night, attracted to light. Wingless females after dusk on tree trunks. Biology: Larva oligophagous. Found on Quercus robur, Q. petraea, Salix caprea and Rosa canina (Ebert 2003). Pupation in the ground. Habitat: Mesophilous, silvicolous. In oak forests and where oak grows, such as gardens, parks and hedges. In southern Scandinavia in the lowlands, in south-western Germany up to 730 m (Ebert 2003), in Greece up to 1000 m and in Spain up to 1300 m (Redondo et al. 2009). Similar species: A. bajaria with broader forewings in male, termen undulating. Genetic data: BIN: BOLD:AAC5604 (n=50 from Germany, United Kingdom, Austria, Czech Republic, France, Italy, Slovenia, Hungary, Croatia, Bulgaria, Tunisia). Genetically heterogeneous with shallow regional divergences, e.g. in southern Italy and Tunisia; maximum variation 2.2%. Nearest congener in current taxonomy: A. budashkini (8.6%), suggesting a long, isolated evolutionary history of A. leucophaearia.

105. Agriopis bajaria (Denis & Schiffermüller, 1775) Geometra bajaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 315 (Austria: Vienna district). Type(s). Geometra aerugaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s). Ph[alaena] (Geom[etra]) ligustriaria Lang, 1789: Verz. Schmett. Gegenden Augsburg: 180 (Germany: Augsburg). Syntype(s). Phalaena (Geometra) sericearia Borkhausen, 1794: Natur. eur. Schmett. 5: 212 (Europe). Syntype(s). Geometra sorditaria Hübner, 1809: Samml. Eur. Schmett. 5 Geometrae (1), pl. 64, fig. 334 (Europe). Syntype(s). Hybernia onytaria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 10: 253. Holotype ♂ (MNHN). Hibernia bajaria coggii Constantini, 1916: Atti Soc. Nat. Modena (5) 3: 16 (Italy: Monte Gibbio). Syntype(s) ♂. Validated at subspecific rank by Scoble (1999), in Leraut (2009) regarded as “form”, infrasubspecific status here confirmed, based on clinal character transitions and absence of constant differential features. Cryopega legrasi Dumont, 1925: Encycl. ent. 1 (1): 29, fig. 2I (France: Basses Corbières, Abbaye de Fontfroide, 30 km from Narbonne). Syntypes 1 ♂1 ♀ (MNHN). Erannis bajaria var. transbaikalica Wehrli, 1938: Mitt. münchn. ent. Ges. 28 (2): 84 (Novorotnaya, Schilkafluss, 2000 m). Holotype ♂ (ZFMK). Valid at subspecific rank (Scoble 1999). Erannis bajaria zehrounensis Wehrli, 1940: in Seitz, Gross-Schmett. Erde 4 (Suppl.):413, pl. 33: h (Morocco: Mrassine, Zehroun). Syntype(s) (ZFMK). Listed in synonymy by Scoble (1999), validated at subspecies rank by Leraut (2009), here confirmed. Unavailable name (infrasubspecific): unicolor: Schawerda (1916) (ab.).

External characters and abdomen: Male: Wingspan ♂ 24–33 mm. Forewing ground colour brownish grey to whitish, more or less speckled with black scales. Dark forms often with diffuse red-brown blotches. Post- and antemedial lines black, usually distinct, sometimes weak or broken. Postmedial line wavy, approaching diffuse and weak medial line on ­inner ­margin. 318

Antemedial line bent, subcostally zigzag angled. Terminal area with whitish wavy line, often broken or indistinct, rarely missing. Hindwing paler with antemedial and diffuse medial line, both often indistinct. Discal spot indistinct or absent. Termen slightly undulating. Terminal line on both wings black, often broken into a row of dots or streaks. F ­ ringes of ground colour. Male antennae partially bipectinate (apical 4–5 flagellomeres free of branches), basal flagellomeres ciliate. Female: Nearly wingless, wing stumps about 1 mm long. Antennae filiform. Hindtibia of both sexes with 2+2 spurs, frons smoothly scaled. Legs greyish. Frons, collar and thorax brownish grey, patagia black. Abdomen with double row of black dots. Variation: Variable species. Most males dark, uniformly brownish, poorly contrasted with lines varying in distinctness. Rarely well contrasted, with whitish ground colour, especially in medial and terminal areas of forewings, lines distinct, black; hindwings paler with distinct lines. Intermediate specimens are known. Male genitalia: Uncus with slightly concave margins, tip with short process. Gnathos ventrally fused, wide and apically rounded. Valva simple, wide, relatively short; costa concave, weakly sclerotized, setose; apex round, setose; in centre weak setose ridge. Juxta plate, inconspicuous. Saccus rounded. Aedeagus short, apex pointed. Vesica without cornutus. Female genitalia: Papillae anales very long, sclerotized, slightly setose. Apophyses posteriores extremely long (7−8 mm). Apophyses anteriores shorter (on average 3.35−3.5 mm), ending spatulate. Lamella antevaginalis and postvaginalis inconspicuous. Ductus bursae narrow, long, membranous. Corpus bursae long, membranous, anteriorly wider, elongated. Signum small, round, weakly sclerotized, sometimes inconspicuous (apparently missing). Distribution and abundance: West Palaearctic. In Europe from south-eastern Russia through central Europe to France and the Iberian peninsula. In the south to the Mediterranean basin, but absent from Corsica. – Rather common, usually occurring in limited numbers. – Outside

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Europe in Jordan, Israel, Cyprus, Turkey, Armenia, Azerbaijan, Turkmenistan and Trans­ baikalia (subsp. transbaicalica). Also in North Africa: Morocco (subsp. zehrounensis). Phenology: Univoltine: In southern Germany from mid-October to mid-November, further to the south until late December. In southern Spain until mid-February (P. Hale pers. comm.) in southern Italy and in Greece as late as mid-April (A. Hausmann; P. Skou own data). In Samos more abundant in early spring than in late autumn (Fritsch et al. 2014). Caterpillar from late April to early June (Ebert 2003). Overwintering in the north as egg, further to the south partly as adult and partly as egg. Males active at night, attracted to light. Wingless females at night on trees and bushes. Biology: Larva polyphagous. Found on Crataegus monogyna, Prunus spinosa, Rhamnus cathartica, Quercus petraea, Ulmus minor, Crataegus monogyna, Rosa canina, Prunus spinosa, Rubus, Cornus sanguinea, Lonicera xylosteum and Ligustrum vulgare (Ebert 2003) and on Pistacia lentiscus (B. Müller own data) and Paliurus spina-christi (I. Kostjuk pers. comm.). Pupation in the ground (T. Drechsel pers. comm.), under moss (M. Leipnitz pers. comm.) or in detritus in the surface of the ground (Z. Las˘tu˚ vka pers. comm.). Habitat: Xerothermophilous. In the north more restricted to xerothermic areas, such as scrubby dry slopes, forest edges or field hedges. In southern Europe widespread and less specialized, in open shrubland and scattered, low growing forests. In central Europe up to 800 m above sea-level, up to 1300 m in the Alps, in the Apennine peninsula from sea-level up to 1600 m (Flamigni et al. 2007). Similar species: A. leucophaearia with smaller forewings in males, termen not undulating. Genetic data: Genetically heterogeneous, diverging into five BINs: BOLD:AAC3211 (n=14 from Germany, Austria, Italy, Hungary, Croatia, Bulgaria, Greece, including Crete island, Kazakh­stan), BIN-sharing populations from Crete diverging by 1.6%. BOLD:AAC3210 (n=6 from Turkey, Cyprus); BOLD:AAC3212 (n=4 from Israel, northern Iran); BOLD:AAC3209 (n=1 from Italy: Calabria); BOLD:AAZ7585 (n=5 from Spain, Italy: Sardinia). BINs at distances of 2.8–3.2%, populations from Israel and northern Iran more strongly diverging (4.1–4.8%), possibly both requiring a separate name. Nearest congener in current taxo­nomy: A. leucophaearia (9.3%), suggesting a long, isolated evolutionary history of A. bajaria. Remarks: Leraut (2009) placed this species in the genus Cryopega Dumont, 1925 because of the genitalia structure.

106. Agriopis aurantiaria (Hübner, 1799) Geometra aurantiaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2): pl. 35, fig. 184 (Europe). Syntype(s). Phalaena Geometra mellearia Scharfenberg, 1805: in Bechstein & Scharfenberg, Vollständige Naturgeschichte der schädlichen Forstinsekten 3, 676 (Austria: Vienna). Syntype(s). Geometra incompletaria Haworth, 1809: Lepid. Br. (2): 305 (United Kingdom). Syntype(s). Hybernia aurantiaria lariciaria Scholz, 1947: Z. wien. ent. Ges. 31 (5–8): 113 (Austria). Syntype(s). Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes and absence of constant and significant differential features. Hybernia aurantiaria f. venosaria Galvagni, 1948: Z. wien. ent. Ges. 31 (8–10): 103 (Austria: Alps 1000 m). Syntype(s).

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Agriopis aurantiaria cleui Leraut, 1993: Entomol. Gall. 4 (4): 234 (France: Hautes-Alpes, l’Argen­ tière-la-Bessée). Holotype ♂ (MNHN). Validated at subspecific rank (Scoble 1999), downgraded to synonymy by Leraut (2009). Unavailable names (infrasubspecific): bellaria: Scholz (1947) (f.); bistrigaria: Lempke (1952) (f.); brunnea: Burmann (1944) (ab.); confluaria: Scholz (1947) (f.); demacularia: Scholz (1947) (f.); ellipsaria: Scholz (1947) (f.); fasciaria: Linstow (1907) (ab.); fumipennaria: Hellweger (1906) (ab.); maculata: Nordström (1941) (ab.); margineobscuraria: Scholz (1947) (f.); mar­ allida: moraria: Scholz (1947) (f.); nigra: Scholz (1947) (f.); obscura: Burmann (1944) (ab.); p Lempke (1952) (f.); poelli: Scholz (1947) (f.); unistrigaria: Lempke (1952) (f.); v­enosaria: ­Galvagni (1948) (ab.); weileri: Scholz (1947) (f.).

External characters and abdomen: Male: Wingspan 27–35 mm. Forewing ground colour deep yellow to yellowish brown, more or less speckled with pale brown scales, often weakly tinted towards pale purple. Medial lines usually distinct, pale brown. Postmedial line angled on M1 and Cu1. Antemedial line bent. Medial line diffuse, straight or slightly bent, running near antemedial line. Subterminal line broken into blotches. Hindwing yellow-white with straight postmedial line, inwardly bent close to costa. Terminal line on both wings usually absent, although sometimes marked by some dots. Fringes of ground colour. Frons, collar, thorax and abdomen of ground colour. Tegulae with long scales. Male antennae partially bipectinate (apical 4–5 flagellomeres free of branches), basal segment reddish brown. Basal flagellomeres ciliate. Female: Wings reduced to stumps with forewing length of about 2.5 mm. Ground colour pale to dark grey usually with indistinct lines. Antennae filiform. Frons, collar and thorax dark grey. Hindtibia of both sexes with 2+2 spurs, frons smoothly scaled. Variation: Little. In males transverse lines can be more or less distinct, rarely absent. In alpine Larix forests usually paler, on average with more distinct pattern (‘f. lariciaria’), sometimes with nearly unicolorous purplish brown forewings and yellowish fringes (‘f. fumipennaria’). Male genitalia: Uncus triangular, tip beak-shaped. Gnathos ventrally fused, wide and apically rounded. Valva simple, slim, base slightly wider with subcostal setose ridge. Apex ending in slightly curved thorn. Juxta bifurcate, ending in two long spines, lateral margins convex. Saccus rounded. Aedeagus narrow, long, curved, apex and caecum rounded. Vesica with spine-shaped cornutus, less curved than in A. budashkini (further differences see there). Female genitalia: Papillae anales oval, membranous, setose. Apophyses posteriores long (on average 3.1 mm), apophyses anteriores shorter (on average 1.6 mm), stout, both spatulate at tip. Ductus bursae conspicuous, strongly sclerotized, large, constricted in posterior third, then widening. Corpus bursae membranous, anterior part wider, rounded, with small, blotch-shaped signum. Posterior part with small, sclerotized, striated area, laterally with membranous diverticulum (smaller and less sclerotized than in A. budashkini). Distribution and abundance: European-Anatolian/Turanian. In Europe from southern Russia through central Europe to Ireland and Portugal and from central Scandinavia, with an isolated occurrence in northern Norway, to the Balkan peninsula, southern Italy, southern France and the northern half of the Iberian peninsula. – Common or very common, sometimes occurring in numbers. In southern Finland sometimes arriving as migrant, sometimes in very large numbers (P. Sihvonen own data). – Outside Europe from Turkey to Georgia, Armenia and Azerbaijan. Phenology: Univoltine: In most of Europe from mid-October to late November, sometimes from late September or until mid-December. In Spain from November until February (­Redondo et al. 2009). Larva from late April or early May to mid-June. Overwintering as 321

egg, in Spain as adult moth. Males active at night, attracted to light; females at night on tree trunks, often together with males. Biology: Polyphagous. Found on Salix caprea, Carpinus betulus, Corylus avellana, Betula, Fagus sylvatica, Quercus, Ulmus, Pyrus communis, P. pyraster, Malus domestica, Sorbus ­aucuparia, Crataegus, Rubus idaeus, R. fruticosus, Rosa vosagiaca, Prunus domestica, P. spinosa, Tilia, Vaccinium myrtillus, Fraxinus excelsior, Ligustrum vulgare (Germany: Ebert 2003) and Populus tremula (U.K.: Skinner 2009). Found on Crataegus also in north-western Italy (King & González-Estébanez 2015). Habitat: Silvicolous. In a wide variety of habitats such as deciduous and mixed forests, shrubberies, parks and gardens. In central Europe from 0 up to 1250 m above sea-level. Similar species: A. budashkini is very similar in its external characters and genitalia structure (see there). Genetic data: BIN: BOLD:AAC2753 (n=38 from Finland, United Kingdom, Germany, Austria, Czech Republic, France, Italy, Iran). Genetically homogeneous in Europe, but populations from northern Iran (same BIN) diverging by 1.9%. Nearest species: A. budashkini (3.5%). Remarks: A. aurantiaria and the following two species form a group of similar species in their genitalia structure and genetics.

107. Agriopis budashkini Kostjuk, 2009 Agriopis budashkini Kostjuk, 2009: Kiev Nat. Univ., Proc. Zool. Mus. 5: 61–64 (Ukraine: Crimea, Karadag). Holotype ♂ (ZMKU).

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External characters and abdomen: Male: Wingspan 30–38 mm. General appearance as in less contrased forms of A. marginaria. Forewing ground colour similarly beige, more or less speckled with pale brown scales. Post- and antemedial lines as in A. marginaria. Medial line inconspicuous or absent. Terminal line absent (as in A. aurantiaria, in A. marginaria broken to black dots). Hindwing pale brownish white (in marginaria paler) with straight postmedial line, inwardly bent near costa. Fringes, frons, thorax and abdomen as in A. marginaria. Basal segment of male antennae reddish brown (in A. marginaria whitish). Specimens from Crete on average less pale. Female: Wings reduced to stumps with forewing length of about 2.5 mm as in A. aurantiaria (wing stumps considerably larger in A. marginaria). Ground colour pale to dark grey usually with indistinct lines. Abdomen with double row of black dots. Antennae filiform. Hindtibia of both sexes with 2+2 spurs, frons smoothly scaled. Frons, collar and thorax dark grey to light grey. Male genitalia: Similar to those of A. aurantiaria. Differing in the structure of the aedeagus and in the shape of bifurcate juxta with its two arms. Base of arms arising at a distance of two-thirds of juxta length from its anterior end (in A. aurantiaria about half length). Base flatter, in central part obtusely angled or arcuate (in A. aurantiaria acutely angled). Valva slightly wider than in A. aurantiaria. Aedeagus more slender, slightly curved, apex pointed, caecum with short process. Female genitalia: Similar to those of A. aurantiaria. Posterior part of corpus bursae sclerotized extensively, right-hand diverticulum sclerotized (posterior part sclerotized weakly, right-hand diverticulum membranous in A. aurantiaria). Distribution and abundance: (South-east) European-West Asiatic. Due to the recent discovery of this species, the distribution area is still insufficiently known, but it has so far been found in Crimea, Crete, Greece, Bulgaria, Serbia, Croatia and Hungary. – Scarce data on abundance, with scattered records in the distribution area. However, found in numbers on several occasions. – Outside Europe recorded in Turkey (Anatolia).

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Phenology: Univoltine: From early November (J. Gelbrecht pers. comm.) to early January (Kostjuk 2009). In an ab ovo rearing, the caterpillars hatched between 16th and 19th April (I. Kostjuk pers. comm.). Larva found in late May in northern Greece (Ziegler 2019). Overwintering partly as pupa, as adult and partly as egg (the last recorded by Y. Budashkin & V. Savchuk pers. comm.). Males active at night, attracted to light (V. Savchuk, M. Leipnitz & I. Kostjuk pers. comm.). Females on trunks at night (recorded by putting tape around tree trunks). Biology: Larva polyphagous. Found on Acer sempervirens, Crataegus, Quercus coccifera and Q. pubescens (M. Leipnitz pers. comm.). Reared on Acer pseudoplatanus and A. mons­ pessulanum (M. Leipnitz pers. comm.). Pupation in the ground (I. Kostjuk pers. comm.). Habitat: Silvicolous, inhabiting scattered forest in steppe regions. In Karadagh mts. in Crimea in open Quercus and Pistacia woodland (I. Kostjuk pers. comm.). From sea-level up to 900 m. Similar species: A. aurantiaria is very similar in its external characters and genitalia structure, similar also A. marginaria (see above). Genetic data: BIN: BOLD:AAJ6603 (n=8 from Croatia, Greece (including Crete), Ukraine, Turkey). Genetically homogeneous. Nearest species: A. aurantiaria (3.5%).

108. Agriopis marginaria (Fabricius, 1776) Phalaena marginaria Fabricius, 1776: Genera Insect.: 286 (Germany: Hamburg). Syntype(s). Phalaena testacea Retzius, 1783: Genera et Species Insectorum: 42 (Europe). Syntype(s). Phalaena Geometra testacearia Villers, 1789: Linn. ent. 2: 321. Emendation of testacea Retzius. Phalaena diversaria Fabricius, 1794: Ent. Syst. 3 (2): 157 (England). Syntype(s). Geometra progemmaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (2), pl. 35, fig. 153 (Europe). Syntype(s). Geometra connectaria Haworth, 1809: Lepid. Br. (2): 285 (Great Britain). Syntype(s). Geometra luctuaria Haworth, 1809: Lepid. Br. (2): 279 (United Kingdom). Syntype(s). Hibernia marginaria var. infumata Fuchs, 1899: Jb. nassau. Ver. naturk. 52: 146 (Scotland). Syntypes 5 ♂. Hybernia marginaria f. pallidata Turati, 1911: Z. wiss. InsektBiol. 7 (7–8): 211 (Italy: Lombardy; Sicily). Syntypes. Validated at subspecific rank by Scoble (1999) and Leraut (2009), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Unavailable names (infrasubspecific): albescens: Lempke (1952) (f.); basifasciata: Leraut (2009) (f.); bicolor: Lempke (1952) (f.); contrasta: Lempke (1952) (f.); denigraria: Uffeln (1905) (ab.); destrigaria: Lempke (1952) (f.); diversaria: Fabricius (1794) (ab.); fasciata: Lempke (1970) (f.); fuscata: Harrison (1878) (ab.); grisea: Lempke (1970) (f.); lesaunieri: Mabille (1916) (ab.); pallida: Lempke (1970) (f.); postgrisescens: Lempke (1970) (f.); pseudobicolor: Lempke (1970) (f.); quadristrigaria: Lempke (1952) (f.); signata: Lempke (1952) (f.); subfasciata: Lempke (1970) (f.); unistrigaria: Lempke (1952) (f.).

External characters and abdomen: Male: Wingspan 27–34 mm. Forewing: Ground colour beige, more or less densely speckled with brown. Transverse lines brownish. Postmedial line distinct, angled on M1 and Cu1. Antemedial line bent. Medial line usually straight, running near antemedial line. Subterminal line variable, brownish, often broken, shaded by the whitish beige wavy line, the latter broken into spots, if any. Hindwing much paler with distinct postmedial line. Terminal line broken into black dots between veins. Both wings with fringes of ground colour and discal spots, the latter elongate on forewings. Frons, thorax 324

and abdomen beige, vertex and patagium paler. Tegulae with long scales. Male antennae partially bipectinate (about six apical flagellomeres free of branches), basal segment whitish. Basal branches reduced. Female: Wings reduced, grey, square, hindwing longer than forewing, termen of forewing concave, straight or bent, of hindwing rounded. Wingspan 9–13 mm. Forewing with distinct black postmedial, medial and antemedial lines, the first two often anastomosing, the last absent from hindwing. Abdomen dark grey, dorsally with double row of black dots and often with paler line in between. Antennae filiform. Hindtibia of both sexes with 2+2 spurs, frons smoothly scaled. Variation: Variable species. Occasional male specimens with reddish beige ground colour. Brownish speckling of forewing varies in a wide range, i.e. wings are darker or paler. All transverse lines or some of them can be reduced, postmedial line can be darkly shaded. Extreme forms are unicolorous brown. Females vary in position and distinctness of lines. Male genitalia: Uncus and gnathos well developed. Uncus triangular, apically with long, narrow, double-pointed process, lateral margins slightly setose. Gnathos ventrally fused, tip rounded. Valva simple, narrow, base slightly wider with subcostal setose ridge. Apex rounded, setose, with spine at tip. Costa sclerotized, concave. Juxta sclerotized wide and long sclerite, distally flattened. Sacculus convex. Saccus rounded. Aedeagus apically rounded. Vesica with spine-shaped cornutus. Female genitalia: Papillae anales oval, membranous, setose. Length of apophyses posteriores about 2.4 mm, apophyses anteriores shorter (on average 1.5 mm), stout, both ending spatulate. Ductus bursae wide, short, sclerotized. Posterior part of corpus bursae sclerotized, triangular, anterior part membranous, partly striated. Signum weakly sclerotized with 2–3 darker, small dots. Distribution and abundance: West Palaearctic. In Europe from the Ural mountains to western­ most Europe and from southern Scandinavia to Greece including Crete (M. Leipnitz pers.

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comm.), Italy including Sicily and Sardinia, France including Corsica and the northern half of the Iberian peninsula with an apparently isolated population in south-western Spain. – Common or very common, usually occurring in limited numbers. – Outside Europe in Turkey, Georgia, Armenia, Azerbaijan and the Altai region and to the south-west in Morocco and Algeria. Phenology: Univoltine: In southern Scandinavica from late February or early March to midApril, in south-western Germany from mid-February to early April (Ebert 2003). In Italy from February to April, in the south as early as December (Flamigni et al. 2007) and in Spain from February to May (Redondo et al. 2009). Such very late records must refer to high altitudes. In southern Spain from late January to early March (P. Hale pers. comm.). In northern Greece in mid-April at 1000 m altitude (P. Skou own data). Caterpillars from late April to mid-June (Ebert 2003). Overwintering as pupa. Males active at night, readily attracted to light; wingless females found sitting or crawling on tree trunks and branches. Males similarly often found by day on trunks or on leaves on the ground. Biology: Larva polyphagous. Found on Salix, Betula pendula, Fagus sylvatica, Quercus, Malus domestica, Sorbus aria, Crataegus, Rosa canina, R. vosagiaca, Prunus spinosa, P.  domest­ica, Tilia, Hippophae rhamnoides, Vaccinium myrtillus and Lonicera xylosteum (Ebert 2003). Pupation in the ground. Habitat: Silvicolous. In a large varity of habitats with deciduous trees and bushes, such as forest, forest edges, shrubberies, parks and gardens. From sea-level, in south-western Germany up to 1100 m (Ebert 2003), in Italy up to 1300 m (Flamigni et al. 2007) and in the Iberian peninsula up to 1400 m (Redondo et al. 2009). Similar species: A. aurantiaria and A. budashkini (differences see under A. budashkini). Genetic data: BIN: BOLD:AAC0355 (n=79 from 17 countries from Iceland and United Kingdom to Hungary and Greece (including Crete) and from Finland to North-West Africa). Genetically rather homogeneous, with populations from Crete slightly diverging (1.2%). Nearest species: North American Parapheromia lichenaria (Pearsall, 1906) (5.8%). Nearest European congener: Agriopis leucophaearia (6.0%). Remarks: Leraut (2009) placed this species in the genus Phigaliohybernia Inoue, 1942 because of the genitalia structure, see comments to genus Agriopis.

Calamodes Guenée, 1858 Calamodes Guenée, 1858, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 229. Type species: Boarmia occitanaria Duponchel, 1829. There are no genus synonyms.

Diversity and distribution: According to Scoble (1999) Calamodes contains four West Medi­ terranean species. Leraut (2009) synonomized the taxa C. boursini Albers & Warnecke, 1941 and C. subscudularia (Turati, 1919) with C. melanaria Oberthür, 1913. Furthermore, he considers all Mediterranean taxa as belonging to the same species, C. occitanaria (­Duponchel, 1829). Recent results from DNA barcoding (see below) show a 2%-difference between C. occitanaria and C. melanaria and seem to confirm C. melanaria and C. subscudularia as conspecific, the former being the valid name. We treat C. occitanaria and C. melanaria as different species. Remarks: In multi-gene analysis Calamodes is sister to Agriopis (Murillo-Ramos et al. 2019). 326

109. Calamodes occitanaria (Duponchel, 1829) Boarmia occitanaria Duponchel, 1829: in Godart & Duponchel, Hist. nat. Lépid. Papillons Fr. 7 (2): 360, pl. 159, fig. 5 (France: Languedoc). Syntype(s). Boarmia occitanaria submelanaria Zerny, 1927: Eos, Madr. 3 (4): 423, pl. 10, figs 30, 31 (Spain: Aragon, Albarracin). Syntypes ♂♀. Validated at subspecific rank by Scoble (1999) and L ­ eraut (2009), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features. Boarmia occitanaria f. bejaranaria Fernández, 1931: Eos, Madr. 7 (2): 220, pl. 2, figs 16 (Spain: Salamanca, Béjar). Syntype(s). Unavailable names (infrasubspecific): pallida: Leraut (2009) (f.). – (incorrect subsequent spelling): occidanaria: Freyer (1837; pl. 210, figs 1, 2).

External characters and abdomen: Wingspan ♂ 26–31 mm, ♀ 24–28 mm. Ground colour greyish white to pale grey, more or less speckled dark grey. Forewing: Postmedial line between costa and vein M1 serrate, between M1 and inner margin curved, grey shaded. Medial line straight or slightly bent, antemedial line oblique, near costa double angled. All these lines black. Wavy line white, distinct, slightly bent. Terminal line black, often broken into dots. Fringes of ground colour, more or less chequered, with indistinct longitudinal line. Hindwing with nearly straight medial lines. Both wings with blackish discal spots. Antennae partially bipectinate (apical 8–9 flagellomeres free of branches) in male, filiform in female. Frons, collar and thorax of ground colour, abdomen of ground colour. Hindtibia of both sexes with 2+2 spurs. Variation: Variable in the degree of dark grey speckling. Therefore darker or lighter forms can be observed. Specimens from Iberian peninsula (‘f. submelanaria’) on average darker, reminiscent of the dark North African C. melanaria Oberthür, 1913 (= C. subscudularia). Male genitalia: Uncus and gnathos well developed. Uncus wide at base, evenly narrowing, apex double beak-shaped, pointed. Gnathos ventrally fused, apex small, rounded. Valva wide, short, apically with curved margin. Costa sclerotized, apex setose. At centre a ridge with group of arcuately arranged setae and a sclerotized harpe with spine. Sacculus slightly curved, sclerotized. Saccus rounded. Juxta ending in two pointed processes of 2/3 of the total length. Aedeagus stout, anteriorly evenly narrowing, without cornutus. Apex with sclerotized striated structure. Female genitalia: Papillae anales elongated, membranous, soft, slightly setose. Apophyses posteriores very long (on average 2.5 mm). Apophyses anteriores shorter (on average 1.2 mm), latter ending spatulate. Ductus bursae shield-shaped, weakly sclerotized, laterally convex, therefore wider at centre (see C. melanaria). Posteriorly with setose funnel-shaped invagination. Lamella antevaginalis inconspicuous. Lamella postvaginalis narrow sclerite, but wider than in C. melanaria, distally rounded. Corpus bursae long, narrow, membranous, without signum. Distribution and abundance: West European. Restricted to the Iberian peninsula, southern France and north-westernmost Italy. Erroneously mentioned for Sicily in Leraut (2009). – Rather common, now and then found in numbers. – No confirmed records from outside Europe. Phenology: Univoltine. From late August to early November. Caterpillars from October to May (D. Fritsch pers comm.). Hibernation as larva (D. Fritsch & G. King pers. comm.). Adults active at night, both sexes attracted to light. 327

Biology: Larva oligophagous. Found on Thymus lacaitae (G. King pers. comm.) and Lavandula stoechas (D. Fritsch pers comm.). Recorded on Thymus, Rosmarinus (Robineau et al. 2007). Pupation in a loose cocoon on the ground (T. Varenne pers. comm.). Habitat: Xerothermophilous. In quite a wide variety of open or semi-open habitats including semi-deserts in south-eastern Spain. From sea-level up to 1600 m, in Sierra Nevada (southern Spain) even one specimen at 2250 m (coll. Skou). Similar species: Calamodes melanaria. Genetic data: BIN: BOLD:AAL9658 (n=3 from Spain). Genetically heterogeneous, maximum variation 1.1%. Nearest species: C. melanaria (2.0%).

110. Calamodes melanaria Oberthür, 1913 Calamodes melanaria Oberthür, 1913: Études Lépid. comp. 7: 262, pl. 166, fig. 1624 (western Algeria: Géryville [El Bayadh]). Syntype(s) including ♂. Calamodes bistriata Rothschild, 1914: Novit. zool. 21: 349 (Algeria central: Guelt-Es-Stel)). Holotype ♂ (NHMUK). Calamodes sordidaria Rothschild, 1914: Novit. zool. 21: 349 (Algeria central: Guelt-Es-Stel)). Holotype ♂ (NHMUK). Boarmia subscudularia Turati, 1919: Naturalista sicil. 24 (7–12): 314, pl. 4, figs 25, 26; text-fig. 2 (‘Sicilia-Cirenaica’: Italy: Sicily; Libya: Cyrenaica). Syntypes. Validated at species rank in Scoble (1999), here tentatively subordinated at subspecies rank under C. melanaria, see, however, also under Genetic data. Requiring further research and lectotype designation.

External characters and abdomen: Generally as in C. occitanaria, wingspan on average a little smaller (♂ 23–27 mm). Only dark forms known. Postmedial line between M1 and inner margin often weakly serrate and less angled. After Albers & Warnecke (1941) termen 328

of forewing more rounded. Especially in females this is sometimes unreliable as diagnostic character. Variation: Little. Postmedial, medial and antemedial lines sometimes indistinct. Male genitalia: Generally as in C. occitanaria with the following small differences: Beakshaped apex of uncus less pointed than in the latter, double beak only hinted. Ridge in the centre more straight with group of setae. Spine on harpe on average larger than equivalent in C. occitanaria. Aedeagus short, straight, caecum with rounded diverticulum. Female genitalia: Generally as in C. occitanaria with the following differences: Apophyses posteriores and anteriores ending spatulate. Ductus bursae shield-shaped, smaller than in C. occitanaria, laterally straight, therefore not wider at centre, posteriorly with tube-shaped invagination. Lamella postvaginalis narrowly sclerotized, smaller than in C. occitanaria, distal half evenly narrowing, apex rounded. Distribution and abundance: Central Mediterranean (when considering the Levantine populations as non-conspecific). In Europe only in Basilicata and Calabria in southern Italy, and in Sicily. – Only known from a few localities, here sometimes abundant (S. Scalercio pers. comm.), especially at sea-level; at inland localities mainly as single specimens. – Outside Europe in Algeria, Tunisia, Eastern Libya (Oberthür 1913; Rothschild 1914; Turati 1919; Flamigni et al. 2007), doubtfully conspecific in Israel and Jordan (A. Hausmann own data), see also Genetic data. Moroccan populations requiring further study.

Phenology: Univoltine. From mid-September to early November (Flamigni et al. 2007). Larval time as well as stage of hibernation unknown. Adults active at night, readily attracted to light.

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Biology: Unknown. Habitat: Xerothermophilous. In Europe in large dry river beds, mainly close to the sea-line. In Tunisia in wadis (D. Stadie pers. comm.). Wadis are probably the original habitat for the species, which in southern Italy has found somewhat similar conditions on the wide, dry shores of the river beds there. The locality in Basilicata is situated at about 300 m, in Calabria close to sea-level. In Sicily a few single specimens above 600 m. Similar species: C. occitanaria. Genetic data: BIN: BOLD:AAD2402 (n=13 from Italy: Sicily, Calabria; Tunisia). Genetically slightly heterogeneous in Europe, maximum variation 0.9%. North African populations BIN-sharing and very close to European populations (0.15%). The populations from Israel and Jordan, currently attributed to ‘C. melanaria subscudularia’ – though BIN-sharing – possibly not conspecific, genetic minimum distance 1.6% (n=2).

Paradarisa Warren, 1894 Paradarisa Warren, 1894, Novit. zool. 1: 433. Type species: Boarmia comparataria Walker, 1866. There are no genus synonyms.

Diversity and distribution: Five species in the Palaearctic and Oriental regions. External characters and abdomen: Medium- to large-sized moths, wingspan 34–45 mm. Ground colour whitish grey to brownish with dark mottling. Transverse lines distinct. Male forewing with fovea. Abdomen with long hairy scales ventrally. Forewing vein R1 arising from Sc, anastomosing with R2. The latter arising from cell, sometimes connected with R3 and R4 (Sato 1980). Antennae ciliate-fasciculate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Uncus sclerotized, triangular. Gnathos fully developed, arms narrow, ventrally fused. Sacculus with process. Harpe with spines. Aedeagus with two sets of cornuti. Female genitalia: Corpus bursae membranous with stellate signum. Ductus bursae short, distinctly sclerotized. Posterior part of ductus bursae striated, with sclerotized diverticulum. Remarks: In multi-gene analysis Paradarisa is sister to Calamodes + Agriopis (Murillo-Ramos et al. 2019).

111. Paradarisa consonaria (Hübner, 1799) Geometra consonaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 30, fig. 157 (Europe). Syntype(s). Boarmia consonata Herrich-Schäffer, 1863: Syst. Verz. Eur. Schmett. (Edn 3): 12. Emendation of consonaria Hübner. Boarmia tetragonaria Curtis, 1829: Br. Ent. 6: folio 280, pl. 280 (United Kingdom). Holotype ♀. Unavailable names (infrasubspecific): intermedia: Lempke (1970) (ab.); nigra: Bankes (1905) (ab.); nigrescens: Lempke, (1970) (ab.); waiensis: Richardson (1940) (ab.).

External characters and abdomen: Wingspan 34–40 mm. Ground colour whitish grey with dark grey or brownish mottling. Females on average lighter than males, more contrasted. Postmedial line distinct, marginally edged, partially interrupted, between M1 and costa straight, at costa obtusely angled. Wavy line whitish, usually distinct, at one-third below 330

costa with dark, nearly quadrate blotch. At centre dark square blotch between postmedial and wavy line. Both blotches sometimes weak or missing. Terminal line black, forming thicker crescents between veins. Medial line weak. Antemedial and postbasal lines bent, close together. Discal spots usually weak. Underside lacking contrast with weak lines and discal spots. Lines on forewing usually ending before inner margin. Fringes, abdomen, thorax, frons and collar concolorous with ground colour. Fringes with weak greyish line, parallel to termen. First abdominal segment silver-grey, second one darker than the rest. Antennae ciliate-fasciculate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variable. Ground colour and degree of irroration on all wings variable from greyish to brownish or ochreous, darker or lighter. Melanic specimens of P. consonaria are usually unicolorous blackish, usually with whitish wavy line. From Central Europe known with greyish white blotch between wavy and marginal lines. Intermediate forms are reported from England with a whitish medial and a black basal and terminal area (Skinner 2009). Male genitalia: Uncus sclerotized, triangular with three short tips. Posteriorly prominent curved setae originating from base of uncus. Gnathos fully developed, arms narrow, ventrally fused, forming granular spoon-shaped plate. Valva membranous. Costa sclerotized, apex setose, round. Sacculus sclerotized, apically with very short process. Harpe with three or four spines. Juxta weakly sclerotized round plate, posteriorly with narrow elongated process. Saccus rounded. Aedeagus short, thick, apex digitiform, vesica with numerous spines, in fan-shaped position, and drop-shaped cornutus. Female genitalia: Papillae anales slightly sclerotized, slightly setose, elongate. Apophyses posteriores long, straight, apophyses anteriores shorter, both ending spatulate. Lamella antevaginalis desmoid, partly sclerotized. Lamella postvaginalis small plate, with rounded margins. Ductus bursae short, distinctly sclerotized. Posterior part of ductus bursae striated, with large sclerotized diverticulum. Anterior part membranous, partly striated concentrically. Signum near anterior end, stellate. Distribution and abundance: Eurasiatic. In Europe from the Urals to southern Ireland and from central Fennoscandia to northern Iberian peninsula, southern France, the Alps and northern Balkan peninsula. Spanish populations not shown on the map in Leraut (2009). – Rather local, usually rare, now and then occurring in numbers. – Outside Europe from Turkey across southern Siberia to Far East Russia, Kurile Islands and Japan. Phenology: Univoltine. In southern Scandinavia from mid-May to mid-June, in south-western Germany from early April to early June (Ebert 2003) and in the Alps from early April to mid-June (Kerschbaum & Pöll 2010). Larva from mid-June to mid-August (Porter 2010). Hibernation as pupa. Adults often found by day on tree trunks, but also active at night, attracted to light. Biology: Larva polyphagous. Found on Sorbus aucuparia (Ebert 2003), and recorded on Betula, Fagus, Quercus, Pinus sylvestris, Taxus baccata (Porter 2010), Corylus, Alnus, Prunus spinosa and Tilia (Bergmann 1955). Pupation in the soil. Habitat: Silvicolous. In forests and forest edges. From sea-level up to 1000 m in the Alps (Kerschbaum & Pöll 2010) and up to 1200 m in south-western Germany (Ebert 2003). Similar species: Ectropis crepuscularia with two distinct teeth on veins M3 and CuA1, blotch on wavy line weak, usually absent. In hindwings basal area concolorous with ground colour, in P. consonaria on anal margin and basally usually darker than the ground colour. Male 331

antenna of E. crepuscularia bears a larger triangular tooth on each flagellomere, from which the bunch of cilia arises. Differences in genitalia structures are distinct. Genetic data: BIN: BOLD:AAC3271 (n=24 from Finland, United Kingdom, Germany, Austria, Italy). Genetically very homogeneous. Nearest species: P. sumatrana Sato, 2004 from Sumatra (7.5%). Remarks: P. consonaria was placed for a long time in the genus Ectropis. Sato (1979; 1980) transferred this species to Paradarisa because of differences in the larval characters and conformity of the genitalia structures with this genus.

Larerannis Wehrli, 1935 Larerannis Wehrli, 1935, Ent. Z. Frankf. a. M. 49 (14): 107. Type species: Phigalia orthogrammaria Wehrli, 1927. There are no genus synonyms.

Diversity and distribution: The genus Larerannis contains four species in the Eastern Palaearctic region, all known species are described from Far East Russia or Japan. All four species are treated in detail by Nakajima (1998), including colour plates of males and females as well as genitalia photos. External characters and abdomen: Similar to Agriopis Hübner, 1825, venation different: On hindwings veins Sc+R1 and Rs fused to half the cell length as in Larentiinae. On forewing R1 and R2 stalked, R1 anastomosing with Sc; cell double angled (Wehrli 1935). Male wings thinly scaled, wingspan 29−38 mm, females brachypterous, belonging to the so called ‘winter moths’. Male genitalia: Uncus triangular, gnathos present, ventrally fused. Valva with sclerotized costa and sacculus. Costa with inwardly directed projection, sacculus terminating in a curved 332

pointed spine. Juxta a plate with long bifurcate process. Aedeagus with one large spineshaped and one smaller cornutus. Female genitalia: Papillae anales narrow, slightly setose. Apophyses posteriores long, apophyses anteriores shorter. Posterior part of corpus bursae membranous, signum with two tips. Anterior part wider, sclerotized. Corpus bursae between anterior and posterior part strongly constricted. Ductus bursae wide, posterior part striated (description after pictures of Nakajima 1998). Remarks: Larerannis has not been included in a multi-gene analysis so far. The position of Larerannis is tentative. Beljaev (2016) placed it between Arichanna and Phigalia.

112. Larerannis orthogrammaria (Wehrli, 1927) Phigalia orthogrammaria Wehrli, 1827: in Bang-Haas, Horae macrolepidopt. Reg. palaearct. 1: 97, pl. 11, fig. 35 ([Russia]: Southern Ussuri, Sutschansk [Partisan]). Syntype(s) ♂.

External characters and abdomen: Male: Wingspan 30−38 mm. Ground colour whitish. Postmedial line distinct, black, slightly wavy, thickened on veins and at costa, in terminal area shaded by a brown band. Medial, antemedial and subterminal lines present, but weaker. Subterminal line towards costa more diffuse and wider. Terminal line black, dotted. On hindwing postmedial, medial and antemedial lines weak, stronger near anal margin, discal spot small. Forewing more speckled brown than the hindwing, all wings thin-scaled. Fringes concolorous with ground colour. Antennae partially bipectinate, about 3–4 apical flagellomeres free of branches, basal flagellomeres with ciliae only. Frons, collar and thorax of ground colour, abdomen brownish, with indistinct double row of dots. Patagia posteriorly with a dark brown ring. Female with short wing stumps, about 2.5 mm long, without lines. Frons, collar, thorax and abdomen grey. Antennae filiform. Abdomen with a double row of black dots. Hindtibia of both sexes with 2+2 spurs. Variation: Very little. Male genitalia: Uncus triangular, slightly setose, apex ventrally bent, in lateral view beakshaped, apically more sclerotized. Gnathos ventrally fused, apex rounded, sclerotized. Valva membranous at centre, costa sclerotized, setose with inwardly directed, partly setose projection. Sacculus sclerotized, apically ending in a curved pointed spine. Juxta trapezoid, posteriorly with a long, narrow, bifurcate process. Saccus pointed. Aedeagus narrow, slightly constricted, caecum rounded. Vesica with a conspicuous spine and a small trapezoid ­cornutus. Female genitalia: Papillae anales narrow, slightly setose. Apophyses posteriores long, apophyses anteriores shorter. Posterior part of corpus bursae membranous, partly striated, signum a weakly sclerotized, large plate, anteriorly with two tips. Anterior part of corpus bursae wider, sclerotized. Corpus bursae strongly constricted between anterior and posterior part. Ductus bursae wide, posterior part striated, sclerotized. Distribution and abundance: East Palaearctic. Recently found in and around Moscow in European part of Russia (Mironov et al. 2008; Anonymous 2018; A. Ponomarev in Lepiforum 2018), reputedly introduced from Japanese populations (Lepiforum 2018). – Outside Europe in Far East Russia, Japan (Hokkaido, Honshu) and China. Phenology: Univoltine. Recorded from late September to mid-October. 333

Biology: Larva polyphagous. Recorded on Betula utilis, Cotoneaster, Populus, Salix and Rosa in China and on Fagus crenata in Japan (Hosts 2017). Habitat: Silvicolous. In European Russia in forests and parks, in the lowlands at about 200 m, in China up to 2200 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:ADD2374 (n=2 from China). Nearest species: Agriopis marginaria (7.4%) and Asian Arichanna transfasciata Warren, 1893 (8.2%).

Chemerina Boisduval, 1840 Chemerina Boisduval, 1840, Genera Index meth. eur. Lepid.: 193. Type species: Ligia caligeniaria Rambur, 1833. Genus synonyms after Scoble (1999). ‡ Chimerina Agassiz, 1847, Nomencl. zool. (Index univl.): 79, 81. [Emendation of Chemerina Boisduval, and junior homonym of Chimerina Eschscholtz, 1829 (Aves)]. Dysemon Lederer, 1853, Verh. zool.-bot. Ver. Wien 3 (Abh.): 177, 211, 212. [Unnecessary replacement name for Chimerina Agassiz, 1847].

Diversity and distribution: Chemerina is a monotypic genus, the only species occurs in the West Palaearctic region. External characters, abdomen and genitalia: see species description below. Remarks: Chemerina has not been included in a multi-gene analysis so far. In an extensive COI neighbour joining tree of Palaearctic Ennominae Chemerina and Fagivorina group together with a long-stalked common node (A. Hausmann own data).

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113. Chemerina caliginearia (Rambur, 1833) Ligia caliginearia Rambur, 1833: Annls Soc. ent. Fr. 2 (1): 35, pl. 2, fig. 34 (Corsica: Ajaccio; near Hières). Syntype(s). Chemerina ramburaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 193. Unnecessary replacement name for caligin[e]aria Rambur. Chemerina caliginearia adriatica Schwingenschuss & Wagner, 1927: Z. öst. EntVer. 12 (6): 63, figured (Croatia: southern Dalmatia). Syntype(s). Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on clinal character transitions and absence of constant and significant differential features. Chemerina punctinervis Rothschild, 1914: Novit. zool. 21: 348 (central Algeria: Guelt-Es-Stel). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999), here confirmed. Unavailable names: (infrasubspecific): pallescens: Leraut (2009) (f.); (incorrect subsequent spelling): caliginaria: Boisduval (1840).

External characters and abdomen: Wingspan ♂ 33–41 mm, ♀ 25–33 mm. Forewing: Ground colour grey, irrorated with darker scales. Post- and antemedial lines dark grey, bent, shaded with brownish band. Wavy line slightly wavy or nearly straight. Terminal line usually broken into dots. Costa nearly straight in males, convex in females. Hindwing light grey, without markings. Fringes of all wings unicolorous grey. Frons, vertex, collar, tegula and thorax concolorous with ground colour. Antennae partially bipectinate in male, apically about ten flagellomeres free of branches. Anennae filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Uncus triangular, laterally slightly concave, tip short, bent. Gnathos arms long, narrow, fused ventrally, forming a small rounded plate, medial part membranous. ­Valva wide with simple structure. Costa straight, sclerotized, weakly setose. Apical end of sacculus rounded, setose, slightly longer than costal part of valva. Juxta triangular, tip rounded. Saccus rounded. Aedeagus thick, short, with sclerotized apex. Vesica with row of numerous cornuti of different length. Female genitalia: Papillae anales slightly sclerotized, setose, wide but short. Apophyses posteriores of medium length, posteriorly thickened, apophyses anteriores shorter, both spatulate at tip. Ductus bursae wide, sclerotized. Lamellae post- and antevaginalis membranous, inconspicuous. Corpus bursae membranous, posterior part slightly striated, with two small, elongated sclerites, anterior part round, concentrically striated with stellate signum inside, stellar arms short. Distribution and abundance: Mediterranean. In Europe from the warmer parts of the Balkan peninsula across Italy incl. Sicily and Sardinia and southern France incl. Corsica to the Iberian peninsula. In Fauna Europaea (Hausmann et al. 2004; 2011a) erroneously recorded from Estonia. – Relatively common, sometimes occurring in numbers. – Outside Europe in Turkey and in Algeria (subsp. punctinervis), Morocco and the Canary Islands. Phenology: Univoltine: From early November to early May. 5th instar larvae in central Spain at 600 m in May (G. King pers. comm.). Hibernation as adult. Adults active at night, both sexes readily attracted to light. Biology: Larva oligophagous. Found on Dorycnium (D. Fritsch pers. comm.) and on Cistus salvifolius (Portugal: M. Corley pers. comm.). Recorded on Dorycnium pentaphyllum, Heli­ anthemum apenninum, Cistus incanus and C. monspeliensis (Flamigni et al. 2007). Reared on Cistus albidus (F. Dujardin in coll. ZFMK) and Helianthemum (B. Müller own data). Pupation among plant litter on the ground (M. Leipnitz pers. comm.; rearing observation). 335

Habitat: Xerothermophilous. In a variety of dry, open habitats, often abundant in costal dune habitats, in northern Greece in open Quercus forests at 800 m (F. Rosenbauer pers. comm.) and in Sierra Nevada (southern Spain) in bushy slopes at 1700 m (J. Gelbrecht pers. comm.), in central Spain in open scrubby landscapes on both gypsum and alluvial soils (G. King pers. comm.) and in Portugal in places where Cistus grows, such as patches of scrub on sand, limestone and on more acid soils, and in open woodland (M. Corley pers. comm.). In Italy up to 1100 m (Flamigni et al. 2007), in Bulgaria up to 350 m (S. Beshkov pers. comm.). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAF2548 (n=7 from southern Italy, Sicily, Bulgaria). Genetically heterogeneous, populations from Sicily diverging by 1.7% from the others. Nearest species: North American Parapheromia ficta (Rindge, 1972) (5.2%). Nearest species in Europe: ­Erannis declinans (5.7%), Tephronia sepiaria (5.8%). Remarks: Resting position of adults reminiscent of that of many Noctuidae, with wings partly folded on top of each other, being unusual for a geometrid species.

Fagivorina Wehrli, 1943 Fagivorina Wehrli, 1943, Gross-Schmett. Erde 4 (Suppl.): 524. Type species: Phalaena arenaria Hufnagel, 1767. There are no genus synonyms.

Diversity and distribution: Fagivorina is a monotypic genus, the only species occurs in Europe. External characters and abdomen: Antennae partially bipectinate to ca. 80% of length (about 8 flagellomeres free of branches) in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Vein R2 shortly anastomosing with R3 and R4. 336

Male genitalia: Uncus wide, rounded, weakly setose. Gnathos arms narrow, fused ventrally. Valvae dorsally wide, becoming narrower toward apex. Apically membranous, weakly setose. Medial ridge sclerotized, entire length with setae. Sacculus with sclerotized setose ridge, apically rounded, serrate. Juxta developed as large plate. Saccus wide, rounded. Aedeagus straight, apex acute. Vesica without cornuti. Female genitalia: Papillae anales membranous, striated, slightly setose, elongate. Apophyses posteriores long, straight. Apophyses anteriores shorter, ending spatulate. Corpus bursae membranous, slightly striated. Diameter of posterior part 1.5 times diameter of ductus bursae. Anterior part elongated. Signum oval, stellate. Ductus bursae short, narrow. Lamella postvaginalis wide, sclerotized, alate, irregularly striated. Centro-anteriorly forming a plate, posteriorly with two processes. Remarks: Fagivorina has not been included in a multi-gene analysis so far. In an extensive COI neighbour joining tree of Palaearctic Ennominae Fagivorina and Chemerina group together with a long-stalked common node (A. Hausmann own data).

114. Fagivorina arenaria (Hufnagel, 1767) Phalaena arenaria Hufnagel, 1767: Berlin Mag. 4 (5): 518 (Germany, Berlin region). Syntype(s). Geometra viduata Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s). ‡ Phalaena angularia Thunberg, 1792: Diss. ent. sistens Insecta Suecica 4: 59, pl. 4 (Sweden: Blekinge). Lectotype ♂ (UZIU). Junior primary homonym of Phalaena (Geometra) angularia Villers, 1789. Phalaena Geometra viduaria Borkhausen, 1794: Natur. eur. Schmett. 5: 172 (Europe). Syntype(s). Unavailable names (infrasubspecific): deumbrata: Lempke (1952) (ab.).

External characters and abdomen: Wingspan 25–34 mm. Wings strongly contrasted, marked with black, brown and sometimes olive-green on white ground. Forewings often with olive-green tinge, antemedial line curved, medial line thick, postmedial line black, distally with white edging, touching medial line near inner margin. Terminal line blackish, discontinuous. Discal spot linear, distinct, distal to medial line. Terminal and basal areas with a pale, dull, orange-brown band. Hindwings more homogeneous, discal spot less distinct. Fringes chequered. Frons, collar, thorax, white with dark scales. Abdomen with a double row of dark spots. Antennae and hindtibia, see genus description. Variation: Rather little. Amount of dark irroration and width of blackish markings are variable. Melanic forms are known (Urbahn 1941). Male and female genitalia: See genus description. Distribution and abundance: European. From eastern central Europe to France and from southern Scandinavia to Italy including Sicily and the Balkan peninsula including northern Greece. The species has disappeared from many of its earlier localities, especially towards the north of its distribution area. Erroneously recorded for Belgium and Netherlands by Leraut (2009), moreover showing a too large area in Denmark and Greece. – Local and not common, usually occurring in limited numbers. – Not recorded from outside Europe.

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Phenology: Uni- or bivoltine. In south-western Germany from late May to late July (Ebert 2003). In central and southern Italy in two generations from late April to mid-June and from late July to mid-September (Flamigni et al. 2016). Caterpillar from July to September. Overwintering as pupa. Biology: Larva oligophagous. Found and reared on young leaves of Quercus petraea, larvae refusing Fagus (Wegner 1998; 2013). Reared on Fagus and Quercus (Urbahn 1941). Erroneously recorded on lichens on Fagus and Quercus in various old literature, but also in Leraut (2009). Pupation among moss or in the ground (M. Leipnitz pers. comm.). Habitat: Silvicolous. In scattered and old Quercus petrea forests on dry soil (Wegner 1998; 2013) and old beech and mixed beech/oak forests on north-facing slopes on moist soil (­Urbahn 1939; Bergmann 1955). In Scandinavia from sea-level, in south-eastern Germany from 200 up to 1000 m (Ebert 2003), in the Alps from 300 up to 1500 m (Kerschbaum & Pöll 2010) and in Italy it can exceptionally reach 2000 m (Flamigni et al. 2016). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAD7665 (n=19 from Germany, Austria, Italy including Sicily, Croatia). Genetically very homogeneous in the main cluster, but populations from southern Italy diverging by 0.6%, those from Sicily by 1.1%. Nearest species: North American Parapheromia falsata McDunnough, 1920 (6.5%). Nearest species in Europe: Hypomecis roboraria (6.6%), Chemerina caliginearia (6.7%). Remarks: Wegner (2013) deals in details with the biology and habitat in north-western Germany.

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Spartopteryx Guenée, 1858 Spartopteryx Guenée, 1858, Hist. nat. Insectes (Spec. gén. Lépid.) 10: 121. Type species: Synopsia kindermannaria Staudinger, 1871. There are no genus synonyms.

Diversity and distribution: Monotypic, the only species occurring in the central and eastern Palaearctic region. External characters, abdomen and genitalia: See species description below. Remarks: Spartopteryx has not been included in a multi-gene analysis so far. In COI neighbour joining tree Spartopteryx and Athroolopha group aside with common node (A. Hausmann own data).

115. Spartopteryx kindermannaria (Staudinger, 1871) Synopsia kindermannaria Staudinger, 1871, in Staudinger & Wocke, Cat. Lepid. eur. Faunengeb. 1: 163, Syntype(s), [Russia] Altai. Unavailable names (infrasubspecific): fuscaria: Fernández (1932) (f.); tibetica: Wehrli (1941) (var.). – (Incorrect subsequent spelling): kindermanniaria: (Viidalepp 1988a; 1996).

External characters and abdomen: Wingspan ♂ 26–34 mm, ♀ about 30 mm. Males: Wings brownish grey, strongly irrorated with whitish grey and brown. Fringes chequered whitish and brownish grey. Wavy line distinct, whitish grey, slightly wavy. Terminal area between wavy line and terminal line usually unicolorous brownish grey, sometimes with more or less distinct pale blotch at centre. Ante- and postmedial lines of forewing whitish grey, usually shaded dark. Inner margin with a whitish grey streak between postmedial and wavy line, ending in the wavy line one-third from costa. Postmedial line with a stalked blackish triangle at centre. Hindwing with two blackish shaded lines. Discal spots absent. Underside greyish brown, terminal area of forewing partly, of hindwing entirely pale. Postmedial line on both wings with distinct whitish grey edging. Female brachypterous, with reduced, very narrow wings. Forewing very long, hindwing about 1/5 of forewing length. Frons, vertex, collar, tegula and thorax concolorous with ground colour, partly irrorated with darker scales. Abdomen dorsally with a double row of dark dots and a light midline. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Little in males. Variable in the extent of lighter blotches on the forewing. Male genitalia: Uncus triangular, tip shortly bent. Gnathos arms long, narrow, ventrally fused, forming a rounded plate. Valva simple, membranous, thicker at base, with smooth margins; costa weakly sclerotized, weakly setose; apical area rounded, setose. Juxta constricted at centre, posteriorly with sclerotized tip. Saccus rounded. Aedeagus sub-cylindrical, four times as long as wide. Vesica with one cornutus of half aedeagus length. Female genitalia: Papillae anales very long, sclerotized, posteriorly membranous, setose. Apophyses posteriores the longest of European ennomines, length on average 8.8 mm. Apophyses anteriores shorter, on average 3.5 mm, both spatulate at tip. Lamella antevaginalis with two membranous, tapered ‘wings’, lamella postvaginalis a strongly sclerotized, anteriorly rounded plate. Ductus bursae narrow. Corpus bursae mostly membranous. Posterior half cylindrical, with some weakly sclerotized striae. Anterior half drop-shaped with a small, stellate signum.

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Distribution and abundance: East Palaearctic. In Europe only recorded in a few localities slightly west of southernmost Ural mountains. – Local in the Urals, now and then found in numbers (K. Nupponen pers. comm.). – Outside Europe widespread from Kazakhstan through southern Siberia, Mongolia, Dahuria and China to Far East Russia; to the south in parts of China and in northern Tibet.

Phenology: Univoltine. From early to late June, in the Altai mountains from early June to mid-July (J. Gelbrecht pers. comm.). Larval time and hibernation stage apparently unknown. Adults active at night, especially males attracted to light. Only few females found in collections. Males found during daytime when disturbed from trunks of large deciduous trees and vertical rocky walls on a river shore (K. Nupponen pers. comm.). Biology: Unknown. Habitat: In southern Ural mountains in a restricted forest steppe by a riverside, surrounded by mixed forest (K. Nupponen pers. comm.). In the Altai mountains in warm, rocky and bushy slopes (J. Gelbrecht pers. comm.). In the Urals at low altitudes, up to 450 m (K. ­Nupponen pers. comm.), outside Europe from 700 up to 2800 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAH0730 (n=7 from eastern Russia, south-eastern Kazakhstan). Genetically homogeneous. Nearest species: Jankowskia athleta Oberthür, 1884 from southern Kazakhstan, Russia and Mongolia (3.0%), genetic data suggesting the two species to be congeneric.

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Athroolopha Lederer, 1853 Athroolopha Lederer, 1853, Verh. zool.-bot. Ver. Wien 3 (Abh.): 179, 227,230. Type species: Geometra pennigeraria Hübner, 1813. There are no genus synonyms.

Diversity and distribution: The genus Athroolopha contains two or three species (see Remarks) in the West Palaearctic (West Mediterranean) region. External characters and abdomen: Large-sized moths. Forewing dark greyish brown with whitish pattern. Hindwing orange with dark terminal bands. Discal spots black, on hindwing sometimes weak. Frons, collar, thorax and abdomen dark grey, with scattered lighter scales. Antennae partially bipectinate in male, filiform in female. Hindtibia with 2+2 spurs. Male genitalia: Uncus triangular, tapered, beak-shaped. Gnathos broad, ventrally fused. Valva near apex and ventrally membranous, wider at base, apex rounded. Costa more strongly sclerotized with inwardly directed setose ridge. Juxta shield-shaped in ventral view. Aedeagus short, proximally thicker, without cornuti. Female genitalia: Papillae anales developed as a long tube, distally soft, membranous, slightly setose. Apophyses posteriores very long, thin, length on average 10 mm (including ovipositor), straight, at tip slightly thickened. Apophyses anteriores shorter, 2.7–3.4 mm, at tip spatulate (n=3). Bursa very small, hyaline, without signum. No differences between A. pennigeraria and A. chrysitaria. Remarks: Athroolopha has not been included in a multi-gene analysis so far. In a COI neighbour joining tree of Palaearctic Ennominae Spartopteryx and Athroolopha cluster together on a common node (A. Hausmann own data).

116. Athroolopha pennigeraria (Hübner, 1813) Geometra pennigeraria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1), pl. 70, fig. 363 (Europe). Syntype(s). Fidonia chrysitaria var. kabylaria Oberthür, 1878: Études ent. 3: 47, pl. 5, figs 6a, b (near Collo). Syntype(s). Validated at subspecific rank (Scoble 1999), see remarks below. Athroolopha chrysitaria latimargo Rothschild, 1914: Novit. zool. 21: 351 (Central Algeria: GueltEs-Stel). Syntypes 20♂♀ (NHMUK). Valid at subspecific rank (Scoble 1999). Fidonia pennigeraria atlanticaria Le Cerf, 1923: Bull. Soc. ent. Fr. 1923 (15): 199 (Morocco: Oulmés). Holotype ♂ (MNHN). Valid at subspecific rank (Scoble 1999). Fidonia pennigeraria maghrebina Rothschild, 1925: Bull. Soc. Sci. nat. Maroc 5 (4/5): 149. Unnecessary replacement name for atlanticaria Le Cerf (Scoble 1999). Fidonia pennigeraria orientalis Rothschild, 1925: Bull. Soc. Sci. nat. Maroc 5 (4–5): 149 (Algeria: Province of Constantine, Hauts Plateau, Batna; Khenchela; Constantine etc.). Syntypes ♂♀ (NHMUK). Downgraded from subspecific rank (Scoble 1999) to synonymy by Leraut (2009).

External characters and abdomen: Wingspan 30–40 mm. Forewings: Ground colour dark brown, with scattered orange and whitish scales. Wavy line, post- and antemedial lines white, thick, zigzagging, partially broken into spots. Postmedial line with teeth. Antemedial line with an acute angle. Hindwings orange with dark brown terminal band from apex to tornus and somewhat lighter medial line. Area along anal margin darker, with scattered orange scales. Fringes chequered dark brown and whitish. Hindwing brown with white patterns. Underside: Ground colour of male orange and whitish, speckled brown, on apex of forewings and along anal margin of hindwings dark brown. Ground colour of female 341

­ indwings dark brown with pattern. Frons, collar, thorax, abdomen and hindtibia, see geh nus description. Male antennae partially bipectinate with long branches, apical 6–8 flagellomeres with very short branches. Antennae filiform in female. Variation: Pattern somewhat variable, especially extent of white blotches on forewings. Terminal band on hindwings can be smaller or even absent. Male genitalia: Smaller than in A. chrysitaria. Uncus triangular, tapered, beak-shaped, weakly setose. Gnathos arms ventrally fused, small (arms wider in A. chrysitaria). Valva near apex and ventrally membranous, wider at base, apex rounded, setose. Valva narrower than in A. chrysitaria. Costa strongly sclerotized with a setose ridge, costa and ventral margin more concave than in A. chrysitaria. Sacculus at base sclerotized, smooth, apically more membranous, setose. Saccus small sclerotized ring with dorsally bent tip. Shield-shaped ­juxta posteriorly ending ring-shaped, anteriorly with two sclerotized, tapered lobes (at a ­wider distance than in A. chrysitaria) and a rounded triangular projection. Aedeagus short, narrower towards apex, caecum ball-shaped, thick (smaller in A. chrysitaria). Without cornuti. Female genitalia: See genus description. Distribution and abundance: West Mediterranean. Distributed through most of the Iberian peninsula and through southern France with a single record in north-western Italy. – Somewhat local, but often abundant in its localities. – Outside Europe in Morocco (subsp. atlanticaria) and Algeria (subsp. latimargo). Mentioned from Tunisia by Redondo et al. (2009), but no record from there has been traced.

Phenology: Univoltine: In Spain from early May to late June, until July at high altitudes. In southern France in June and July (Robineau et al. 2007). Caterpillars found in late March at 1150 m in central Spain (G. King pers. comm.), in late April and early May (J. Gelbrecht 342

pers. comm.). Males flying by day in sunshine. Females sitting on the ground in sunshine and flying only for short distances when flushed up (M. Leipnitz pers. comm.). Rarely, the females can be seen flying actively by day (K. Nupponen pers. comm.). Overwintering as larva. Biology: Larva polyphagous. Found on Artemisia (J. Gelbrecht & T. Drechsel pers. comm.). Recorded on Lavendula, Halimium lasianthum and Rosmarinus officinalis (­Redondo et al. 2009). Reared on Artemisia camphorata and A. abrotanum (M. Leipnitz pers. comm.). Pupa­tion in a loose cocoon among plant parts and moss on the ground (M. Leipnitz pers. comm.; rearing observation). Habitat: Xerothermophilous. In a variety of open habitats or habitats with scattered trees and bushes. From sea-level up to 1800 m. Similar species: A. chrysitaria with lines of forewings continuous, less sinuose than in A. pennigeraria. Terminal band of A. pennigeraria extended over the whole hindwing, just weaker on anal margin, whilst in A. chrysitaria not reaching the anal and costal margins. Genetic (5.4%).

data:

BIN: BOLD:AAO0991 (n=1 from Spain). Nearest species: A. chrysitaria

Remarks: Wehrli (1939–54) and Leraut (2009) consider A. kabylaria (Oberthür, 1878) from North Africa as a bona species, requiring further study.

117. Athroolopha chrysitaria (Hübner, 1831) Geometra chrysitaria Hübner, 1831: Samml. Eur. Schmett. 5 Geometrae (1), pl. 107, figs 557, 558 (Italy: Sicily). Syntype(s).

External characters and abdomen: Wingspan 31–41 mm. Forewings: Ground colour dark greyish brown, with scattered orange and whitish scales. Wavy line slightly wavy, clearer towards costa. Postmedial line with two teeth, antemedial line with one tooth, both thick, whitish. Medial area often darker. Hindwings orange with greyish brown terminal band along the outer margin from tornus, ending before reaching the apex, with scattered orange scales. Fringes chequered brown and whitish orange. Ground colour of underside orange, brown speckled. Basal area dusted with black scales. Frons, collar, thorax, abdomen and hindtibia see genus description. Male antennae partially bipectinate with long branches, the apical 6–7 flagellomeres with very short branches. Antennae filiform in female. Variation: Little. Extent of white forewing pattern variable. Greyish brown terminal band along outer margin more or less distinct. Male genitalia: Larger than in A. pennigeraria. Uncus wide, triangular, weakly setose, tip bent beak-shaped. Gnathos arms wide, fused ventrally, granulate (arms smaller in A. penni­ geraria). Valva near apex and ventrally membranous, base extended, apex rounded. ­Costa sclerotized, subapically setose over half of valva length. Sacculus at base sclerotized, smooth, apically more membranous, setose. Valva wider than in A. pennigeraria, dorsal margin slightly concave. Saccus tapered, tip dorsally bent, longer than in A. pennigeraria. Shield-shaped juxta posteriorly forming a ring, anteriorly with two sclerotized, tapered lobes (at smaller distance than in A. pennigeraria) and a rounded triangular projection. Aedeagus short, narrower towards apex, caecum round (wider in A. pennigeraria). Without cornuti. Female genitalia: See genus description.

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Distribution and abundance: Central Mediterranean, endemic to Sicily. In Europe only known from Sicily, occurring in a limited area in the north-west of the island. – Very local and not common.

Phenology: Univoltine: From early May to early July (Flamigni et al. 2007). Hibernation stage unknown, but probably as larva as in A. pennigeraria. Males flying by day in sunshine, especially in the morning until eleven o’clock, while females are rarely seen. Biology: Larva monophagous. Found on Erica multiflora (M.  Romano & P. Mazzei pers. comm.). Recorded on Erica arborea (Flamigni et al. 2007). Habitat: Xerothermophilous. In various open land localities or localities with scattered trees and bushes (M. Romano & P. Mazzei pers. comm.). In Sicily from sea-level up to 1500 m (Flamigni et al. 2007). Similar species: A. pennigeraria with lines of forewings more strongly angled. Terminal band of hindwing of A. chrysitaria does not reach the anal and costal margins, in A. pennigeraria extended along the whole margin with various intensity. Male genitalia of A. pennigeraria with tip of gnathos smaller, valvae apically narrowing, juxta distally with two tapered sclerotized lobes. Proximal end of the aedeagus thickened. Genetic (5.4%).

344

data:

BIN: BOLD:ADL9523 (n=1 from Sicily). Nearest species: A. pennigeraria

Eurranthis Hübner, 1823 Eurranthis Hübner, [1823] 1816, Verz. bekannter Schmett.: 296. Type species: Phalaena plummistaria Villers, 1789. Genus synonyms after Scoble (1999). Fidonia Treitschke, 1825, in Ochsenheimer, Schmett. Eur. 5 (2): 435. Type species: Phalaena plummistaria Villers, 1789. Phytonia Sodoffsky, 1837, Bull. Soc. imp. Nat. Moscou 1837 (6): 90. [Unnecessary replacement name for Fidonia Treitschke].

Diversity and distribution: The genus Eurranthis contains one West Mediterranean species and two more species in South Africa, which probably do not belong to this genus. External characters, abdomen and genitalia: See description below (based on type species). Remarks: In multi-gene analysis Eurranthis and Nychiodes are sister taxa (Murillo-Ramos et al. 2019). The close relationship is supported by the genitalia, too (this volume).

118. Eurranthis plummistaria (Villers, 1789) Phalaena Geometra plummistaria Villers, 1789: Linn. ent. 2: 326 (France). Syntype(s). Phalaena Geometra plumistaria Borkhausen, 1794: Natur. eur. Schmett. 5: 174. Emendation of plummistaria Villers. Geometra auritaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 81, fig. 416 (­Europe). Syntype(s). Eurranthis plummistaria var. albosignata Neuburger, 1907: Societas ent. 22 (1): 2 (Portugal: Soalheiro). Syntypes 2♂. Validated at subspecific rank (Scoble 1999), downgraded to synonymy (“f.”) by Leraut (2009), here confirmed based on clinal transitions and absence of constant and significant differential features. Eurranthis plummistaria atlanticaria Lucas, 1920: Bull. Soc. ent. Fr. 1920: 254 (Morocco: near Meknès, 1800 m). Syntypes 3♂1♀. Valid at subspecific rank (Scoble 1999). Fidonia plumistraria moriscaria Le Cerf, 1923: Bull. Soc. ent. Fr. 1923 (15): 199 (Morocco: ­Azrou). Holotype ♂ (MNHN). Fidonia plummistaria algeriensis Rothschild, 1925: Bull. Soc. Sci. nat. Maroc 5 (4/5): 150 (Alge­ ria: Prov. of Constantine, Djebel Mahmel; near Batna). Syntypes ♂♀. Valid at subspecific rank (Scoble 1999). Fidonia plumistaria var. acronevadaria Wehrli, 1926: Ent. Z., Frankf. a. M. 39 (40): 163 (Spain: Castilla, Sierra Nevada, 2600 m). Syntypes 25♂17♀. Fidonia (Eurrhanthis) concolor Schwingenschuss, 1935: in Zerny, Mém. Soc. Sci. nat. Phys. ­Maroc 42: 91, pl. 1, fig. 30 (Morocco: Great Atlas, Tachdirt, 2700 m). Holotype ♂. Unavailable names (infrasubspecific): albicans: Oberthür (1878) (ab.); confluens: Oberthür (1878) (ab.).

External characters and abdomen: Very characteristic species. Wingspan ♂ 32–40 mm, ♀ 29–32 mm. Ground colour of forewing orange with white, of hindwing orange. Medial lines black, thick, partly forming spots. Terminal area of forewing with row of orange spots, in hindwing black spots. Postmedial and medial lines of forewing touching on inner margin. Antemedial line on forewing curved, on hindwing absent. Fringes black. Ground colour of underside whitish on hindwing, orange on forewing, pattern similar. Both wings with discal spot. Frons, collar, thorax with long hairy scales, black with little orange. Abdomen black with orange spots. Antennae bipectinate to tip in male, branches very long, filiform in female. Hindtibia of both sexes with 2+2 spurs.

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Variation: Wing colour varying from off-white to yellow or yellow-orange. On forewing intensity of black lines and spots and extent of white and orange areas varies considerably. High mountain specimens of Spanish Sierra Nevada small (wingspan 26–30 mm), wings with extended white and pale orange parts (similar to ‘f. albosignata’). Male genitalia: Uncus sclerotized, triangular, tip sclerotized, rounded, laterally concave. Gnathos narrow, strongly sclerotized, fused, ventrally upturned. Valva wide, short. Costa sclerotized, apex small, setose. Sacculus with two central projections: one pointed, bent, other finger-shaped with setose apex. Saccus flattened. Juxta plate, posteriorly with long narrow projection. Aedeagus thick, slightly tapered, vesica with long cornutus of 4/5 aedeagus length. Female genitalia: Papillae anales slightly sclerotized, setose, nearly round. Apophyses posteriores narrow, straight, length 2.2 mm. Apophyses anteriores stouter, little more than half length of the latter (1.3 mm). Corpus bursae membranous, anterior part oval, signum weak, stellate. Posterior part tubular with globular diverticulum, with origin of ductus seminalis. Lamella antevaginalis small, weakly sclerotized, tapered on both sides. Lamella postvaginalis sclerotized plate, margin rounded, posteriorly concave. Distribution and abundance: West Mediterranean. In Europe from the Iberian peninsula through southern France to north-western Italy. – Relatively common, usually found in limited numbers. – Outside Europe in Morocco (subsp. atlanticaria) and Algeria (subsp. algeriensis).

Phenology: Univoltine: From early March to early June, depending on altitude, sometimes even until late June. In the Iberian peninsula mainly from April to June (Redondo et al. 2009). In Morocco from March to May (Rungs 1981). Larva from April to June, depending 346

on altitude (M. Leipnitz pers. comm.). Small larvae found in southern France in late May (T. Drechsel pers. comm.). Hibernation as pupa. Adults fly by day in sunshine, especially before noon, when mainly males are found. Females sitting in low vegetation (T. Varenne pers. comm.) or on the ground in sunshine, flying only over short distances when flushed up (M. Leipnitz pers. comm.). Biology: Larva monophagous. Found on Dorycnium (T. Drechsel pers. comm.). Recorded on Dorycnium pentaphyllum (Flamigni et al. 2007; Redondo et al. 2009). Reared on Dorycnium germanicum and D. hirsutum (M. Leipnitz pers. comm.). Pupation in a loose cocoon among plant litter on the ground (rearing observations). Habitat: Xerothermophilous. In open uncultivated places, rocky slopes, places with scattered bushes and trees. In southern Spain on calcareous ground, in open Mediterranean vegetation; in southern France in bushy slopes with calcareous low-nutrient meadows (J. Gelbrecht pers. comm.). From sea-level up to 800 m in north-western Italy (Flamigni et al. 2007), in Spain up to 2770 m in the Sierra Nevada (Redondo et al. 2009). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAL5357 (n=6 from France, Spain, Morocco, including holotype of atlanticaria). Genetically homogeneous. Nearest species: North American Parapheromia ficta (Rindge, 1972) (6.1%), one of the two (putative) congeners, E. acuta (Warren, 1900) from South Africa at a distance of 13.4%, possibly requiring the description of a separate genus.

Nychiodes Lederer, 1853 Nychiodes Lederer, 1853, Verh. zool-bot. Ges. Wien, 3: 177, 216, 219. Type species: Geometra lividaria Hübner, 1799 [Europe]. Genus synonyms after Scoble (1999). Comeesia Wehrli, 1941, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 440. Type species: Nychiodes admirabila Brandt, 1938. Eunychiodes Wehrli, 1941, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 441. Type species: Boarmia amygdalaria Herrich-Schäffer, 1948. Fritzwagneria Wehrli, 1941, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 438. Type species: Nychiodes dalmatina Wagner, 1909.

Diversity and distribution: The genus contains 27 species, distributed in the Palaearctic region from western Europe and North Africa (Morocco to Tunisia) into Iran, Afghanistan and Pakistan. Non-European species of this genus need taxonomic revision urgently. External characters and abdomen: Large-sized, wingspan 30–50 mm, females slightly larger than males. Wings light brown or beige, to dark grey or different shades of brown, markings blackish. Wing termen slightly dentate. Forewing ante- and postmedial lines prominent, ­medial line often partially present. Outer margin of wings in most species slightly darker. Discal spots usually present on all wings, medium-sized, dark, distinct. Hindwing post­medial line blackish, medial line partially visible. Frons, thorax and abdomen concolorous with wings; chaetosemata as two separate patches. Collar covered with thin blackish transverse line. Antennae covered dorsally by a mix of yellowish and brown scales, bipectinate in both sexes; branches long in male, short in female. Proboscis completely absent. Epiphysis long (as long as foretibia). Spur formula 0–2–4. Abdominal segments in both sexes without modifications.

347

Male genitalia: Uncus apically acute, subapically triangular. Gnathos well sclerotized, at centre tongue-shaped, covered with tiny spines. Valva split into dorsal and ventral part; costa of valva partially or completely sclerotized, sometimes medially humped; in some species apex of valva with digitiform process (see below). Ventral part of valva ornamented with three processes: ampulla superior (setose); ampulla inferior (depending on species setose or not); sacculus process (presence and form diagnostic in different species), not setose. Juxta anchor-shaped, width of apical and basal part diagnostic. Aedeagus generally large with needle-shaped cornutus, relative lengths of cornutus versus aedeagus diagnostic in different species-groups (see below). Female genitalia: Generally large. Ovipositor large-sized, round or ovally elongated. Relative lengths of apophyses anteriores and posteriores diagnostic to species (within species-groups). Lamella postvaginalis generally sclerotized, and diagnostic to species-groups (see below). Ductus bursae membranous, but in some species partially sclerotized. Corpus bursae membranous; signum stellate (i.e. round, dentate at margins), located on dorsal part of corpus bursae. Classification: Wehrli (1929a; 1929b; 1929c) tentatively classified the known Nychiodes species on the base of male genitalia structures into three species-groups (amygdalaria group, dalmatina group and obscuraria group). Following on from this system, he later tried to split up the genus into four genera (Wehrli 1941). This division was not followed by ­other authors (e.g. Fazekas 1996; 1997; Expósito Hermosa 1984; 1985a; 1989; Rákosy 1997; Rákosy & Bere 1996; Rákosy & Szekely 1993; Leraut 2009; Redondo et al. 2009), and the additional taxa were regarded as synonyms of the genus Nychiodes by Scoble (1999) as well. The present study shows that some of the species are not satisfactorily grouped by Wehrli (1929a; 1929b; 1929c; 1941), e.g. according to the male and female genitalia dalmatina and waltheri cannot be regarded as belonging to the same group, and the former is more similar to amygdalaria. To avoid any confusion, and only for identification purposes, we tentatively classify the European Nychiodes species into three morphological groups here: obscuraria species-group, amygdalaria species-group and waltheri species-group. Identification of most European Nychiodes species based on wing pattern is difficult and for a reliable identification, dissection of genitalia (ideally male) is recommended. Diagnostic characters in genitalia of these three species-groups are listed and illustrated as follows: obscuraria species-group Species included: Nychiodes obscuraria (Villers, 1789); N. ragusaria Millière, 1884; N. anda­ lusiaria Millière, 1865; N. notarioi Expósito, 2005 and N. hispanica Wehrli, 1929. The species of this group are distributed in central and western Europe, and in North-west Africa. Male genitalia: Costal part of valva sclerotized only up to the subapical part, medially humped. A digitiform extension on the apex of valva present, covered with thin setae. Ampulla superior setose, ampulla inferior and sacculus process acute, strongly sclerotized, not setose. Aedeagus thick and stout, needle-shaped cornutus at least half length of aedeagus. Female genitalia: ovipositor short, rounded. Apophyses anteriores half length of apophyses posteriores. Lamella postvaginalis strongly sclerotized, extended laterally. Ductus bursae long, membranous, or partially sclerotized, of same length as corpus bursae. The latter always membranous. Signum stellate, corpus bursae small and roundish.

348

Text-figs 148–150. Diagnostic characters of male genitalia (indicated) of European Nychiodes species-groups. Text-fig. 148. obscuraria group (N. obscuraria), Text-fig. 149. waltheri group (N. waltheri), Text-fig. 150. amygdalaria group (N. amygdalaria).

349

Text-figs 151–153. Diagnostic characters of female genitalia (indicated) of European Nychiodes species-groups. Text-fig. 151. obscuraria group (N. obscuraria), Text-fig. 152. waltheri group (N. waltheri), Text-fig. 153. amygdalaria group (N. amygdalaria).

350

waltheri species-group Species included: Nychiodes waltheri Wagner, 1919 and N. aphrodite Hausmann & Wimmer, 1994 (non-European). The species of this group are distributed only in eastern Europe into the Middle East. Male genitalia: Costal part of valva sclerotized only up to the subapical part. Digitiform extension on the apex of valva absent. Ampulla superior and ampulla inferior apically covered with spines. Sacculus process present, acute, strongly sclerotized, not setose. Aedeagus generally shorter than in obscuraria and dalmatina groups, length of the needle-shaped cornutus half length of aedeagus. Female genitalia: Ovipositor short, rounded (in contrast to amygdalaria group). Apophyses anteriores very short (1/3 length of apophyses posteriores). Lamella postvaginalis sclerotized, conically extended. Ductus bursae very short, partially membranous. Signum stellate, corpus bursae small and roundish. amygdalaria species-group Species included: Nychiodes amygdalaria (Herrich-Schäffer, 1848) and N. dalmatina ­Wagner, 1909. The species of this group are distributed in eastern Europe and in the Middle East. Male genitalia: Costa of valva sclerotized up to and over the apical part. Valva subapically spinose. Digitiform extension absent from apex of valva. Ampulla superior and ampulla inferior apically covered with spines. Sacculus process absent. Aedeagus generally narrower than in obscuraria group, needle-shaped cornutus maximum half length of aedeagus. Female genitalia: Ovipositor ovally elongated (in contrast to the other two groups). Apophyses anteriores very short (1/3 length of apophyses posteriores). Lamella postvaginalis sclerotized, extended anteriorly and posteriorly. Ductus bursae long, partially or fully sclerotized. Signum stellate, corpus bursae large and elongate. Phenology: Normally uni- or bivoltine (even trivoltine in warmer habitats). Adults without proboscis, therefore not attracted to bait, but to light. Females are very inactive and prefer not to fly (M. Leipnitz pers. comm.). Biology: As far as known, larvae of Nychiodes species live on species of Rosaceae family (Fazekas 1996; Flamigni et al. 2007; Redondo et al. 2009; Taurand 2009). However, larva of N. anda­lusiaria on Genista (Fabaceae), see page 358, N. obscurata collected on Prunus spinosa, N. dalmatina on Cotoneaster racemiflorus and Crataegus sp., N. aphrodite on Crataegus sp. In captivity larvae were reared on other Rosaceae: Prunus spinosa, P. domestica, Cotoneaster sp., Amelanchier sp., Crataegus sp., Pyracantha sp. Habitat: Moths are found on overgrown rocks, rocky walls and in rocky valleys covered with small- to middle-sized food-plants, dense hedges, and stony grounds covered with shrubs and meadows, acre, wild barren lands, open meadows, also overgrazed areas and open forests with small plants (M. Leipnitz pers. comm.). Immature stages (Unpublished data from M. Leipnitz pers. comm.): Female hides the eggs in the ground or between leaves and soil as singletons or in small groups. Sometimes eggs are attached to the leaves, branches or fruits of the food-plant. Eggs are relatively large. Freshly laid eggs are normally light to dark turquoise, after some days becoming dark orange to red-brown and finally before hatching grey to anthracite-grey. Larva light brown to anthracite-grey with warts, with blackish spots laterally and dorsally, and yellow spots laterally on 351

the abdominal segments. Anterior part of prothorax has a characteristic yellow to orange ring. A unique behavioural observation in studied Nychiodes larvae was the excretion a light yellow to creamy coloured fluid just before prepupation and in pupation phases. This fluid stained the last segments of the prepupa to light yellow or creamy. Overwintering as young larva (L1 or L2). The pupa, depending on species, is red-brown to blackish brown. Pupation in the soil, without any cocoon. Genetic data: Judged from the available mtDNA data (COI barcodes), the group of East Mediterranean species (including all non-European species!) shows very little relationship with the other group including all species from the Central and West Mediterranean region. Remarks: In multi-gene analysis Eurranthis and Nychiodes are sister taxa (Murillo-Ramos et al. 2019). The close relationship is supported by the genitalia also (this volume).

119. Nychiodes obscuraria (Villers, 1789) Phalaena (Geometra) obscuraria Villers, 1789: Linn. ent. 2: 325 (southern France). Neotype ♂ (ZFMK, designated by Fazekas (1997), examined). Phalaena (Geometra) obscurata Borkhausen, 1794: Natur. eur. Schmett. 5: 537. Emendation of obscuraria Villers. Geometra lividaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 26, fig. 141 (­Europe). Syntype(s). Junior synonymy proposed by Prout (1915). Nychiodes lividaria coloxaria Constantini, 1916: Atti Soc. Nat. Modena (5) 3: 17 (Italy: M ­ onte Gibbio; Castelvetro; Ligorzano). Syntype(s). Junior synonym of N. obscuraria (Fazekas 1997), confirmed by Leraut (2009) and Flamigni et al. (2007), here confirmed. Nychiodes obscuraria teriolensis Wagner, 1927: in Schwingenschuss & Wagner, Z. öst. EntVer 12 (7): 69 (southern Dalmatia; Tyrol). Syntype(s). Junior synonym of N. obscuraria (Fazekas 1997), confirmed by Leraut (2009) and Flamigni et al. (2007), here confirmed. Nychiodes obscuraria ticina Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 437, pl. 37c (Switzerland: Tessin near Biasca). Lectotype ♂, Paralectotype ♀ (ZFMK, designated by Fazekas (1997), externally examined). Junior synonym of N. obscuraria (Fazekas 1997), confirmed by Leraut (2009) and Flamigni et al. (2007), here confirmed.

External characters and abdomen. Wingspan ♂ 37–49 mm, ♀ 40–51 mm. Wings grey-brown to dark beige, terminal area of wings slightly darker, markings dark grey to black. Forewing antemedial line curved outwards; medial line completely absent (important diagnostic character of N. obscuraria from other European congeners); postmedial line angled outwards on M1, curved slightly inwards, ending at centre of forewing inner margin (dorsum). Forewing subterminal area covered with creamy scales, replaced by brown scales towards costa; wavy line very thin, finely dentate, in some specimens hardly visible; outer margin (termen) dark brown to grey. Basal, antemedial and medial lines absent from hindwing; postmedial line straight or slightly curved. Hindwing subterminal area covered with creamy scales; wavy line hardly visible. Discal spots blackish, sometimes faint on forewing. Terminal line of all wings blackish, narrow, strongly undulating, continuous but stronger between vein endings. Fringes concolorous with wings, slightly darker at vein endings. Underside of wings in the basal half light brown, outer/terminal area dark brown, diffuse. Only postmedial line visible on underside, as narrow blackish line bordered with pale stripe. Variation: The species shows high variation in size and colour (from light brown to dark forms). The distance between forewing’s medial lines varies in different individuals. Additionally the degree of light irrorations on wings varies. 352

Male genitalia: Dorsal part of valva widely extended. Only basal part of costa of valva sclerotized, medially slightly humped. Digitiform extension on the apex of valva present, covered with thin setae. Ampulla superior clubbed, covered with tiny stout setae; ampulla inferior and sacculus process star-shaped, strongly sclerotized, not setose. Ampulla superior and inferior very close together. Sacculus process strongly reduced. Juxta anchor-shaped, apically concave. Aedeagus larger than in all other species of the obscuraria group, needle-shaped cornutus only slightly shorter than aedeagus, curved at centre. Female genitalia: Genitalia large. Lamella postvaginalis at centre less sclerotized, laterally with stronger sclerotization, here folded and with arch-shaped extension. Ductus bursae membranous, longitudinally striated, of the same length as the membranous corpus bursae. Signum stellate, small. Distribution and abundance: Central Mediterranean: southern Europe (eastern France, Italy, southern Switzerland, Slovenia, Croatia and a fragmented population in Hungary (Fazekas 1997; Flamigni et al. 2007; Leraut 2009; Fauna Europaea: Hausmann et al. 2004; 2011a; SwissLep Team 2010). Reputedly in Greece (Müller 1996). Records from the eastern Pyre­ nees and Spain (specimens deposited in SMNS) probably are subject of mislabelling and cannot be confirmed here. – Occurring in numbers. – Not recorded outside Europe.

Phenology: Uni- or bivoltine: From June to August in single brood populations and additionally in September in populations with second generation. Attracted to UV light, but also found in daylight Taurand (2009), being very sensitive and are easily flushed up when somebody walks nearby. Overwintering as larva (Taurand 2009).

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Biology: Larva oligophagous on Rosaceae. Found on Prunus spinosa. Reared on Cotoneaster sp., Amelanchier sp., Crataegus sp., Prunus spinosa, P. domestica, Pyracantha sp. (M. Leipnitz pers. comm.) Habitat: Xerothermophilous, preferring warm and dry habitats with the hostplants (Flamigni et al. 2007). From sea-level up to 1500 m. Similar species: In contrast to all other species in this group, in N. obscuraria the medial line is absent from the forewing. Additionally, the hindwing postmedial line is straight or slightly curved (same condition in N. notarioi, angled in N. hispanica and N. andalusiaria, curved in N. ragusaria) (see Text-figs 154–158). In male genitalia, the short ampulla inferior very close to ampulla superior and the strongly reduced sacculus process are characteristic for N. obscuraria. Ampulla superior clubbed in N. obscuraria (comparable with N. ragusaria, but tubular or thumb-shaped in all other species of this group). Relative width of apical versus basal part of juxta 1:1.5 (1:1.5 in N. andalusiaria; 1:1 in N. ragusaria and N. notarioi; 1:2 in N. hispanica). In female genitalia, lamella postvaginalis less sclerotized at centre (well sclerotized in all other species of the obscuraria group). Ductus bursae membranous (partially sclerotized in all other species of the obscuraria group). As this species shows high colour variation, we recommend genitalia dissection and/or DNA barcoding for certain identification. Especially male genitalia structures allow reliable identification. Genetic data: BIN: BOLD:ACE5020 (n=5 from south-eastern France, northern, central and southern Italy, including one specimen from central Calabria). Genetically homogeneous. Nearest species: N. ragusaria (1.2%).

120. Nychiodes ragusaria Millière, 1884 Nychiodes lividaria var. ragusaria Millière, 1884: Naturalista sicil. 3 (7): 196, pl. 3, fig. 1 (Italy: Sicily, Castelbuono). Neotype ♂ (ZFMK, designated by Fazekas (1997), examined). Nychiodes bellieraria Ragusa, 1884: Naturalista sicil. 3 (12): 352, pl. 3, fig. 2 (Italy: Sicily, Castelbuono). Junior synonym of N. ragusaria proposed by Fazekas (1997), confirmed by Flamigni et al. (2007).

External characters and abdomen. Wingspan ♂ 32−48 mm, ♀ 36−49 mm. Wings light to dark brown, irrorated with red-brown scales on costal area, markings blackish, faint towards costa. Forewing antemedial line curved outwards near costa, partially visible; medial line present, often well developed towards inner margin (completely absent in N. obscuraria); postmedial line angled outwards on M1 vein, curved slightly inwards, ending at middle of inner margin. Forewing subterminal area partially covered with yellow scales as linear shadow, replaced by brown scales towards costa; wavy line faint, hardly visible, yellowish. Hindwing basal and antemedial lines absent; medial line dull, hardly visible; postmedial line slightly angled at M2 vein (as in N. andalusiaria, whilst straight or slightly curved in N. obscuraria and N. notarioi, outwards extended in N. hispanica). Hindwing subterminal area partially covered with creamy scales; wavy line hardly visible. Tiny discal spots blackish, hardly visible on hindwings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings with yellow shadow. Fringes brown to grey, slightly darker on vein endings. Underside of wings dull, beige to dark brown, outer area slightly darker. Postmedial line partially visible as a yellow line. Variation: Wing colour fairly variable, as well as the markings. Sometimes wings markings fully faint. 354

Text-figs 154–158. Diagnostic characters of the European species of the obscuraria species-group. Text-fig. 154: N. obscuraria (Villers, 1789), Italy: South Tirol, Vinschgau, 16.7.1976. Text-fig. 155: N. andalusiaria Millière, 1865, Spain: Sierra de Gredos, 21.7.2003. Text-fig. 156: N. nota­ rioi Expósito, 2005, Spain: Sierra del Montseny, 1.7.1994. Text-fig. 157: N. hispanica Wehrli, 1929, Spain: Sierra Nevada, 12.7.2009. Text-fig. 158: N. ragusaria Millière, 1884, Italy: Coccorino, 15.6.2008.

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Male genitalia: Valva extended, its dorsal part triangular, apically merged with digitiform extension (important diagnostic character against all other species of obscuraria group), covered with thin setae. Only the basal part of the costa of valva sclerotized, basally humped. Ampulla superior clubbed (comparable with equivalent in N. obscuraria, whilst tubular or thumb-shaped in all other species of this group), apically covered with tiny spines. Ampulla inferior and sacculus process very short, apically blunt, strongly sclerotized, not setose; valva between these two processes strongly concave (diagnostic character against all other European species of the genus Nychiodes). Juxta anchor-shaped, width of apical part corresponding to basal part (1:1) (1:1 in N. notarioi; 1:1.5 in N. andalusiaria and N. obscuraria; 1:2 in N. hispanica). Aedeagus large, at centre curved. Needle-shaped cornutus slightly shorter than aedeagus, at centre slightly curved. Female genitalia: Lamella postvaginalis apically split and bilobed, dorsolaterally larger than that in N. notarioi, N. andalusiaria and N. obscuraria. Ductus bursae posteriorly sclerotized, longitudinally striated, towards corpus bursae membranous. The latter large, membranous. Signum stellate. Distribution and abundance: Central Mediterranean. Endemic to Sicily and southern Italy (Basilicata; Calabria). – Common in Sicily (Flamigni et al. 2007).

Phenology: Most probably bivoltine, from late May to late July and from mid-September to late October, with some changes in flight time depending on different climate and altitudes. Biology: Larva oligophagous on Rosaceae. Reared on Prunus spinosa, P. domestica and Crataegus sp. (D. Stadie pers. comm.), and Cotoneaster sp. (E. Friedrich pers. comm.). Recorded on Prunus (Rosaceae) (Bellier de la Chavignerie 1884), and ‘probably’ on Quercus ilex (Fagaceae) (Flamigni et al. 2007), the latter requiring confirmation. Adults attracted to light. 356

Habitat: According to Flamigni et al. (2007), it flies in xerothermic habitats from sea-level up to 1550 m. Similar species: Externally the species may be confused with other species of the obscuraria group (see above under ‘external characters’), but male genitalia allow reliable identification. Genetic data: BIN: BOLD:AAE5581 (n=7 from Italy: southernmost Calabria, Sicily). Genetically homogeneous. Nearest species: N. obscuraria (1.2%).

121. Nychiodes andalusiaria Millière, 1865 Nychiodes lividaria var. andalusiaria Millière, 1865: Iconogr. Descr. Chenilles Lépid. inédits II: 77, pl. 60, fig. 2. (Spain: Hoch-Castilien [High-Castilia]), Neotype ♂, Paralectotype ♀ (ZFMK, designated by Fazekas (1997), externally examined, male genitalia made by Fazekas: Nr. 4057 not traced in ZFMK). Nychiodes obscuraria estrellae Wehrli, 1934: Int. ent. Z. 27: 534, figs 15, 16 (Portugal: Sierra d’Estrella, Manteigas). Lectotype ♂, paralectotypes 8♂7♀ (ZFMK, Fazekas (1997) transferred this taxon from N. obscuraria to N. andalusiaria and designated a lectotype, examined). ­Redondo et al. (2009) and Corley (2015) regarded this population with smaller sized specimens as a synonym of andalusiaria, herewith we confirm it.

External characters and abdomen. Wingspan ♂ 33−45 mm, ♀ 35−47 mm. Wings light brown to brown, sometimes irrorated with creamy scales, costa and outer area of wings slightly darker, markings blackish. Forewing antemedial line deeply curved outwards near costa (whilst less outwards curved in all other Iberian Nychiodes); medial line present, often faint towards costa but always visible towards inner termen; postmedial line angled outwards on veins R1, M1 and M3, curved slightly inwards, ending at middle of inner margin. Forewing subterminal area partially covered with creamy scales as a linear shadow, replaced by brown scales towards costa; wavy line faint, hardly visible; termen dark brown. Hindwing basal and antemedial lines absent; medial line only towards inner termen partially visible as a shadow; postmedial line slightly angled on M2 vein (straight or slightly curved in N. obscuraria and N. notarioi, and outwards extended in N. hispanica). Hindwing subterminal area covered with creamy scales; wavy line nearly invisible. Discal spots blackish, present on all wings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings with yellow edging. Fringes light brown between veins and dark brown on vein endings. Underside of wings dull, light brown. Postmedial line present, blackish (whilst absent or hardly visible in the other species of the obscuraria group). Variation: Highly variable in size and colour (light brown irrorated with creamy to brown scales). Populations from Serra da Estrela slightly lighter and smaller in size. Male genitalia: Valva slightly narrower than in N. obscuraria. Costa of valva sclerotized only in its basal part, at centre strongly humped (clearly more than in N. obscuraria). Digitiform extension on the apex of valva present, covered with thin setae. Ampulla superior tubular, apically covered with setae; ampulla inferior and sacculus process acute, strongly sclerotized, not setose. Ampulla inferior shorter than sacculus process, the latter slightly curved towards saccus. In N. andalusiaria, distance between ampulla superior and inferior is less than distance between ampulla inferior and sacculus process, both structures are highly variable in size. Juxta anchor-shaped, apically concave, medial connection triangular. Relative width of apical versus basal part of juxta 1:1.5. Needle-shaped cornutus slightly shorter than aedeagus, sub-medially curved. 357

Female genitalia: Lamella postvaginalis with lateral arch-shaped extension, evenly sclerotized. Ductus bursae posteriorly sclerotized, towards corpus bursae membranous; slightly shorter than corpus bursae. The latter membranous, signum stellate, small, medially less sclerotized. Distribution and abundance: Endemic and restricted to the western Iberian peninsula (­Redondo et al. 2009). – Not well represented in most collections, apparently rare.

Phenology: Univoltine, from June to September. Biology: Larva oligophagous on Fabaceae. Recorded on Genista (Redondo et al. 2009), found also on Cytisus oromediterraneus (Corley 2015). Habitat: Inhabiting mountain habitats, from 900 up to 2000 m (Redondo et al. 2009). Similar species: The most similar species is N. notarioi, the latter differing by the yellowish postmedial line on the underside (blackish in N. andalusiaria). Structure of male genitalia allows reliable identification: ampulla superior tubular (as in N. notarioi, whilst thumb-shaped in N. hispanica, clubbed in N. obscuraria and N. ragusaria). Ampulla inferior shorter than sacculus process (similar to N. notarioi, but in N. hispanica ampulla inferior longer than sacculus process). Relative width of apical versus basal part of juxta 1:1.5 (as in N. obscuraria whilst 1:1 in N. ragusaria and N. notarioi 1:2 in N. hispanica). The only nearly reliable external character is the visibility of the postmedial line on the underside, which is weak or absent in the other species of this group (Fazekas 1997). Genetic data: BIN: BOLD:ABZ0919 (n=4 from Spain: Castilla y Leon, Galicia). Genetically homogeneous. Nearest species: N. notarioi (1.9%).

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Remarks: Scoble (1999) presents an erroneous author and original citation for this species: “Staudinger, 1892: Dt. ent. Z. Iris 5 (1): 173 (Spain: Andalusia)”. This error was repeated by different authors (e.g. Leraut 1999; 2009; Redondo 1999; Flamigni et al. 2007), as well as on the labels in many collections.

122. Nychiodes notarioi Expósito, 2005 Nychiodes notarioi Expósito, 2005: SHILAP Revta. lepid. 33 (132): 461–466 (Spain: Prov. T ­ eruel, Albarracín, Valdevecar, 1061 m). Holotype ♂, paratypes 12♂8♀ (coll. Expósito Hermosa, Móstoles-Madrid).

External characters and abdomen. Wingspan ♂ 33−48 mm, ♀ 38−52 mm. Wings dark beige to grey, markings blackish. Forewing antemedial line curved outwards near costa; medial line present, thin, sometimes part faint towards costa (completely absent in N. obscuraria); postmedial line angled outwards on M1 vein, curved slightly inwards, ending at middle of inner margin. Forewing subterminal area partially covered with creamy scales as a linear shadow, replaced by brown scales towards costa; wavy line faint, hardly visible; termen slightly darker. Basal, antemedial and medial lines absent from hindwing (medial line present in N. hispanica); postmedial line slightly curved (as in N. obscuraria, whilst angled in N. andalusiaria and N. ragusaria, outwards extended in N. hispanica). Hindwing subterminal area partially covered with creamy scales; wavy line faint or absent. Discal spots blackish, present on all wings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings with yellow shadow. Fringes brown, slightly darker on vein endings. Underside of wings basally dull, light beige, towards termen slightly darker. Postmedial line present, developed as a yellow shadow on all wings. Variation: The species shows a large variation in size, but also in colour (from lighter to darker wing pattern). Also male genitalia show some variation in the ornamentation of the valva, however the distance between ampulla superior, inferior and sacculus process revealed to be constant in all studied specimens. Male genitalia: Very similar to those of N. andalusiaria (e.g valva extended, costa of valva sclerotized up to the sub-apical part, medially humped, digitiform extension on apex of valva present, ampulla superior tubular), but in in N. notarioi the distance between ampulla superior and inferior is less than that between ampulla inferior and sacculus process (the opposite in N. andalusiaria). Juxta anchor-shaped, apically widely concave, width of its apical part of the same size as the basal part (1:1) (as in N. ragusaria, whilst 1:1.5 in N. anda­ lusiaria and N. obscuraria; 1:2 in N. hispanica). Needle-shaped cornutus slightly shorter than the aedeagus, slightly curved. Female genitalia: Lamella postvaginalis rectangular at the centre, with lateral arch-shaped extensions, evenly sclerotized. Ductus bursae posteriorly sclerotized, longitudinally striated, towards corpus bursae membranous. The latter membranous, with round signum, stellate; the sclerotization of signum reduced towards ductus bursae. Distribution and abundance: West Mediterranean. From eastern Spain to south-western France. – Not rare, sometimes common in its distribution area.

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Phenology: Apparently univoltine, from June to August (Taurand 2009), or even until mid-September (Redondo et al. 2009). Biology: Larva possibly oligophagous on Rosaceae and Fabaceae. In captivity feeding on Cotoneaster sp. (E. Friedrich pers. comm.), Prunus spinosa and Genista sp. (Taurand 2009). Habitat: Xerothermophilous. Found in dry steppes, wastelands and warm hillsides (Taurand 2009). From sea-level up to 1800 m. Similar species: The most similar species is N. andalusiaria. The latter differing by the blackish postmedial line on the underside (yellowish in N. notarioi). Structure of the male genitalia allows a reliable identification (see male genitalia section of N. notarioi and N. andalusiaria). Additionally the use of DNA barcodes is recommended for identification. Genetic data: BIN: BOLD:AAP3782 (n=13 from Spain: Castilla la-Mancha, Aragon, Catalonia). Genetically slightly heterogeneous, populations from Catalonia diverging by 0.8%. Nearest species: N. andalusiaria (1.9%).

123. Nychiodes hispanica Wehrli, 1929 Nychiodes obscuraria hispanica Wehrli, 1929: Mitt. münch. ent. Ges. 19: 51, pl. 4, fig. 9 (Spain: Andalusia, Sierra Nevada). Lectotype ♀, paralectotype 1♂, (ZFMK, designated by Fazekas (1997), examined). Nychiodes obscuraria atlanticaria Schwingenschuss, 1936: Mem. Soc. Sci. Nat. Maroc., 42: 86– 87. (Morocco, Gr. Atlas, Tachdirt, 2200–2700 m) Lectotype ♂, paralectotype 1♂, (NHMW, designated by Fazekas (1997)). Fazekas (1997) correctly realized the major differences between N. obscuraria and N. atlanticaria, but he mentioned a single diagnostic character in female genitalia “membrana intersegmentalis” and regarded N. atlanticaria as a bona species. “Membrana intersegmentalis” is a membranous structure connecting the sterigma with

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sternum A8. This structure easily gets macerated when exposed to potassium hydroxide. Additionally, Fazekas (1997) had erroneously depicted female genitalia (preparation Nr. 2594) as N. atlanticaria stating in the same paper, that this slide belongs to the paralectotype of N. mauretanica. Genitalia structure of several male and female specimens of N. atlanticaria (including the specimens studied by Fazekas (1997), deposited in ZFMK), suggest N. atlanticaria should be treated as a junior synonym of N. hispanica. DNA barcoding data, however, seem to justify subspecies rank for atlanticaria (see below). Nychiodes obscuraria torrevinagrensis Expósito, 1984: Revta Lepid. 12 (3): 203 (Spain: Cazorla, Casa Forestal de Torre del Vinagre). Holotype ♂ (coll. Expósito Hermosa, Móstoles-Madrid). In his original species description, Expósito Hermosa (1984) correctly realized high similarity of N.  hispanica and torrevinagrensis, therefore he regarded the latter taxon as a ‘sister subspecies’ of N. hispanica under N. obscuraria. Later, Expósito Hermosa (1989) followed Dufay (1974) and regarded N. hispanica as a bona species as well as ‘N. torrevinagrensis’, but mentioned that these two taxa cannot be easily diagnosed based on external characters. Also Fazekas (1997) mentioned high similarity of these two taxa, and downgraded torrevinagrensis to a subspecies of N. hispanica. He could not find any single diagnostic character in the female genitalia of torrevinagrensis and hispanica and hypothesized torrevinagrensis to be a transitional form between N.  hispanica and N.  atlanticaria. Whilst Expósito Hermosa (1984, 1985a) described and recorded torrevinagrensis from Torre del Vinagre and Jaén, Fazekas (1997) recorded it also from Aragon and Sierra Nevada. The latter is the type locality of N. hispanica. Moreover Expósito Hermosa (1984) and Fazekas (1997) recorded N. hispanica from Alava, Barcelona, Burgos, Cuenca, Gerona, Granada, Guadalajara, Guipúzcoa, Huesca, La Rioja, Llérida, Madrid, Navarra, Pamplona, Segovia, Teruel, and from France: East Pyrenees, Hérault, Dordogne, Hautes-Pyrenées, Lozère). All these published data, along with our examined materials from different localities in western Europe, do not confirm any clear geographical separation between these taxa (this issue had been left open by Redondo et al. 2009) suggesting treatment of torrevinagrensis as a junior synonym of N. hispanica. However, DNA barcoding data of a large series of specimens including five ‘torrevinagrensis’ from localities close to type locality suggest reduced gene-flow between torrevinagrensis and N. hispanica which may justify subspecies rank for this taxon (see below).

External characters and abdomen. Wingspan ♂ 38−48 mm, ♀ 44−50 mm. Wings brown to grey, sometimes irrorated with yellowish brown, costa and terminal area of wings slightly darker, markings dark grey to black. Very few specimens with darker ground colour irrorated with ochre or creamy yellow ground colour irrorated with dark brown. Forewing antemedial line curved outwards near costa; medial line present, close to costa often faint but towards inner termen present (absent in N. obscuraria); postmedial line angled outwards on M1 vein, curved slightly inwards, ending at middle of inner margin. Forewing subterminal area partially covered with creamy scales as a linear shadow, replaced by brown scales towards costa; wavy line faint, hardly visible; margin dark brown. Basal and antemedial lines absent from hindwing; medial line dull, mostly faint; postmedial line angled at M2 vein and often extended outwards (straight or slightly curved in N. obscuraria and N. notarioi, slightly angled in N. andalusiaria and N. ragusaria). Hindwing subterminal area covered with creamy scales; wavy line hardly visible. Discal spots blackish, present on all wings. Terminal line of all wings blackish, narrow, discontinuous, more strongly marked between vein endings with yellow edging. Fringes brown. Underside of wings dull, beige to dark brown, terminal area and at costa darker. Postmedial line partially present, blackish. Variation: Extremely high variation in wing size and pattern, as well as in the size of ampulla inferior and sacculus process of male genitalia. In addition, DNA-barcoding pattern is highly heterogeneous.

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Male genitalia: Costa of valva sclerotized up to the subapical part, medially humped. Digitiform extension on apex of valva present, covered with thin setae. Ampulla superior thumb shaped. Ampulla inferior and sacculus process acute, strongly sclerotized, not setose; the former clearly larger than the equivalents in N. andalusiaria and N. notarioi (important diagnostic character); sacculus process curved towards saccus (both these characters are highly variable in size, see figures). Juxta anchor-shaped, apically rectangular, relative width of apical versus basal part 1:2. Aedeagus large. Needle-shaped cornutus slightly shorter than aedeagus, at centre slightly curved. Female genitalia: Apophyses anteriores clearly longer than in the other species of the obscuraria group (3/4 of apophyses posteriores). Lamella postvaginalis heart-shaped at centre, laterally with arch-shaped extensions, strongly sclerotized (slightly variable in size). Length of ductus bursae 3/4 of corpus bursae, posteriorly sclerotized, longitudinally striated, towards corpus bursae membranous. The latter membranous, with large, stellate signum (comparatively larger than in the other species of the obscuraria group). Distribution and abundance: West Mediterranean. In Europe restricted to southern Spain. In Cazorla region replaced by subsp. torrevinagrensis, in southernmost Andalusia by subsp. atlanticaria. – Outside Europe from Morocco to northern Algeria (subsp. atlanticaria).

Phenology: Univoltine, adults from July to late September. Biology: Larva oligophagous on Rosaceae. Reared on Prunus spinosa, P. domestica and Crataegus sp. (D. Stadie pers. comm.), and on Cotoneaster sp. (E. Friedrich pers. comm.). Habitat: Usually from 700 up to 2100 m (Redondo et al. 2009), sometimes also in the lowlands, a population recorded at 50 m near Cadiz (2 km south of Almoraima, leg. P. Skou).

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Similar species: Easily confused with other species of the obscuraria group. Study of the structure of male and female genitalia and/or DNA barcodes necessary for reliable identification (see male genitalia section of N. notarioi and N. andalusiaria). In contrast to all other species of this group, female genitalia of N. hispanica show a large heart-shaped lamella postvaginalis. Genetic data: BIN: BOLD: AAI3448 (n=19 from Spain and Morocco). Genetically heterogeneous, in several haplotype-clusters, maximum variation 2.2%. Five DNA barcodes from localities close to the type locality of torrevinagrensis diverging from those of eastern Andalusian localities (Sierra Nevada, type locality of N. hispanica) by 1.4% supporting reduced gene-flow between the populations of these two taxa despite close geographical proximity. The same applies for the DNA barcodes of the Sierra Nevada populations (’hispanica’) versus the populations from Morocco (type locality of ‘atlanticaria’) diverging by 1.0−1.5%, supporting validation of the taxon atlanticaria at subspecies rank. Populations from southernmost Andalusia (southern prov. Malaga) clustering together with Moroccan populations. Nearest species: N. notarioi 4.9%.

124. Nychiodes waltheri Wagner, 1919 Nychiodes obscuraria waltheri Wagner, 1919: Dt. ent. Z. Iris 33: 110 (Turkey: Istanbul). Lecto­ type ♂ (Haidar-Pascha), Paralectotypes 1♂2♀ (NHMUK, designated by Fazekas (1997)). Nychiodes waltheri transcaspia Wehrli, 1929: Int. ent. Z. 22 (42): 386 (Turkmenistan: Ashkha­ bad). Syntype(s) ♂ (ZFMK, examined). Valid at subspecific rank (Scoble 1999). Nychiodes waltheri osthelderi Wehrli, 1929: Mitt. münch. ent. Ges. 19 (2–4): 42, pl. 3, fig. 6 (Turkey: Toros Dağları, Marasch). Holotype ♂ (ZSM, examined). Colour patterns fall into the variation range of the nominotypical subspecies. Here regarded as a synonym of the nominotypical subspecies, based on sympatric occurrence of such forms. Nychiodes waltheri syriaca Wehrli, 1929: Mitt. münch. ent. Ges. 19 (2–4): 41 (Turkish/Syrian border: Akbès). Holotype ♀ (ZFMK, examined). Colour patterns fall into the variation range of the nominotypical subspecies. Here regarded as a synonym of the nominotypical subspecies, based on sympatric occurrence of such forms. Nychiodes waltheri saerdabica Wehrli, 1938: Ent. Rdsch. 55 (31): 355 (Iran: Tacht i Suleiman; Särdap-Tal). Syntypes 2♂1♀ (ZFMK, examined). Valid at subspecific rank (Scoble 1999). Confirmation of this subspecies needs further study based on larger series. Nychiodes cuencaensis Leraut, 2009: Moths of Europe 146 (‘Spain: Castile’, mislabelled), Holotype ♀ (in MNHN, examined). Female genitalia of this monotypic species are identical with those of N. waltheri (as illustrated in this book). As N. waltheri is not distributed in western Europe, the single female type specimen is likely mislabelled. Herewith, we regard this name as a junior synonym of N. waltheri.

External characters and abdomen. Wingspan ♂ 35−46 mm, ♀ 41−48 mm. Ground colour of wings light to dark brown; basal and terminal areas of forewing irrorated with redbrown scales, medial area light brown. Forewing antemedial line curved outwards twice; medial line very thin, sometimes discontinuous, faint towards costa; postmedial line angled outwards on M1 vein, at centre curved slightly inwards, minutely dentate on veins, ending at middle of inner margin. Forewing subterminal area with thin light brown shadow, replaced by yellowish scales towards costa; wavy line present, discontinuous, yellowish; termen slightly darker. Hindwing basal and medial areas light brown, terminal area irrorated with red-brown scales; basal and antemedial lines absent; medial line dull, hardly visible; postmedial line sharp, blackish, characteristically angled outwards between M1 and M2 veins (reminiscent of equivalent in N. hispanica, whilst curved or slightly angled in all other 363

Text-figs 159–160. Diagnostic characters of N. dalmatina and N. waltheri. Text-fig. 159: N. dalmatina Wagner, 1909, Bulgaria: Pirin Mts. Liljanovo, 14.8.1981. Text-fig. 160: N. waltheri Wagner, 1919, Bulgaria: Sozopol, 14.11.1987.

European Nychiodes species); wavy line hardly visible. Discal spot blackish, conspicuous on all wings. Terminal line of all wings blackish, narrow, discontinuous, with tiny yellow edging. Fringes brown, slightly darker at vein endings. Underside of wings brown, terminal area mixed with orange-brown and appearing slightly darker, basal area light brown; postmedial line partially visible as a yellow shadow. Variation: The species is highly variable in size and wing colour, from light brown to dark brown and grey. Validity of the subspecies of N. waltheri needs to be examined based on more material from the type localities of all described subspecies which is out of scope of present study. Male genitalia: Valva wide at base, round, widely extended (valva narrow at base in N. aphro­ dite Hausmann & Wimmer, 1994 from Cyprus). Costa of valva sclerotized up to the subapical part, humped at centre (not humped, but slightly curved in N. aphrodite). Ampulla superior tubular, ampulla inferior digitiform, thin, shorter than ampulla superior; both apically covered with few tiny spines (ampulla inferior not spinose in the obscuraria group; ampulla inferior in N. aphrodite as thick as ampulla superior and apically with more spines); sacculus process apically acute, strongly sclerotized, arched towards uncus (much shorter in N. aphro­ dite and arched towards saccus). Juxta anchor-shaped at base, apically cup-shaped, width of apical part corresponding to that of basal part. Aedeagus thick, needle-­shaped cornutus half length of aedeagus (in all species of the obscuraria group cornutus slightly shorter than the aedeagus). Female genitalia: Very small genitalia (larger in all other Nychiodes species). Apophyses anteriores very short (1/3 of apophyses posteriores). Lamella postvaginalis well sclerotized, conically extended. Ductus bursae very short, with extended sclerotized patch. Corpus bursae membranous. Signum stellate, very large (compared with the size of coupus bursae), well sclerotized, variable in size from round to slightly elongated.

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Distribution: East Mediterranean: In Europe distributed in the Balkan peninsula (Bulgaria, eastern Greece, European Turkey). – Outside Europe from Turkey, Syria, northern Iran (subsp. saerdabica) to southern Turkmenistan (subsp. transcaspia).

Phenology: Bi- or trivoltine, depending on the locality, from late May to mid-November (Fazekas 1996). Biology: Larva oligophagous on Rosaceae. Reared on Prunus spinosa, P. domestica and Cotoneaster sp. (D. Stadie, E.  Friedrich pers. comm.). Feeding also on leaves of Prunus mahaleb und P. tenella (Fazekas 1996). Habitat: According to Fazekas (1996) the distribution area of this species can be attributed to the following three plant associations: 1. Mediterranen evergreen sclerophyllous forests (Oleo-Ceratonion zone); 2. Submediterranen deciduous mixed forests (Ostrio-Carpinion aegocum subzone); 3. Continental mixed forests and forest steppe (Quercetum frainetto-­ cerris subzone; Quercetum petraeae subzone). It occurs from sea-level up to 2000 m. Similar species: Differing from western Mediterranian Nychiodes species by the irroration with orange-brown scales, moreover, its distribution area being well separated. For a reliable identification, genitalia dissection is recommended. Both male and female specimens have unique genitalia and cannot be confused with any other European species. Male genitalia show clear differences from N. aphrodite Hausmann & Wimmer, 1994 from Cyprus: valva wide at base in N. waltheri (narrow at base in N. aphrodite). Costa of valva humped at centre (not humped, but slightly curved in N. aphrodite). Ampulla inferior digitiform, thin and shorter than ampulla superior, both apically covered with few tiny setae (ampulla inferior as thick as ampulla superior and apically with more setae in N. aphrodite); sacculus process apically acute, arched towards uncus (much shorter in N. aphrodite and arched towards saccus). 365

Genetic data: BIN: BOLD:ACF3394 (n=9 from Greece, Bulgaria, western Turkey). Genetically heterogeneous, in eastern Turkey replaced by populations with separate BIN (n=5; BOLD:AAI3454) at a minimum pairwise distance of 1.6%, subsp. saerdabica from northern Iran diverging by 2.0% from all other populations (n=2; BOLD:ABZ5798). Nearest species: N. dalmatina (5.7%), N. amygdalaria (6.3%), N. aphrodite (6.9%).

125. Nychiodes amygdalaria (Herrich-Schäffer, 1848) Boarmia amygdalaria Herrich-Schäffer, 1848: Syst. Bearb. Schmett. Eur. 3 (31: 82; ibidem, pl. 70, figs 432, 433 (non binominal) (Crete). Syntype(s). Nychiodes amygdalaria almensis Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 441, pl. 38: c (Syria; Lebanon, 1400 m; Palestine). Syntype(s) 1♂1♀ (ZFMK, examined). Wing pattern and genitalia structure of the studied type specimens confirmed this taxon to be a colour variation of amygdalaria. We therefore downgrade it from subspecies rank (Scoble 1999) to a junior synonym. Nychiodes amygdalaria malatyaca Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 441, pl. 38: d (Turkey: Malatya-Tecde). Holotype ♂, paratype ♂ (ZFMK, examined). Wing pattern and genitalia structure of the studied type specimens confirmed this taxon to be a colour variation of amygdalaria. We therefore downgrade it from subspecies rank (Scoble 1999) to a junior synonym.

External characters and abdomen. Wingspan ♂ 35−46 mm, ♀ 41−47 mm. Ground colour of wings characteristically light to dark ivory, irrorated with grey-brown; basal and terminal area of wings lighter, markings dark grey to blackish. Forewing antemedial line angled outwards near costa; medial line present, but weak, sometimes discontinuous; postmedial line curved inwards near costa, deeply angled outwards on M1 vein, at centre slightly curved inwards, ending at middle of inner margin. Forewing subterminal area with light ivory scales, replaced by darker scales near costa; wavy line absent or hardly visible; termen ivory mixed with grey scales. Basal and antemedial lines absent from hindwing; medial line dull, dark brown; postmedial line sharp, dark brown, distally edged with light brown, weakly angled on M3 vein. Hindwing subterminal area ivory; wavy line hardly visible. Discal spots dark brown, faint on upperside of hindwing, but clearly visible on the underside. Terminal line of all wings blackish, narrow, discontinuous on vein endings. Fringes ivory, slightly darker at vein endings. Underside of wings pale ivory, termen slightly darker, postmedial line partially visible as a light brown shadow. Variation: The species is variable in size, but also in colour (from grey to light cream specimens). Male genitalia: Valva generally smaller and narrower than in N. dalmatina. Costa of valva less extended; costa of valva not humped, sclerotized up to the apex, at apex covered with spines (in N. dalmatina costa of valva humped at centre, subapical spines arranged on a ridge). Ampulla superior and inferior tubular, apically covered with tiny spines, the latter shorter and much narrower than the former; sacculus process absent (in contrast to all species of the obscuraria and waltheri groups). Juxta basally anchor-shaped, connected with the cup-shaped apical part by a medial rod-shaped sclerotized part. Aedeagus narrow and long (compared with the species of the obscuraria and waltheri groups), needle-shaped cornutus less than half length of aedeagus. Female genitalia: Ovipositor ovally elongated. Lamella postvaginalis ovally extended anteriorly and posteriorly (conical in N. dalmatina). Ductus bursae membranous, roughly of same 366

size as corpus bursae, longitudinally striated. Corpus bursae membranous, signum stellate, medium-sized (compared with corpus bursae) more or less sclerotized. Distribution and abundance: East Mediterranean. Distributed in the Balkan peninsula, Bulgaria, Macedonia and Greece including Crete. – Extremely rare, only single specimens found at light. – Outside Europe across central and eastern Turkey to Georgia, Armenia to Iran. In the south to the Levant (Israel).

Phenology: Apparently uni- or bivoltine (depending on temperature), from mid-May to late September. Biology: Larva oligophagous on Rosaceae. Reared on Prunus spinosa, P. domestica and Crataegus sp. (D. Stadie pers. comm.). Habitat: In open woodland. From sea-level up to 1700 m. Similar species: This species cannot be confused with any other European Nychiodes species and is easily diagnosable based on its wing pattern. Genetic data: BIN: BOLD:AAE5586 (n=9 from Turkey, Israel). Genetically homogeneous. Nearest species: N. divergaria Staudinger, 1892 from Turkey (5.9%), N. waltheri (6.3%).

126. Nychiodes dalmatina Wagner, 1909 Nychiodes lividaria dalmatina Wagner, 1909: Ent. Z., Frankf a. M. 23 (4): 18 ([Croatia], Dalmatia: near Zaras). Lectotype ♂ (NHMW, designated by Fazekas 1997) Nychiodes dalmatina andreasaria Warnecke, 1925: Int. ent. Z. 19 (4): 20 (Macedonia: Monastir). Holotype ♀. Junior synonym of N. dalmatina (Fazekas 1997).

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Nychiodes dalmatina vorbrodtaria Wehrli, 1929: Mitt. münch. ent. Ges. 19 (2–4): 38 (Switzer­ land: La Batiaz, Martigny). Syntypes. Junior synonym of N. dalmatina (Fazekas 1997). Presence of N. dalmatina in Switzerland was rejected since the taxon was based on old ‘ex. ovo.-reared’ specimens labelled as subspecies “vorbrodtaria” by Wehrli (1929b) originally received from Dalmatia by the collector Wullschlegel, but labelled with the Swiss locality of rearing “La Batiaz” (Fazekas 1997). Confirmed by the SwissLep Team (2010), as no more new material of this taxon was collected in Switzerland.

External characters and abdomen. Wingspan ♂ 33−47 mm, ♀ 37−48 mm. Ground colour of wings light to dark brown, irrorated with yellow scales. Forewing antemedial line curved outwards twice; medial line very thin, sometimes discontinuous, faint towards costa; postmedial line deeply angled outwards on M1 vein (important diagnostic character), curved slightly inwards at centre, minutely dentate on veins, ending at middle of inner margin. Forewing subterminal area with thin light brown shadow, replaced by yellowish scales near costa, interrupted by the projection of postmedial line; wavy line present, yellowish; termen dark grey, except yellowish apex (the latter character present only in N. dalmatina and N. amygdalaria). Basal and antemedial lines absent from hindwing; medial line dull, weak, dark brown; postmedial line sharp, blackish, characteristically angled outwards on M1 vein (slightly angled also in N. amygdalaria, N. hispanica, and curved in all other European congeners). Hindwing medial and subterminal areas irrorated with creamy scales; wavy line hardly visible. Discal spot blackish, conspicuous on all wings. Terminal line of all wings blackish, narrow, discontinuous. Fringes brown, slightly darker at vein endings. Underside of wings light brown to beige, subterminal area slightly darker, terminal concolorous with basal and medial area; postmedial line partially visible as a yellow shadow. Variation: Wing colour shows high variation. However, the pattern of markings revealed to be constant in all studied specimens. Male genitalia: Generally larger than those of N. amygdalaria. Costa of valva strongly sclerotized up to the apex, strongly humped at centre (not humped in N. amygdalaria); apical tiny spines arranged on a ridge (distributed all over the apex in N. amygdalaria). Ampulla superior thicker, but of same length as ampulla inferior, both tubularly elongated, apically covered with stout spines; sacculus process absent. Tiny warts sometimes present ventrally on the valva (variable in size and often absent). Juxta basally anchor-shaped, connected with the bowl-shape apical part by a medial rod-shaped sclerotized part. Aedeagus narrow and long, needle-shaped cornutus nearly half length of aedeagus. Female genitalia: Ovipositor ovally elongated, clearly thinner than in N. amygdalaria. ­Lamella postvaginalis sclerotized, conical. Ductus bursae extremely long (longest in the genus), sclerotized nearly all over, but with reduced sclerotization towards corpus bursae. Corpus bursae membranous, signum stellate, medium-sized (compared with corpus bursae), less sclerotized. Distribution and abundance: (South-east) European. North-easternmost Italy (near Trieste), Slovenia, Croatia, Bosnia, Serbia, Montenegro, Kosovo, Albania, Macedonia, south-western Romania, Bulgaria, Greece. It is expected to be found also in European Turkey. – Outside Europe, e.g. from western Turkey, there are no reliable records (Fazekas 1996; Rákosy & Bere 1996). Phenology: Uni- or bivoltine (depending on temperature), from mid-May to late September (overlapping generations?) (P. Skou own data).

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Biology: Larva oligophagous on Rosaceae. Found on Cotoneaster racemiflorus and Crataegus sp. (M. Leipnitz pers. comm.). In captivity reared on various Rosaceae: Prunus spinosa, P. domestica, Cotoneaster sp., Amelanchier sp., Crataegus sp., Pyracantha sp. (M. Leipnitz, E. Friedrich, D. Stadie pers. comm.). Habitat: In a variety of open habitats, scrubland and places with scattered shrubs of the food-plants. From sea-level up to 1700 m (P. Skou own data). Similar species: Differing from other species of obscuraria group (most being allopatric) by the characteristic shape of the postmedial line of both wings (see Text-figs 154–160). In doubtful cases, genitalia dissection is recommended. Genetic data: BIN: BOLD:AAI3512 (n=9 from Croatia, Greece, Bulgaria). Genetically heterogeneous, maximum variation 1.8%. Nearest species: The three Levantine species N. muelleri Hausmann, 1991, N. persuavis Wehrli, 1929 and N. palaestinensis Wagner, 1919 at distances of 5.1–5.5%.

Phaselia Guenée, 1858 Phaselia Guenée, 1858, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 228. Type species: Synopsia phaeoleucaria Lederer, 1855. There are no genus synonyms.

Diversity and distribution: Eight species in the southern central and south-western Palae­ arctic region including Arabian peninsula. External characters and abdomen: Large, robust species. Wingspan 25–35 mm. Females tend to be larger than males. Proboscis vestigial. Ground colour whitish, more or less mottled with dark scales. Forewing with sigmoid postmedial line (usually broken on vein A) and zigzagging antemedial line. Terminal area with varying umber of teeth. Hindwing distinctly 369

lighter. Antennae of males and females bipectinate. Hindtibia of both sexes with 2+2 spurs. Species can often be separated only by genitalia dissection. Male genitalia: Uncus beak-shaped, pointed, gnathos with non-fused arms. Valva short, wide, costal arm narrow, rounded, apex setose. Ventral arm shows differences between species: rectangular base structure, without or with spines. If bearing spines, with varying number, length and position. Saccus wide, rounded. Juxta a plate of varying size. Aedeagus short, thick, apex tapered. Vesica usually with two cornuti: one bent thorn, base thick, and a narrow setose part (absent in P. joestleinae Hausmann, 1996). Female genitalia: Papillae anales sclerotized, soft, slightly setose. Apophyses posteriores of medium length. Apophyses anteriores distinctly shorter. Lamella antevaginalis sclerotized, in transverse position. Lamella postvaginalis a plate. Ostium bursae extended by a tranverse sclerite. Ductus bursae funnel-shaped, anteriorly narrowing. Corpus bursae membranous, posterior end slightly striated, anterior end expanded. Signum circular, weakly stellate. Biology: Larvae of many, perhaps all, species are monophagous (2) on Ephedra. Remarks: In multi-gene analyses Phaselia and non-European Melanolophia were sister taxa (Sihvonen et al. 2011), whereas in more extensive study (Murillo-Ramos et al. 2019) Phaselia was recovered as a sister to a large lineage, which contains mostly non-European genera. The few included European genera in the large lineage are Nychiodes, Eurranthis and Lycia.

127. Phaselia serrularia (Eversmann, 1847) Fidonia serrularia Eversmann, 1847: Bull. Soc. imp. Nat. Moscou 20 (2): 81, pl. 6, fig. 6 (Kazakhstan [East]: Altai Mts., Noor-Saisan [Saisan Lake]). Syntype(s) ♂. Synopsia phaeoleucaria Lederer, 1855: Verh. zool.-bot. Ges. Wien 5 (Abh.): 114, pl. 2, fig. 3 (Russia: Siberia, Altai Mts.). Holotype ♂. Phaselia serrularia catharia Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 467, pl. 41: c (Armenia: Darasham). Syntypes 2♂. Valid at subspecific rank (Scoble 1999). Phaselia serrularia pithana Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 467 (NE Iran: Kuh i Mirabi, 2200 m). Syntype(s). Valid at subspecific rank (Scoble 1999). Phaselia serrularia shurensis Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 466, pl. 41: c (Dagestan: Temir-Shan-Shura district, Kaptshugaj). Holotype ♂. Valid at subspecific rank (Scoble 1999).

External characters and abdomen: Wingspan ♂ 28–33 mm, ♀ on average slightly larger. Forewing: Costa slightly concave, ground colour whitish, more or less mottled with brown. Postmedial line black, s-shaped, with small black triangular teeth on veins, distally a fine brownish line with row of distinct triangular teeth on whitish ground. Between this and terminal line a brown band. Medial shadow near postmedial line. Discal spot on medial shadow. Antemedial line zigzagging, joining costa at right-angle or obtusely. Basal area brown, paler near wing base. Hindwing: White with postmedial line angled on M2. Medial shadow of forewing continuing on hindwing, half way joining postmedial line. Discal spot small. Terminal area sometimes speckled with fine dark scales. Both wings with black terminal lines. Fringes chequered whitish and brownish with line parallel to termen. Underside pattern as on forewing, less contrasted, lines paler. Antemedial line indistinct or absent. Frons, vertex and thorax with a mixture of black and white scales. Patagia and mesothorax often dark brown, abdomen paler brown. Antennae of males and females bipectinate. Hindtibia of both sexes with 2+2 spurs.

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Variation: Little variation in degree of dark mottling and extent of brown shadows. Male genitalia: Uncus beak-shaped, weakly setose, sclerotized. Gnathos consists of two small, non-fused sclerotized upturned bands. Valva short, wide, strongly sclerotized. Costal arm narrow, apex rounded, setose. Ventral arm strongly sclerotized, almost rectangular, covered with large number of spines, distally getting longer. Saccus wide, rounded. Juxta a sclerotized band, base slightly concave, posterior margin serrate. Aedeagus short, thick, anterior end membranous, distinctly longer than in P. algiricaria, apex tapered. Vesica with two sclerotized cornuti as in P. algiricaria, but spinose part shorter, not s-shaped. Female genitalia: Larger than those of P. algiricaria. Papillae anales sclerotized, slightly setose. Apophyses posteriores of medium length, apophyses anteriores shorter, less than half length of the former. Lamella antevaginalis large, sclerotized, laterally with two round sclerites. Lamella postvaginalis posteriorly with rounded posterior margin, anteriorly serrate. Ostium bursae wide. Antrum strongly sclerotized, broader than in P. algiricaria. Ductus bursae anteriorly slightly narrowing. Corpus bursae membranous, anterior end expanded. Signum distinct, circular, stellate. Distribution and abundance: Turanian. In Europe only in southern Urals and southernmost European Russia. Erroneously recorded for Belarus and Ukraine in Leraut (2009). – Outside Europe in Kazakhstan, subsp. shurensis in Dagestan, Georgia, and Azerbaijan. Subsp. catharia described from Armenia, with Turkish populations tentatively attributed. Subsp. pithana from northern Iran to Turkmenistan. For unclear taxonomy see under genetic data.

Phenology: Probably bivoltine. In southern Urals from late May to early June and from mid-July to mid-August. Adults active at night, readily attracted to light, usually late in the night (J. Gelbrecht pers. comm.). 371

Biology: Apparently unknown. Habitat: Xerothermophilous. A steppe species, preferring xerothermic slopes (K. Nupponen pers. comm.). In southern Urals up to 350 m (K. Nupponen pers. comm.). In western Kazakhstan found at 200 m and in Turkey from 700 up to 2400 m (J. Gelbrecht pers. comm.). In Armenia from 700 up to 1300 m. Similar species: European P. algiricaria with only three teeth in the terminal area near tornus and with different shape of the antemedial line (joining forewing costa at acute angle). Genetic data: BIN: BOLD:AAY2405 (n=2 from north-eastern Turkey, Russia: southern Urals). There are three or four other BINs at minimum distances of 5.1% (Mongolia, Kazakhstan, Kirghizstan, but also Iran, Armenia, south-eastern Turkey) which could be the ‘true’ P. serrularia (described from Altai region). Potentially, these are not conspecific with the European and north-eastern Turkish populations which may still be undescribed. Remarks: Requiring comprehensive revision and further study (see under genetic data).

128. Phaselia algiricaria Oberthür, 1913 Phaselia deliciosaria algiricaria Oberthür, 1913: Études Lépid. comp. 7: 260, pl. 166, fig. 1619 (Algeria: Géryville). Syntype(s) including ♂. Synopsia relicta Turati, 1930: Atti Soc. ital. Sci. nat. 69: 64, pl. 2, figured (Libya: Cyrenaica, Scleidima). Syntype(s), including ♂ (ZFMK). Valid at subspecific rank (Scoble 1999). Phaselia deliciosaria gigantaria Schwingenschuss, 1936: in Zerny, Mém. Soc. Sci. nat. Phys. Maroc 42: 87 (Morocco: Ijjoukak). Syntypes 6♂2♀. Valid at subspecific rank (Scoble 1999). Phaselia algiricaria aragona Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 468, pl. 41: d (Spain: Prov. Teruel, Albarracín). Syntype(s) ♂ (ZFMK). Valid at subspecific rank (Scoble 1999).

External characters and abdomen (subsp. aragona): Wingspan 25–35 mm. Forewing: Costa slightly concave. Wavy line whitish, curved, undulating between vein CuA1 and inner margin. Postmedial line black, s-shaped, with brownish subterminal line nearby. Antemedial line forms an acute angle at costa, straight between costa and vein CuA2, then zigzagging to inner margin. Basal and terminal areas brownish, medial area whitish with beige or brown parts. Discal spot linear. Hindwing white with more or less distinct, curved postmedial line and sometimes a small discal spot. Terminal area often speckled with fine dark scales. On both wings black terminal lines. Fringes chequered whitish and beige, with dark line parallel to termen. Underside: Forewing with postmedial and weak medial line. Both lines black towards costa, the latter silvery grey apically. Antemedial line indistinct or absent. Hindwing white, with black blotch at costa. Discal spots weak. Frons, vertex and thorax with a mixture of black and white scales. Patagia and mesothorax often dark brown, abdomen paler brown. Antennae of both sexes bipectinate. Hindtibia of both sexes with 2+2 spurs. Variation: Little, in degree of dark mottling and distinctness of pattern. Male genitalia: Uncus and gnathos as in P. serrularia. Valva different, short, wide, strongly sclerotized. Apex of costa rounded, with group of setae. Sacculus setose, trapezoidally indented below apex. Saccus wide, rounded. Juxta sclerotized, almost circular, slightly concave at base. Aedeagus short, thick, anterior end membranous, apex tapered. Vesica with two sclerotized cornuti: one a sclerotized, bent thorn, the other narrow, s-shaped, spinose.

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Female genitalia: Significantly smaller than those of P. serrularia. Papillae anales sclerotized, slightly setose. Apophyses posteriores of medium length, apophyses anteriores shorter, one third length of the former. Lamella antevaginalis weakly sclerotized, in transverse position, at both sides with an unevenly rounded structure. Lamella postvaginalis a weakly sclerotized plate, anteriorly curved. Ostium bursae wide. Antrum a strongly sclerotized band, smaller than in P. serrularia, pointed on both sides. Ductus bursae with two sclerotized blotches, anteriorly slightly narrowing. Corpus bursae membranous, slightly striated, anterior end expanded. Signum distinct, circular, stellate. Distribution and abundance: West Mediterranean. In Europe (approximately) restricted to the eastern half of Spain (subsp. aragona). – In Spain local, usually found in limited numbers. – Outside Europe in Morocco (subsp. gigantaria), Algeria (nominotypical subspecies), Tunisia and Libya (subsp. relicta).

Phenology: Bivoltine. From early April to mid-July and from early August to mid-September. Possibly with overlapping generations in some localities. Caterpillar from May to mid-September (Gómez de Aizpúrua 2002). Hibernation as pupa. Adults active at night, both sexes readily attracted to light. Biology: Larva monophagous on Ephedra. Found on Ephedra major (M. Leipnitz pers. comm.) and E. fragilis (G. King pers. comm.). Recorded on Ephedra nebrodensis and E. distachya (Spain: Gómez de Aizpúrua et al. 2002). Habitat: Xerothermophilous. In a variety of open habitats with scattered trees and bushes, including semideserts. From sea-level up to 1400 m. Similar species: P. serrularia differs by a row of teeth over the whole terminal area and by the different shape of antemedial line (joining forewing costa right-angled). 373

Genetic data: BIN: BOLD:AAL2739 (n=3 from Spain), genetically homogeneous in Europe (subsp. aragona) but in Morocco (High Atlas, subsp. gigantaria) strongly diverging, with separate BIN (BOLD:ACM0587) at a distance of 7.3% casting doubt on the current taxonomic status of all intraspecific taxa and requiring further research. Nearest congener: P. delicio­ saria (9.3%), suggesting a long isolated evolutionary history for P. algiricaria.

Ekboarmia Wehrli, 1943 Ekboarmia Wehrli, 1943, Gross-Schmett. Erde 4 (Suppl.): 517. Type species: Boarmia atlanticaria Staudinger, 1859. [Replacement name for preoccupied Rhoptria Gumppenberg, 1892. Ekboarmia was originally proposed as a subgenus of Boarmia Treitschke, 1825]. ‡ Rhoptria Gumppenberg, 1892, Nova Acta Acad. Caesar. Leop. Carol. 58: 299. Type species: Boarmia atlanticaria Staudinger, 1859. [Junior homonym of Rhoptria Guenée, 1858 (Geo­ metridae: Ennominae)].

Diversity and distribution: Four species in the West Mediterranean area, including the French Alps, Iberian peninsula, the Balearic Islands, Sardinia in Italy, and Morocco, Algeria and Tuni­sia in North Africa (Skou et al. 2017a). External characters and abdomen: Medium-sized, wingspan 18–29 mm. Wings grey-brown or different shades of brown, markings blackish. Postmedial line prominent, on forewing rather smooth, dentate or even zigzagging, turning inwards near costa, outer margin bordered with whitish. Forewing discal spot small, dark, distinct. Hindwing postmedial line blackish. Male antennae bipectinate, female antennnae filiform. Male hindtibia with hair pencil, hindtibia of both sexes with 2+2 spurs. Male sternite 1+2 with free lateral process, 3rd sternite with setal comb and 8th tergite weakly triangular, posterior margin narrower. Other abdominal segments in both sexes without modifications. Male genitalia: Uncus triangular, apex round. Socii absent. Gnathos arms fused ­ventrally, medial element upturned, plate-shaped. Valva evenly narrowing towards apex, costa sclerotized, apical part of valva with narrow, setose ridge. Juxta sclerotized, with two large arms, inner margins serrate or dentate. Posterior arms of juxta absent. Saccus elongated, upturned medially. Aedeagus cleft apically, tip bents when vesica is everted. Vesica slightly enlarged at base, without cornuti. Female genitalia: Papillae anales oval, setose. Lamella antevaginalis weakly sclerotized, loosely v-shaped transverse band. Lamella postvaginalis weakly sclerotized. Ductus bursae narrow, short, membranous. Corpus bursae elongated, without sclerotizations. Signum small and stellate or absent. Biology, habitat, phenology: Probably all species feeding on Juniper needles. Larva of E.  atlan­ticaria recorded on (Culot 1919) and found on Juniperus phoenicea (Corley 2004), that of E. sagnesi reared on Juniperus communis (Colomb 2005). Occurring in a variety of habitat types from coastal dunes to open pine forests to mountain slopes with scattered trees and bushes, from sea-level up to 1400 m. Adults of E. atlanticaria recorded from February to December in several generations (Redondo et al. 2009), the other species apparently bivoltine. Adults nocturnal. Immature stages: Larvae of E. sagnesi (see Colomb 2005) and E. atlanticaria green, needle-­ mimics matching the small twigs of their Juniperus foodplants. Both larvae with a row of red dots dorsally, interspersed with yellow and white markings both dorsally and laterally. D2 setae in pupal cremaster fused and elongated. 374

Remarks: Ekboarmia has recently been classified in the Ennominae, tribe Boarmiini (Vives Moreno 1994; 2014; Leraut 1997; Hausmann et al. 2004; 2011a). Pupal cremaster with setae D2 fused and elongated supporting this position. Systematic position: Boarmiini.

129. Ekboarmia atlanticaria (Staudinger, 1859) Boarmia atlanticaria Staudinger, 1859: Stettin. ent. Ztg. 20 (7–9): 218 (Spain, Andalusia: near Chiclana). Syntypes 3♂3♀ (MNHU, examined externally), 1 ♂ (Zoologisches Forschungs­ museum Alexander Koenig, Bonn, Germany; genitalia dissected by E. Wehrli, prep. number 5199, examined, including genitalia). One syntype ♂ and syntype ♀ are illustrated also in Albers & Warnecke (1941), plate II, figs. 9, 12 (on p. 332). Hemerophila atlanticaria holli Oberthür, 1909: Ètudes Lépid. comp. 3: 412, pl. 27, fig. 152 (Alge­ ria: Alger). Holotype ♂ (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany, genitalia dissected by E. Wehrli, prep. number 5193, examined, including genitalia). Tentatively validated at subspecies rank, status requiring further study. Boarmia atlanticaria gracilis Albers & Warnecke, 1941: Mitt. münch. ent. Ges. 31 (1): 115, pl. 4, fig. 40 (Morocco: Mehedya). Syntypes 4♂5♀ (2 syntypes ♂ in Universität Hamburg, Centrum für Naturkunde, Germany, examined externally, based on a photograph in the original publication). Junior synonym of Boarmia atlanticaria Staudinger, 1859 according to Skou et al. (2017a), here confirmed. Potentially unavailable names (infrasubspecific): pallidior: Lucas (1956) (f.). – atlanticaria gracilis: Albers & Warnecke (1941) (lapsus calami of incorrect year in Vives Moreno (2014): "1841").

External characters and abdomen: Wingspan 19–27 mm. Wings grey-brown, forewing ­medial area darker brown, general appearance somewhat smooth, little glossy. Forewing antemedial line inclined, deeply v-shaped, angled acutely towards base but often this part very narrow and weakly visible. Forewing medial line narrow, barely visible. Forewing postmedial line dark brown to blackish, inclined, weakly dentate on veins, often does not reach wing margin, deeply v-shaped and angled towards base near apex, angled again towards apex before reaching costa. Forewing postmedial line not angled outwards in inner margin and medial area narrow. Dark brown to blackish streak in forewing subapical area, often appearing as an extension of postmedial line. Outer margin of postmedial line bordered with whitish. ­Terminal line blackish, narrow, often discontinuous, stronger at vein endings. Fringes uniform grey-brown. Hindwing medial line weakly curved, strongest on inner margin. Hindwing postmedial line weakly dentate, outer margin whitish. Hindwing subterminal line brown, suffuse. Forewing discal spots blackish, small. Hindwing discal spots smaller. Wings below grey-brown, suffuse. Postmedial line, discal spots and terminal line blackish. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male, filiform in female. Male hindtibia with weak hair pencil, hindtibia with 2+2 spurs in both sexes. Tympanal organs medium-sized. Setal comb present on male abdominal sternite 3. Male 8th tergite weakly triangular, posterior margin narrower, other abdominal segments without modifications. Variation: Spring generation larger. Wings vary from grey-brown to different shades of brown. Medial area sometimes completely suffused with dark grey or brown. Terminal line continuous or discontinuous.

375

Text-figs 161–164. Diagnostic external characters (indicated) of three European and one non-­ European Ekboarmia Wehrli, 1943 species (after Skou et al. 2017a). See also Text-figs 169–171 for not-related but similar species Ecleora solieraria, Menophra thuriferaria and Peribatodes ara­ gonis. Text-fig. 161. E. atlanticaria (Staudinger, 1859), male, Portugal: Salema, 40 m, 19.iv.2007 (coll. Skou). Text-fig. 162. E. fascinataria (Staudinger, 1900), male, Algeria: Hammam Righa, April 1928 (coll. ZFMK). We illustrate this non-European species because it can potentially be found in Europe. Text-fig. 163. E. sagnesi Dufay, 1979, male, Spain: Valle de Luna, 1200 m, 2.vii.2011 (coll. Skou). Text-fig. 164. E. miniaria Skou, Stüning & Sihvonen 2017, male, Portugal: Grandola, Ameiras de B., 20.vi.2011 (coll. Skou).

Male genitalia: Uncus narrow, elongated, apex round. Socii absent. Gnathos arms fused ventrally, upturned, plate-shaped, surface rather smooth, margin round. Valva evenly narrowing towards apex, costa sclerotized, apical part of valva with narrow, setose ridge. Juxta arms large, rather narrow, dentate at apex only, base solid. Saccus elongated, upturned medially. Aedeagus narrow, caecum very short. Aedeagus apices bent when vesica is everted. Vesica enlarged at base, opens at about 135 degree angle, without cornuti. Female genitalia: Papillae anales elongated, setose. Apophyses posteriores slightly longer than apophyses anteriores. Lateral arms of lamella antevaginalis weakly sclerotized (Textfigs 165–167). Ductus bursae short, narrow, sclerotized laterally. Corpus bursae elongated, without sclerotizations. Signum distinctly stellate, small. Subsp. holli historically recognized for populations of Tunisia and Algeria. Holotype large, greyish brown, but in ZFMK a series of distinctly smaller specimens, collected later in the year, probably developed under dry conditions. DNA barcodes of Tunisian specimens diverging from other populations of E. atlanticaria (see Genetic data). No specimens from Algeria were available for DNA barcoding. Validity of taxon at subspecies level needs further research. 376

Text-figs 165–167. Diagnostic female genitalia characters (indicated) of European Ekboarmia Wehrli, 1943 species. Text-fig. 165. E. atlanticaria (Staudinger, 1859), Spain: Prov. Cádiz, Punta Paloma, 30 m, 22–23.v.2006, slide Sihvonen 2049 (coll. Skou). Text-fig. 166. E. sagnesi Dufay, 1979, France: Dept. Isère, Valle de la Romanche, 1200 m, 22–23.vi.1999, slide Sihvonen 2096 (coll. Skou). Text-fig. 167. E. miniaria Skou et al., Portugal: Grandola, 15.vi.2009, slide Sihvonen 2051 (paratype, coll. Skou). The shape of signum is diagnostic in E. atlanticaria and E. sagnesi, whereas the signum is absent in E. miniaria (see Genitalia plate).

Distribution and abundance: West Mediterranean. In Europe only in south-western Iberian peninsula, Balearic Islands and Sardinia. A single specimen recorded from southern France, undoubtedly a migrant from outside the known distribution area, of which the closest are the Balearic Islands. – In Europe somewhat local but not rare, found in limited numbers. – Outside Europe in Morocco, Algeria and Tunisia.

377

Phenology: Multivoltine. Recorded from early March to early November with a few additional records in late January, February and December. Larva found in February (­Staudinger 1859), in March and mid-October (M. Corley pers. comm.). Adults nocturnal, both sexes readily attracted to light. Biology: Larva monophagous. Found on Juniperus phoenicea (Staudinger 1859; Zangheri 1968; Corley 2004; D. Fritsch pers. comm.). Habitat: In a variety of habitat types where the foodplant occurs, including coastal dunes as well as non-sandy inland localities. From sea-level up to 300 m; according to Redondo et al. (2009) up to 500 m. Similar species: No similar species in Europe. North African E. fascinataria is similar and it can potentially be found in Europe. Forewing postmedial line weakly dentate and not angled outwards at termen in E. atlanticaria (forewing postmedial line strongly dentate or zigzagging and sharply angled outwards at termen in E. fascinataria). Forewing medial area narrow at termen in E. atlanticaria (forewing medial area wide at termen in E. fascinataria). Male and female genitalia of E. fascinataria and E. atlanticaria without diagnostic dif­ ferences. Not closely related Menophra harterti (Rothschild, 1912) with striated hindwings and continuous forewing postmedial line, reaching wing termen below apex. Not closely related Ecleora solieraria (Rambur, 1834) with pale medial area and rather straight hindwing postmedial line (see Flamigni et al. 2007 for illustrations) and Text-figs 169–171. Genetic data: BIN: BOLD:AAD7906 (n=11 from Morocco, Portugal and Spain including Balearics belonging to nominotypical subspecies). In Tunisia (putative subsp. holli) with two separate, sympatrically occurring BINs diverging by 1.0% (BOLD: ABZ3161, n=3) and by 2.2% (BOLD: ABZ6474, n=2) from the Iberian and Moroccan populations. E. sagnesi samples were nested within E. atlanticaria groups (only short sequences available for the former, see E. sagnesi). Nearest species: E. sagnesi (1.5%), E. miniaria (3.5%).

Ekboarmia fascinataria (Staudinger, 1900) Ekboarmia fascinataria (Staudinger, 1900), which is similar to E. atlanticaria, is known from Algeria and Morocco, and it could potentially be found in Europe. E. fascinataria has been recorded from France (Llauro, Department Pyrénées-Orientales by Bérard 1995) and it has been included in the French list of Lepidoptera (Leraut 1997) and in Fauna Europaea (Hausmann et al. 2004; 2011a). In our opinion the specimen in Bérard (1995) is E. atlanticaria and also Leraut (2009) states in his more recent publication that the species is not yet known from Europe. Further, a second specimen has been recorded from the Balearic Islands in Spain (Expósito Hermosa 2007) but identity of the specimen has remained unconfirmed. Vives Moreno (2014) considers taxon fascinataria sensu Expósito, 1998 a junior synonym of E. atlanticaria subsp. gracilis (Albers & Warnecke, 1941). The male and female genitalia of E. fascinataria and E. atlanticaria are without diagnostic differences. See Textfigs 161, 162, 165 and 168. E. fascinataria is rare in collections and there are no recent records. The most recent record we have managed to locate is from Algeria: Prov. Alger, Gorges de la Chiffa 200–400 m, 6.vi.1979 (in coll. Sommerer). E. fascinataria is probably bivoltine, specimens have been observed in February, March, May, June and in October. Larval time and hibernation unknown. Larva potentially feeding on Juniperus needles as do other Ekboarmia species. Adults probably nocturnal. 378

Text-fig. 168. Adults and genitalia of Ekboarmia fascinataria (Staudinger, 1900), a species that could potentially be found in Europe (diagnostic characters are indicated, figures are not in scale). See also Text-figs 161–167. Male and female genitalia of E. fascinataria and E. atlanticaria without diagnostic differences. Male adult (wingspan 25 mm): Algeria: Hammam Righa, April 1928 (coll. ZFMK, illustrated in Wehrli (1939–1954) also). Female adult (holotype, wingspan 29 mm): Algeria: Teniet el Had, May-June (coll. MNHU). Male genitalia, vesica and juxta illustrated from the same specimen: Algeria: Hammam Righa, May 1928 (slide D. Stüning 2307), aedeagus illustrated from a specimen: Algeria: Hammam Righa, June 1928 (dissected by E. Wehrli, slide 5919 (in glycerol), mounted on slide by D. Stüning in 2015), all specimens in coll. ZFMK, photographed by D. Stüning. Female genitalia (point of origin of ductus seminalis indicated by circle), signum and ostium bursae and adjacent structures (all pictures illustrated from the same specimen): Algeria: Hammam Righa, May 1928 (slide D. Stüning 2310), all specimens in coll. ZFMK, photographed by D. Stüning.

379

130. Ekboarmia sagnesi Dufay, 1979 Ekboarmia sagnesi Dufay, 1979: Bull. Soc. ent. Fr. 84: 129, figs 1–3 (France: Hautes Alpes, high valley of the Romance). Holotype ♂ (MNHN, examined externally, illustrated in Leraut (2009, plate 47, fig. 18)), paratype ♂ genitalia illustrated in Dufay (1979), 7♂ paratypes (coll P. Sagnes, L’Arbresle, Rhône, France, one specimen examined, including genitalia), 1♂ paratype (coll. C. Dufay, Chaponost, Rhône, France), 1♂ paratype (coll. Herbulot in ZSM, Germany, France: Les Fréaux, près La Grave, examined externally). Ekboarmia sagnesi herrerai Expósito Hermosa, 2007: SHILAP Revta. lepid. 35 (138): 269, fig. 1–2 (Spain, Andalusia: Jaén, Hornillos, de Puente de Guada). Holotype ♂ (coll. A. ­Expósito Hermosa, Móstoles, Madrid, Spain, genitalia examined). Junior synonymy proposed by ­Leraut (2009), here confirmed.

External characters and abdomen (adopted from Skou et al. 2017a): Wingspan 25–29 mm, largest species in the genus. Wings dark grey, sometimes with chocolate-brown tinge, rather uniform in colour. Forewing antemedial line black, narrow, deeply angled inwards before costa. Medial line narrow, barely visible and rather straight, strongest near inner margin. Postmedial line distinct, particularly at inner margin, black, weakly dentate, bordered white outside, not reaching termen, weakly v-shaped and angled towards base subapically, angled again towards costa. Postmedial line not angled outwards near inner margin and medial area rather narrow. Hindwing medial line faint, almost straight, not reaching costa. Hindwing postmedial line distinct, black, weakly dentate, distally bordered with white, weak or absent near costa. Subterminal line absent or very faint. Terminal line faint, continuous. Fringes concolorous with wings. Discal spots weak or absent. Wings below uniform pale greyish brown, postmedial line and discal spots most visible, antemedial line of forewing absent. Course of postmedial line in forewing does not conform to upperside. Frons, collar, thorax and abdomen concolorous with wings. Other structures as in E. atlanticaria, see above. Male abdominal sternites 2 + 3 with sterno-tympanal processes and setal comb illustrated in Skou et al. (2017a). Variation: Forewing medial area concolorous with wings or darker, particularly near inner margin. Forewing postmedial line rather straight or distinctly dentate. Male genitalia: Generally as E. atlanticaria (see above). Setose ridge extends into medial part of valva in E. sagnesi (medial ridge absent in other Ekboarmia species). Vesica opens at 90 degree angle (at 135 degree angle in E. atlanticaria and E. fascinataria). Juxta arms broader distally than basally, dentate or smooth along inner margin, always dentate distally, base with roundish lobe in E. sagnesi (juxta arms gradually tapered towards apex, margin dentate at apex only in E. atlanticaria and E. fascinataria, base with elongated lobe in E. miniaria). Female genitalia: Generally as E. atlanticaria (see above). Lamella antevaginalis broader. Signum small, weakly stellate in E. sagnesi (signum distinctly stellate in E. atlanticaria and E. fascinataria, signum absent in E. miniaria), see Text-figs 165–167. Distribution and abundance: (South-west) European. Only few and isolated populations are known: In Val de la Romanche, Dept. Hautes-Alpes; St. Baume, Dept. Var, France; from Valle de Luna and Velilla de Rio Carrión (Gaston pers. comm.), both in Leon Province; and Sierra de Cazorla, Jaén Province, Spain. – Apparently extremely local, appearing as single specimens or in limited numbers.

380

Phenology: Uni- or bivoltine. In Spain from early June to mid-July, in France from early May to late June and in August. Larval period and hibernation stage unknown. Adults nocturnal, attracted to light. Biology: Larva monophagous on Juniperus. Reared on Juniperus communis (C. Tautel pers. comm.; Colomb 2005), but other Juniperus species are possible foodplants. Habitat: Mountain slopes with scattered trees and bushes. In France from 1150 up to 1400 m, in Dept. Var by St. Baume at only 700 m (J. Junnilainen pers. comm.) and in Spain known from around 1200 m. In Spain (Leon) found on slopes with Juniperus sabina (T. Molina pers. comm.). Similar species: No similar species in Europe. Rather uniform brown colour and small angle in forewing postmedial line near costa are diagnostic. Genetic data: No BIN assigned yet (n=3 from France, Spain, short sequences, 127–273 bp; including the holotype of taxon herrerai). Genetically grouping near or within the cluster of E. atlanticaria. Nearest species: E. atlanticaria (1.5% from Spanish populations; 2.0% from Tunisian populations); E. miniaria (3.9%). The barcodes of E. sagnesi are incomplete, so the genetic results must be considered tentative. Remarks: Colomb (2005) illustrated a superficial, hand-drawn picture of the female genitalia and it had an unusual curved, potentially sclerotized structure on the corpus bursae. We have not observed such structure in any of the examined E. sagnesi specimens and the signum (which Colomb apparently did not illustrate) of E. sagnesi is weakly stellate, not resembling such curved structure. See Skou et al. (2017a) for details.

381

131. Ekboarmia miniaria Skou, Stüning & Sihvonen, 2017 Ekboarmia miniaria Skou, Stüning & Sihvonen, 2017: Nota lepid. 40: 58, figs. 12, 16, 21, 25 (Portugal: Grandola, Ameiras de Baixo). Holotype ♂ (coll. Skou, Denmark, to be deposited at the Zoological Museum, University of Copenhagen, Copenhagen, Denmark, examined, including genitalia, slide Sihvonen 2050). Paratypes 5♂ 5♀ (Portugal: Grandola, Ameiras de Baixo; Portugal: Alentejo, Lagoa de Santo André). All paratypes currently in coll. Skou, one female will be deposited at ZFMK, one male at NHMUK, one male at ZSM and all remaining specimens will be deposited at Zoological Museum, University of Copenhagen, Denmark (all paratypes examined).

External characters and abdomen: Wingspan males 17–19 mm (n=6), females 17–18 mm (n=5), smallest species in the genus. Sexually dimorphic. Male dark greyish brown or blackish brown, medial area darker. Medial line blackish, weakly dentate, turned inwards on costa. Postmedial line distinctly dentate, angled inwards before costa, distally narrowly bordered with whitish. Medial area dark, wide on inner margin. Subterminal line absent. Terminal line narrow, blackish, slightly widened at vein endings. Fringes grey-brown, unicolorous. Hindwing medial line diffuse, postmedial line weakly dentate. Forewing discal spot small, distinct. Wings below uniform grey-brown, postmedial line weakly visible. Female almost uniform grey-brown, postmedial line weakly visible, dark brown, dentate. Forewing discal spot small, distinct, dark brown. Hindwing discal spots smaller. Wings below uniform greybrown, discal spots minute and dark brown. Frons, collar, thorax and abdomen concolorous with wings, irrorated with grey. Abdomen paler ventrally. Male antennae bipectinate, female antennae filiform. Hindtibia with 2+2 spurs, male hindtibia not swollen, with very small hair pencil (if not everted, a weak groove visible only). Tympanal organs medium-sized, not meeting medially, slightly smaller in female. Male 8th tergite weakly triangular, posterior margin narrower. Other sternites and tergites of both sexes unmodified. Variation: June specimens are dark brown, the single specimen taken in October is light brown, except terminal area, which is distinctly dark brown. It is not clear whether this specimen shows the normal appearance of the second generation or whether it is a strongly aberrant individual. Male genitalia: Generally as in E. atlanticaria, but genitalia smaller. Uncus wide in E. miniaria (uncus narrow in E. atlanticaria and E. sagnesi). Aedeagus apex not bent after everting vesica (aedeagus apex bent after everting vesica in E. atlanticaria and E. sagnesi). Vesica opens at about 90 degree angle (vesica opens at about 135 degree angle in E. atlanticaria). Juxta arms dentate along inner margin, base with elongated lobe (juxta arms dentate at apex only, base solid in E. atlanticaria; juxta arms dentate (or smooth) along inner margin, base with roundish lobe in E. sagnesi). Female genitalia: Generally as in E. atlanticaria, but genitalia smaller. Lateral arms of lamella antevaginalis reduced in E. miniaria (lateral arms of lamella antevaginalis weakly sclerotized in E. atlanticaria and strongly sclerotized in E. sagnesi) (Text-figs 165–167). Signum absent in E. miniaria (signum distinctly stellate in E. atlanticaria and weakly stellate in E. sagnesi). Distribution and abundance: Endemic to Portugal, where it is presently known from only three localities in the southern part of the country. In addition to the type specimens, a single male has been found from Almograve (D. Grundy pers. comm.). – Occasionally in numbers, e.g. seven specimens were found in one night (20.vi.2011).

382

Phenology: Possibly bivoltine. So far recorded in mid-May (D. Grundy pers. comm.), in mid-June and once between 16th September and 10th October. Larval time and hibernation stage unknown. Adults nocturnal, attracted to light. Biology: Unknown. Habitat: Open pine forests (Pinus pinaster) on sandy soil with undergrowth of herbs and scattered bushes, including Halimium atriplicifolium, Stauracanthus genistoides and Cistus psilosepalus among the dominant species. One of the three known localities had suffered a bushfire, probably 8–10 years earlier. Found close to sea-level and at 90 m. Similar species: No similar species in Europe. E. miniaria female shows some resemblance to weakly patterned Eupithecia species (Larentiinae), but these are structurally different. Genetic data (after Skou et al. 2017a): BIN: BOLD:AAZ6253 (n=2 from Portugal). Intra­ specific variation low (0.15%). Distances from E. atlanticaria 3.5% and from E. sagnesi 3.9%.

Ecleora Wehrli, 1941 Ecleora Wehrli, 1941, Gross-Schmett. Erde 4 (Suppl.): 472. Type species: Boarmia solieraria Rambur, 1834. There are no genus synonyms.

Diversity and distribution: The genus Ecleora includes three species in the Palaearctic region (Scoble 1999). External characters and abdomen: Medium-sized, typical boarmiine moths with brownish grey ground colour. Proboscis vestigial. Fovea absent. Forewing with radial veins 1 and 2 shortly stalked, not free from cell (Wehrli 1939−1954).

383

Male genitalia: Uncus double-pointed, Gnathos narrow. Valva with sclerotized harpe, with teeth. Juxta double-pointed. Aedeagus without cornuti. Female genitalia: Apophyses posteriores of medium length, apophyses anteriores shorter. Corpus bursae membranous, narrow, very long, partly striated, without signum. Lamella antevaginalis alate, lamella postvaginalis developed as flat sclerite. Remarks: Ekboarmia and Ecleora appear as sister taxa in multi-gene analysis (Murillo-Ramos et al. 2019).

132. Ecleora solieraria (Rambur, 1834) Boarmia solieraria Rambur, 1834: Annls Soc. ent. Fr. 3 (2): 390, pl. 8, fig. 5 (France). Lectotype ♂ (ZSM). Boarmia solieraria f. nana Fernández, 1933: Boln Soc. esp. Hist. nat. 33: 373, pl. 26, fig. 6 (Spain: La Vid.). Syntype(s).

External characters and abdomen: Wingspan first generation: ♂ 21–27 mm, ♀ 25–32 mm, second generation 2–3 mm smaller. Ground colour brownish grey. Fringes concolorous with ground colour. Terminal line black. On forewing post- and antemedial lines black, oblique, almost parallel, ending before reaching costa. Postmedial line on both wings edged by brownish band. On forewing this band is continued by black streak to apex. Medial area with distinct discal spot. Terminal area grey with often inconspicuous subterminal line. Hindwing with black, slightly bent postmedial line and greyish brown medial and subterminal lines, usually without or with indistinct discal spot. Underside with distinct postmedial line, subterminal and medial lines weaker, discal spots on both wings present. Frons dark brown, vertex whitish, collar scales grey with dark brown tips. Thorax and tegula whitish, abdomen concolorous with thorax, each segment with small dark brown rings. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Little. Wing pattern can be darker or lighter. Male genitalia: Uncus triangular with short double tip. Gnathos arms fused ventrally, upturned, tip small, rounded. Valva short, divided. Dorsal arm sclerotized, narrow, expanded at apex, costa and apex setose. Ventral arm with sclerotized margin, horn-shaped at tip, at centre with a dorsally directed, pointed process and a large sclerotized, nearly oval plate; the latter with conspicuous teeth at dorsal margin. Juxta large, sides parallel, posteriorly with two large tips, in between deeply concave. Saccus rounded, margin concave. Aedeagus cylindrical, apex with slightly bent, finger-shaped and tapered band-shaped process, without cornuti. Female genitalia: Papillae anales oval, slightly sclerotized, densely covered with setae. Length of apophyses posteriores on average 1.15 mm, apophyses anteriores shorter (on average 0.58 mm), at tergum 8 triangularly dilated. Corpus bursae membranous, narrow, very long, partly striated, without signum. Lamella antevaginalis alate, lamella postvaginalis nearly rectangular plate, both sclerotized. Distribution and abundance: South-west European. Distributed only in southern France and eastern Spain with a single isolated population in southern Portugal (E. Marabuto pers. comm.), not distributed across the whole peninsula as suggested by Leraut (2009). – Rather common, sometimes occurring in numbers. – Unknown from outside Europe despite a general record for North Africa by Redondo et al. (2009). Not listed for the fauna of Morocco by Rungs (1981). We have not been able to trace any North African record. 384

Phenology: Bivoltine. From early April to late June and from mid-July to late September, possibly with overlap of the two generations. A last stage larva in early November (C. Tautel pers. comm.). Adults active at night, both sexes readily attracted to light. Biology: Larva oligophagous. Found on Juniperus phoenicea (J. Gelbrecht pers. comm.). Recorded on Juniperus thurifera (Gómez de Aizpúrua 2002), Cupressus and Spartium junceum (Redondo et al. 2009). Reared on Juniperus chinensis and J. horizontalis (M. Leipnitz pers. comm.). Pupation on the ground in plant litter (Gómez de Aizpúrua 2002). Habitat: Xerothermophilous. In a variety of open habitats with more or less scattered trees and bushes on both sandy and rocky soil as well as gypsophilous ground. From 200 up to 1600 m altitude. Similar species: Menophra thuriferaria (Zerny, 1927), M. harterti (Rothschild, 1912) and Peribatodes aragonis (Wehrli, 1943). Diagnostic characters for these three species see Textfigs 169–171. M. harterti cannot be separated from M. thuriferaria with certainty by external features. Ekboarmia atlanticaria is also similar (see Text-fig. 161). Genetic data: BIN: BOLD:ABA9342 (n=3 from France and Spain). Genetically homogeneous. Nearest species: Apophyga sericea Warren, 1893 from India (6.6%).

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Text-figs 169–171. Diagnostic characters (indicated) of similar Iberian species. Text-fig. 169: Ecleora solieraria (Rambur, 1834), Spain: Aragón, Albarracín, 11.vi.2007. Text-fig. 170: Menophra thuriferaria (Zerny, 1927), Spain: Aragón, Albarracín, 11.vi.2007. Text-fig. 171: Peribatodes aragonis (Wehrli, 1943), Spain: Aragón, Albarracín, 06.ix.2009 e.o. (all coll. B. Müller.

Afriberina Wehrli, 1943 Afriberina Wehrli, 1943, Gross-Schmett. Erde 4 (Suppl.): 473. Type species: Boarmia tenietaria Staudinger, 1900.

Diversity and distribution: Six putative species in the West Mediterranean area, including the Iberian peninsula, Hautes-Alpes in France, Morocco, Algeria and Tunisia. Central Asian 386

‘Afriberina’ nobilitaria (Staudinger, 1892) was recently transferred to Alcis (Tautel et al. 2016). Taxonomy of the genus is notorious and in urgent need of a modern revision. External characters and abdomen: Medium-sized, wingspan 25–31 mm. Wings grey to greybrown, markings blackish. Postmedial line prominent, on forewing oblique and turned towards costa near apex. Postmedial line often bordered with dark fascia, followed by pale wavy line. Medial line often prominent on hindwing, but diffuse. Discal spots distinct on underside. Male antennae bipectinate, female antennae filiform. Hindtibia of both sexes with 2+2 spurs, male hindtibia without hair pencil. Tympanal organs medium-sized. 3–4 intersegmental abdominal membrane with sacs (Text-fig. 172), other abdominal segments of both sexes without distinct modifications.

Text-fig. 172. Membrane sacs of Afriberina tenietaria (Staudinger, 1900), located on 3–4 inter­ segmental abdominal membrane. Similar sacs are found in all European Afriberina, potentially in North African taxa also. These sacs are not restricted to Afriberina because small and inconspicuous extensions are found commonly throughout the Geometridae, but such distinctive sacs are unusual. Spain: Prov. Teruel, 4 km W Albarracin, 1185 m, 7.viii.2007, slide Sihvonen 2147 (coll. Skou).

Male genitalia: Uncus wide, apex round. Socii absent. Gnathos arms sclerotized, fused, apex upturned, granulate. Valva wide, ventral and dorsal margin (costa) sclerotized, apex membranous and round. Costa ridge setose subapically, curved ventrally. Medio-ventral part of valva (harpe) with distinct tooth and setose ridge. Juxta sclerotized, constricted medi­ ally. Aedeagus with long caecum, apex with dentate ridge. Vesica long, narrow, with 3–4 apical cornuti. Female genitalia: Papillae anales and segment 9 elongated. Apophyses posteriores extremely long, apex of apophyses anteriores widened, spatulate. Posterior margin of antrum concave. Ductus bursae sclerotized, extremely long. Posterior part of corpus bursae wrinkled, anterior part membranous sac, turned laterally. Biology: Larva of A. tenietaria feeding on Juniperus sabina and J. thurifera (Gómez de Aizpúrua 2002), potentially other species of the genus may feed on Juniperus also. Adults inhabiting Mediterranean woodland with Juniperus, from sea-level up to 1500 m. E ­ uropean species with two generations per year. Adults nocturnal, attracted to light. 387

Immature stages: Eggs of an unidentified Afriberina species from Morocco green-brown, with longitudinal rows of tiny depressions (Euroleps 2018). Larvae of A. tenietaria and A. salemae cryptic, well camouflaged among twigs and needles of Juniperus: colour varying from green to brown to pinkish brown, with diamond-shaped pattern dorsally, interspersed by whitish markings (Gómez de Aizpúrua 2002, see A. salemae below). Pupa brown, cremaster details unknown (Gómez de Aizpúrua 2002). Remarks: Recently Afriberina has been classified in the Boarmiini (Vives Moreno 1994; Hausmann et al. 2004; 2011; Vives Moreno 2014). Male genitalia of some Nychiodes L ­ ederer, 1853 similar to those of Afriberina, particularly N. dalmatina Wagner, 1909 (lectotype genitalia illustrated in Flamigni et al. 2007), but female genitalia of Afriberina unlike any other genus in Europe. Systematic position: Boarmiini. Taxonomy of Afriberina is notoriously difficult and in urgent need of a modern revision involving the North African taxa, and the matters are complicated by large number of named subspecies (see Scoble 1999; Leraut 2009). External appearance of taxa varies quite a lot, but the genitalia are very uniform throughout the genus and thus of little diagnostic value. Preliminary work on the DNA barcodes seems to show constant correlations with external characters, but the differences between putative species and subspecies are very small, often < 1%, which is considerably lower than usual average values between congeneric species in European Geometridae (Hausmann et al. 2013). It would be desirable to DNA barcode as many name bearing types as possible.

133. Afriberina tenietaria (Staudinger, 1900) Boarmia tenietaria Staudinger, 1900: Dt. ent. Z. Iris 12: 394, pl. 6, fig. 1 (Algeria: Teniet el Had). Syntypes 2♂ (MNHU, examined, including genitalia, one syntype DNA barcoded: BC ZSM Lep 93741_NGS). Boarmia tenietaria var. punctata Fernández, 1933: Boln Soc. esp. Hist. nat. 33: 374 (Spain: Albarracin, La Vid). In recent literature it has been considered a junior synonym of tenietaria Staudinger, 1900 (Scoble 1999), valid at subspecies level (Leraut 2009), or not mentioned at all (Redondo et al. 2009). We consider it a junior synonym of tenietaria, but more research is needed (see Remarks under genus chapter). Boarmia tenietaria fasciata Albers & Warnecke, 1941: Mitt. münch. ent. Ges. 31 (1): 119 (Spain: Albarracin). Holotype ♂. In recent literature it has been considered a junior synonym of punctata Fernández, 1933 (Leraut 2009), valid at subspecies level (Scoble 1999) or a junior synonym of tenietaria Staudinger, 1900 (Redondo et al. 2009). We consider it a junior synonym of tenietaria, but more research is needed (see Remarks under genus chapter). Afriberina tenietaria hammamrirha Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 475 (Algeria: Hammam Rirha). Syntypes, including ♀ (ZFMK). Valid at subspecies level (Scoble 1999, Leraut 2009). Potentially unavailable names (infrasubspecific): junia: Wehrli (1943) (var.), lineata: Fernández (1933) (ab.), melaleucaria: Zerny (1927) (ab.).

External characters and abdomen: Wingspan first generation males 28–34 mm, females 34–36 mm (n=3), second generation males 24–25 mm (n=2), females no data. Females are larger than males, also specimens in central Spain are on average 2 mm larger than those in the southern Iberian peninsula. Wings various shades of grey, markings blackish brown. Forewing antemedial line straight, curved towards costa (often curved part not visible). Medial line diffuse. Postmedial line strongest, oblique, weakly concave, curved towards costa before apex. Postmedial line bordered with dark brown band, followed by pale wavy 388

line and light brown band. Terminal line distinct, continuous, blackish. Fringes concolorous with wings. Hindwings with diffuse medial line and distinct, weakly concave postmedial line. Postmedial line bordered with dark brown band. Discal spots blackish, often weak or absent on forewings. Wings below uniform greyish, except large discal spots, blackish terminal line and faint postmedial line that are distinct. Frons brown, collar, thorax and abdomen concolorous with wings. Male antennae with long pectinations, female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Tympanal organs and abdomen as in genus description. Variation: Variable. Wing colour varies from whitish grey to grey (common in central Spain and in France: Hautes-Alpes) to grey-brown (common in southern Iberian peninsula). Forewing postmedial line can be wide or very narrow, rather straight, distinctly concave, and it can meet inner margin as bent inwards (most common) or straight. Postmedial line is often bordered with dark brown area but this can be almost absent. Discal spots can be distinct and blackish or almost absent (absence is quite common in southern Iberian peninsula). Wings below often uniform greyish in males, but with dark suffusion in terminal area, particularly near forewing apex in females. In southern Spain both sexes have a pale blotch on forewing underside on terminal area, more pronounced in females (Text-figs 177–178). Male genitalia: Uncus wide, apex blunt. Socii absent. Gnathos arms wide, sclerotized, fused, apex upturned, granulate. Valva wide, ventral and dorsal margin (costa) sclerotized, apex membranous and round. Costa ridge setose subapically, curved ventrally. Medio-ventral part of valva (harpe) with distinct spike pointing towards valva apex, followed by setose ridge. Juxta sclerotized, base wide, constricted medially, apex wide again and margins weakly upturned. Saccus wide, round. Aedeagus with long caecum, apex with dentate ridge. Vesica long, narrow, grooved longitudinally, with 3–4 apical cornuti. Female genitalia: Papillae anales striated, weakly sclerotized, elongated. Segment 9 elongated. Apophyses posteriores extremely long, apex of apophyses anteriores widened, spatulate. Lamella postvaginalis membranous. Lamella antevaginalis membranous, lunular. Ostium bursae sclerotized, narrow, ventral margin concave. Ductus bursae sclerotized, narrow, extremely long. Posterior part of corpus bursae wrinkled, anterior part membranous sac, turned laterally, without signum. Distribution and abundance: West Mediterranean (Ibero-Magrebian). In Europe only in the Iberian peninsula with one remote population in south-eastern France. The distribution map in Leraut (2009) show a distribution in the whole of Spain though the species only occurs in central Spain and along the south-western Atlantic coast. – Outside Europe in Algeria (nominotypical locality), recorded from Morocco and Tunisia (Rungs 1981). – In the Iberian peninsula local, but sometimes in numbers at its localities. Phenology: Uni- and bivoltine. In northern central Spain in one generation from early June to mid-August and in south-western Spain in two generations from late March to early June and again in September and October. In southern France found in early July. Hibernation as half grown larva. Active at night, both sexes readily attracted to light. Biology: Larva monophagous. Recorded on Juniperus thurifera (Redondo et al. 2009), in Portugal occurring in places where J. phoenica is present (M. Corley pers. comm.). Rearedon Juniperus chinensis and J. horizontalis (M. Leipnitz pers. comm.). In rearings pupation takes place in a cocoon on the ground. Habitat: Xerothermophilous. In northern central Spain in open wood- and scrubland, from 300 to 1500 m. In south-western Iberian peninsula in dunes and other places with sandy 389

soil close to sea-level. In Portugal and France in rocky places. In France between 900 and 1000 m, in Portugal in lowlands. Similar species: Afriberina salemae Skou & Sihvonen (see below). In Menophra thuriferaria (Zerny, 1927) and Ecleora solieraria (Rambur, 1834) forewing apex is with a dark stripe, and in M. harterti the hindwing base is striated. Genetic data: BIN: BOLD:AAD3818 (n=4 from Spain, one further from Algeria: syntype). Genetically slightly heterogeneous, maximum variation on the Iberian peninsula 0.77%. Algerian syntype diverging by 1.2% from the DNA barcode of Andalusian A. tenietaria. 30 further DNA barcoded specimens from Morocco and Tunisia BIN-sharing, but diverging in several clusters at minimum distances of 1.6–2.2%, but doubtfully conspecific (probably referring to A. zernyi Wehrli, 1943, A. rungsi (Albers & Warnecke, 1941) and A. terraria (Bang-Haas, 1907)). Nearest species: Afriberina salemae (0.76%) and North African ­congeners.

Afriberina terraria (Bang-Haas, 1907) The identity of Afriberina terraria and its occurrence in Europe are doubtful, and it has been considered to potentially be a junior synonym of A. tenietaria (Staudinger, 1900) (Albers & Warnecke 1941), an externally variable species. Boarmia terraria Bang-Haas was described from Algeria: Southern Oran (Bang-Haas 1907), see Text-fig. 174. The holotype is externally different from A. tenietaria, whose two syntype males are from Algeria also, but the genitalia are without noticeable differences (examined, our observation). It must be considered, however, that the genitalia in genus Afriberina are uniform and not very informative (see illustrations in Leraut 2009). Both A. tenietaria and A. terraria show a valva hook, which is directed towards the valva apex. A. terraria has got putative records from Spain and 390

Portugal (for instance Vives Moreno 1994; Redondo et al. 2009; Leraut 2009), but due to the difficulties in identification, the records have remained dubious. Redondo et al. (2009) examined a male and a female of A. terraria from Algeria: Guelt-es-Stel (in coll. ZSM) and concluded (p. 73): “the female genitalia give good genital characters (particularly ductus bursae); this is not the case with the male genitalia”. They illustrate hand-drawn pictures of both male and female genitalia but without mentioning what the diagnostic characters are. We have not noticed diagnostic differences in the illustrations. Holotype of A. terraria (Bang-Haas, 1907) and a syntype of A. tenietaria (Staudinger, 1900) were submitted to Next Generation Sequencing (NGS) analysis, which was only successful for the latter, unfortunately. Until identity of A. terraria has been cleared, we consider the European records doubtful and do not give A. terraria a full species treatment.

Text-figs 173–174. The identity of Afriberina terraria and its occurrence in Europe are doubtful. The genitalia of the illustrated type specimens are without noticeable differences (our observation), but it must be considered that the genitalia in genus Afriberina are uniform and not very informative. Until identity of A. terraria has been cleared, we consider the European records doubtful and do not give A. terraria a full species treatment. Text-fig. 173. A. tenietaria (­Staudinger, 1900), syntype, Algeria: Teniet el Had, assumedly taken in May-June in 1898. Text-fig. 174. A. terraria (BangHaas, 1907), holotype, Algeria: Southern Oran, 30.vi.1905. Both illustrated specimens in MNHU.

134. Afriberina salemae Skou & Sihvonen, sp. n. Holotype ♂: Portugal,/ Prov. Algarve,/ Salema Camping, 55 m,/ 21.iv.2017,/ Peder Skou leg. (currently in coll. Skou, Denmark, to be deposited at Zoological Museum, University of Copen­ hagen, Denmark); Afriberina salemae/ Skou & Sihvonen/ HOLOTYPE [red rectangle label]. Holotype illustrated on Text-fig. 179 and on the Colour plate. Paratypes (52 males 4 females): 1♂ Portugal, Prov. Algarve,/ 1 km N Salema,/ Camping place, 30 m,/ 22.–23.ix.2009,/ Peder Skou leg. (dissected, slide Sihvonen 2149) (coll. Skou, Denmark). 1♂ Portugal,/ Prov. Algarve,/ Salema, 40 m,/ 19.iv.2007,/ Peder Skou leg. (dissected, slide Sihvonen 2247). 24♂3♀ Portugal, Prov. Algarve,/ Salema Camping, 55 m,/ 21–22.iv.2017,/ Peder Skou leg. (coll. Skou, Denmark) (1♂ dissected, slide Sihvonen 2240, 1♀ dissected, slide Sihvonen 2241). 13♂ Portugal, Algarve/ 15 km N Vilamoura/ 80 m, 10.–15.7.1991/ leg. J. Lenz (one with DNA barcode BC ZSM Lep 09597) (coll. ZSM). 6♂ id., 18.10.1991/ leg. J. Lenz (one dissected gen. prp. ZSM G 12687; one with DNA barcode BC ZSM Lep 09596) (coll. ZSM). 5♂1♀ id., 20.4.1991/ leg. J. Lenz (coll. ZSM). 1♂ [Portugal] P4025/ Vale

391

da Fonte/ Algarve/8.6.[19]96/ M. F. V. Corley (dissected, slide Sihvonen 2257) (coll. Corley, Portugal). 1♂ Portugal/ Vale Santo,/ Sagres, Algarve/ 7. x. 1999,/ M. F. V. Corley (coll. Corley, Portugal). All paratype labelled with red or yellow rectangle label: Afriberina salemae/ Skou & Sihvonen/ PARATYPE.

Etymology: The species name salemae refers to a small coastal town Salema in Algarve Province on the southern coast of Portugal, from where numerous specimens have been found. Diagnosis: Similar to Afriberina tenietaria (Staudinger, 1900), specimens from south-western Spain, but with the following differences (A. tenietaria characters in parentheses). External characters: Forewing postmedial line reduced to dots (line continuous), forewing discal spot often strong (spot may be absent), wings brownish grey (grey), fascia on terminal area paler brown (dark brown), forewing postmedial line not visible on underside (visible), female forewing apex weakly darkened on underside (dark), female terminal area with weak blotch on underside (with pale blotch), see Text-figs 180–181. Male genitalia: Medio-ventral part of valva (harpe) with distinct spike curved inwards, pointing towards costa (spike pointing towards valva apex). Female genitalia: No diagnostic differences observed. Genetic data (COI barcode): 0.76% divergence between A. salemae and closest haplotypes of A. tenietaria. Description, external characters and abdomen: Wingspan in first generation males 27–32 mm, females 34 mm (n=3), in second generation males 26–28 mm, females no data. Generally as A. tenietaria (Staudinger, 1900), see Diagnosis above and Text-figs 179–181. Variation: Rather little. Second generation males slightly smaller (second generation females currently not available, but potentially similarly smaller, too). Fascia on terminal area usually pale brown, sometimes dark brown. One examined male specimen with forewing postmedial line more or less continuous (Portugal: Vale da Fonte, coll. Corley), similar to A. tenietaria, but genitalia typical for A. salemae.

Text-fig. 175. Larva of Afriberina salemae Skou & Sihvonen sp. n., from Portugal: Sagres, Cabo de Sao Vicente, 25.ii.2012, reared on Juniperus phoenicea. Larva is well camouflaged to the food-plant: colour varies from green to pinkish, it has diamond-shaped pattern dorsally and whitish markings laterally. Specimen reared and photographed by Dieter Fritsch.

392

Text-figs 176–181. Diagnostic characters (indicated) of Afriberina tenietaria (Staudinger, 1900) and A. salemae Skou & Sihvonen sp. n. We illustrate A. tenietaria from south-western Spain because specimens in that area are the most similar to A. salemae. A. tenietaria is paler in Central Spain and in France, see the Colour plates. Text-fig. 176. A. tenietaria male. Spain: Prov. Huelva, Matalascañas, 35 m, 18.iv.2017 (coll. Skou). Text-fig. 177. A. tenietaria male underside, same specimen. Text-fig. 178. A. tenietaria female underside. Spain: Prov. Huelva, Matalascañas, 35 m, 18.iv.2017 (coll. Skou). Text-fig. 179. A. salemae male. Portugal: Algarve, Salema Camping, 55 m, 21.iv.2017 (holotype, coll. Skou). Text-fig. 180. A. salemae male underside, same specimen (holotype). Text-fig. 181. A. salemae female underside. Portugal: Algarve, Salema, 55 m, 21.iv.2017 (paratype, coll. Skou). The course of forewing postmedial line is rather variable in A. tenietaria and it can meet inner margin as angled inwards (most common) or straight (rare).

393

Description, male genitalia: Generally as A. tenietaria (Staudinger, 1900), but with following differences: Medio-ventral part of valva (harpe) with distinct spike curved inwards, pointing towards costa (spike pointing towards valva apex in A. tenietaria). Description, female genitalia: As in A. tenietaria (Staudinger, 1900) (see above), no diagnostic differences observed. Angle between ductus bursae and corpus bursae variable and not diagnostic (see Genitalia plate). It is always tilted to left in ventral view. Distribution and abundance: Portuguese endemic, so far known from Algarve and Estremadura provinces. – Apparently relatively common, but usually found in limited numbers.

Phenology: Bi- or trivoltine. So far recorded from early and mid-March, early April to midMay, early and late June, mid-July to late August and from late September to mid-November. Larva found in mid- and late March (D. Fritsch pers. comm.). Biology: Larva monophagous. Found on Juniperus phoenicea (D.  Fritsch pers. comm.) (Text-fig. 175). Habitat: Macchia with Mediterranean vegetation, close to sea-level. Similar species: Afriberina tenietaria (Staudinger, 1900), particularly in southern Iberian peninsula. Diagnostic characters are explained above. Genetic data: Genetically homogeneous (n=2 from southern Portugal). Nearest species: Afriberina tenietaria (Staudinger, 1900) (0.76%) and North African congeners (see A. tenietaria above). Remarks: A. salemae has so far been found in a geographically restricted area in south-­ western Portugal. The previous Afriberina records from that area are likely to be A.  sale­mae, 394

and we have already discovered numerous photographs of this species on the internet. The genetic and genitalia differences are small in genus Afriberina, when compared to the Geometridae in general, but the external features are more diagnostic. The diagnostic differences of A. salemae are also small, but in line with the ‘taxonomic yardstick’ in Afri­berina. Potentially Afriberina is a young group, the speciation is ongoing, and it would not be surprising if hybrids or intermediate forms will be found, particularly between A. salemae and A. tenietaria.

Text-fig. 182. Map showing the records of Afriberina tenietaria (Staudinger, 1900) and A. salemae Skou & Sihvonen sp. n. in southern Portugal and south-western Spain.

Sardocyrnia Wehrli, 1943 Sardocyrnia Wehrli, 1943, Gross-Schmett. Erde 4 (Suppl.): 523. Type species: Boarmia bastelicaria Bellier, 1862. There are no genus synonyms.

Diversity and distribution: Two species in south-western Europe. External characters and abdomen: Ground colour of forewings brownish grey, of hindwings more whitish grey with dark mottling. Postmedial line s-shaped, between costa and centre of wing weakly dentate. Subterminal line close to the postmedial line, thick, brown. Terminal line black, between veins slightly thickened. Wavy line whitish, often strongly waved. Discal spots present on all wings. Forewing medial line distal to discal spot, on hindwings crossing it. Antemedial and postbasal lines straight, close together, hardly visible on costa, on hindwings lacking. Forewing veins R1 and R2 separately arising from cell. R2 shortly anastomosing with R3 and R4. Thorax, frons and collar concolorous with ground colour, darker mottled. Fringes chequered light and dark grey, with a weak greyish line, parallel to termen. Abdomen dorsally with a double row of dark dots, first abdominal segment silver-grey. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs, distal spurs usually shorter. 395

Male genitalia: Uncus bottle-shaped, sclerotized, weakly striated, setose, Gnathos a ­ trophic. Valva divided. Dorsal arm narrow, setose, apex bifurcate with small teeth. Ventral arm wide, membranous, narrowing to apex, setose. Sacculus weakly sclerotized. Juxta constricted, anterior part wide, posteriorly furcated. Saccus wide, rounded. Aedeagus narrow, apex pointed, without cornutus. Female genitalia: Papillae anales soft, setose. Apophyses posteriores long, apophyses anteriores shorter, both thickened at 2/3 from the end, there spatulate. Lamella antevaginalis with two membranous, tapered ‘wings’, lamella postvaginalis a small sclerotized plate. Ductus bursae narrow. Posterior end of corpus bursae with small, narrow, sclerotized structure at junction with ductus seminalis. Corpus bursae long, membranous, folded, anterior part widened, oval, without signum. Remarks: Leraut (2009) downgrades Sardocyrnia from genus rank to synonymy of Peri­ batodes. The authors hesitate to follow this classification based on the comparatively large genetic distance (minimum distance of >6% in the COI gene) between Sardocyrnia and Peribatodes. Multi-gene analysis justifies classification of three different genera Sardocyrnia, Selidosema and Peribatodes clustering together in one common branch of genera (Murillo-­ Ramos et al. 2019).

135. Sardocyrnia bastelicaria (Bellier, 1862) Boarmia bastelicaria Bellier, 1862: Annls Soc. ent. Fr. (4) 2 (2): 380, pl. 9, fig. 3 (France: Cor­ sica, Bastelica). Syntype(s). Unavailable names (infrasubspecific): aestivalis: Bytinski-Salz (1937) (f.).

External characters and abdomen: Wingspan 28–35 mm, summer generation smaller, 24– 26 mm (Flamigni et al. 2007). Both Sardocyrnia species are very similar. Wing coloration, pattern and external characters see under genus description, for diagnostic features between the two species see below. Variation: Little in ground colour and distinctness of markings. Male and female genitalia: See genus description and under ‘similar species’. Distribution and abundance: Central-Mediterranian (Tyrrhenian). Endemic to Sardinia and Corsica. All records for Spain refer to S. fortunaria as a separate species (Skou 2007), but in Leraut (2009) recorded for Spain due to subordination under S. bastelicaria. – Rather local and usually found in single specimens or limited numbers. – Not recorded from outside Europe. Phenology: Bivoltine. From early May to late June and from early September to late October. Larva from October to April or May and again from June or July to August. Hibernation as larva (M. Leipnitz pers. comm.). Larvae continue to feed during winter unless it is unusually cold (T. Varenne pers. comm.). Adults active at night, attracted to light. Biology: Larva oligophagous. Found on Santolina corsica, Helichrysum italicum (T. Varenne pers. comm.) and Erica (M. Leipnitz pers. comm.). Reared on Erica carnea (M. Leipnitz pers. comm.). Habitat: Xerothermophilous. In various open habitats with scattered trees and bushes. From sea-level up to 1400 m.

396

Similar species: S. fortunaria differing from S. bastelicaria by the course of the postmedial line and differences in genitalia. See Text-figs 183–184. Male genitalia of S. bastelicaria with apical end of the dorsal arm of valva with shorter process and fewer teeth, these sometimes missing or shorter. Juxta arms distinct, wider and longer. Male genitalia of S. fortunaria: Apical end of the dorsal arm of valva with longer process, more and longer teeth. Juxta arms less distinct, smaller and shorter. Female genitalia without noticeable differences between the two species. Both Sardocyrnia species can be separated externally from the similar Menophra annegreteae Skou, 2007 by the costal part of the postmedial line (dentate in Sardo­cyrnia and acute-angled in M. annegreteae) and the male antennae (longer pectination in M. annegreteae). Genetic data: BIN: BOLD:AAI0724 (n=3 from France: Corsica; Italy: Sardinia). Genetically somewhat heterogeneous, populations from Corsica diverging from Sardinian populations by 0.8%. Nearest species: S. fortunaria (2.6%).

136. Sardocyrnia fortunaria (Vázquez, 1905) Boarmia fortunaria Vázquez, 1905: Boln Soc. esp. Hist. nat. 5 (2): 116, fig. 2a. (Spain). Syntype(s) including at least 1♀.

External characters and abdomen: Wingspan 28–35 mm, second and third generations smaller, 24–26 mm (Redondo pers comm.). Both Sardocyrnia species are very similar. Wing coloration, pattern and external characters see under genus description, for diagnostic features of the two species see Text-figs 183–184 and above under ‘Similar species’ of S. ­bastelicaria. Variation: Little in ground colour and distinctness of markings.

397

Male and female S. bastelicaria.

genitalia:

See genus description and above, under ‘Similar species’ of

Distribution and abundance: Endemic in eastern Spain. – Rather common, often found in numbers. – Not recorded from outside Europe.

Phenology: Bivoltine, at lower altitudes perhaps trivoltine. According to Redondo et al. (2009) in three generations: April to May, August to September and October to November. Own data from early April to late June, mid-July and mid-September to late October. Larval time apparently unknown, but most likely as in S. bastelicaria. Hibernation as larva (M. Leipnitz pers. comm.; rearing observation). Adults readily attracted to light. Biology: Natural foodplant of larva unknown (Redondo et al. 2009). Reared on Artemisia camphorata (M. Leipnitz pers. comm.). Pupation in a loose cocoon between plant litter on the ground (M. Leipnitz pers. comm.; rearing observation). Habitat: Xerothermophilous. In a variety of open habitats with sparse vegetation. From sea-level up to 1400 m, according to Redondo et al. (2009) up to 2000 m in the Sierra Nevada in southern Spain. Similar species: See above, under ‘Similar species’ of S. bastelicaria. Genetic data: BIN: BOLD:AAE4206 (n=7 from Spain). Genetically homogeneous. Nearest species: S. bastelicaria (2.6%), contradicting the downgrading of fortunaria to subsp. of S. bastelicaria in Leraut (2009). Remarks: Some authors (e.g. Leraut 2009) consider S. fortunaria as subspecies of S. bastelicaria. Because of the differences given above and the genetic data both taxa should better be treated as different species. 398

Text-figs 183–184. Diagnostic characters (indicated) of Sardocyrnia Wehrli, 1943 species. Textfig. 183. S. bastelicaria (Bellier, 1862), France, Corsica, Ghisoni, 1.vi.2008. Text-fig. 184. S. fortunaria (Vázquez, 1905), Spain, Province Teruel, Albarracín, 31.v.2007 (all coll. B. Müller).

Selidosema Hübner, 1823 Selidosema Hübner, [1823] 1816, Verz. bekannter Schmett.: 299. Type species: Geometra plumaria Denis & Schiffermüller, 1775. Genus synonymy after Scoble (1999). Spermo Gumppenberg, 1887, Nova Acta Acad. Caesar. Leop. Carol. 49: 334 (key). Type species: Geometra ambustaria Geyer, 1831.

Diversity and distribution: Eleven species in the central and western Palaearctic region (cf. Scoble & Hausmann 2007, updated). External characters and abdomen: Medium-sized, wingspan 25–43 mm, females with slightly narrower forewings. Ground colour brown to grey, more or less irrorated with dark scales. Medial, post- and antemedial lines black or dark grey, sometimes indistinct or faint in part. Terminal area in some species darker. Discal spots present. Antennae partially bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Uncus triangular, tip elongated, weakly setose. Gnathos usually absent, but present in S. erebaria casting doubt on current association with genus. Valva divided. Dorsal arm of valva slender, partly thickened and setose, with prominent structure. Ventral part 399

of valva nearly membranous, margin weakly setose. Saccus wide, rounded. Juxta large, distally with a wide structure of varying size. Aedeagus slender, vesica with more or less conspicuous cornuti or microcornuti. Female genitalia: Papillae anales long, weakly setose. Apophyses posteriores long, apophyses anteriores 50–85% length of the former, both ending spatulate. Lamella postvaginalis a conspicuous, sometimes trilobed sclerite. Corpus bursae membranous, anterior part in some species developed as a round sac. Remarks: In multi-gene analysis Sardocyrnia + Selidosema + Peribatodes cluster together (Murillo-Ramos et al. 2019).

137. Selidosema brunnearia (Villers, 1789) Phalaena Geometra brunnearia Villers, 1789: Linn. ent. 2: 325 (Europe). Syntype(s). Phalaena Geometra ericetaria Villers, 1789: Linn. ent. 11: 329, pl. 6, fig. 9 (Europe). Syntype(s). Phalaena subfimbriata Haworth, 1812: Trans. ent. Soc. Lond. 1 (3): 336 (United Kingdom). Syntype(s). Selidosema ericetaria var./ab. pallidaria Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (Edn 3) 1: 351 (Sicily; Dalmatia; centr. Italy; Pontus; Hyrcania). Syntype(s). Downgraded from subspecific rank (Scoble 1999) to synonymy by Flamigni et al. (2016). This taxon needs a lectotype designation, preferably from Sicily or Dalmatia. Selidosema ericetaria var. scandinaviaria Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (Edn 3) 1: 351 (Southern Scandinavia; Denmark; Gotlandia). Syntype(s). Valid at subspecific rank (Scoble 1999). Selidosema ericetaria var. syriacaria Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. (Edn 3) 1: 351 (Turkey: ‘Syria; Pontus (trans.)’). Syntype(s). Valid at subspecific rank (Scoble 1999). Conspecifity with S. brunnearia questionable, see remarks under ‘Distribution’ and ‘Genetic data’. This taxon needs a lectotype designation, preferably from the southern Turkish border towards Syria (from where Staudinger likely got his material) and not from Pontus from where Staudinger characterizes the populations as ‘transitional’. Selidosema ericetaria oelandica Wahlgren, 1913: Ent. Tidskr. 34 (2–4): 165, fig. 2 (Sweden: Öland Island, Alvar region). Syntype(s) ♂. Selidosema brunnearia dilucescens Wehrli, 1924: Mitt. schweiz. ent. Ges. 13 (7): 345 (Switzerland: Wallis, Zermatt). Syntypes 9 (ZFMK). Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Selidosema tyronensis Cockayne, 1948: Entomologist’s Rec. J. Var. 60: 79 (Northern Ireland: County Tyrone, Dylonghan, near Lough Neagh). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999). Selidosema parenzani Hausmann, 1993: Naturalista sicil. (4) 17 (1–2): 94, figs 3, 4, 15 (Italy: ­Sicily, Madonie, 1200 m). Holotype ♂. (ZSM, dissected, examined and DNA barcoded). Junior synonym (hybrid), see remarks under ‘Genetic data’. Unavailable names (infrasubspecific): cleui: Leraut (2009) (f.); demarginata: Lempke (1952) (ab.); depuncta: Lempke (1952) (ab.); deumbrata: Lempke (1952) (ab.); obscura: Lempke (1970) (f.); oelandica Wahlgren, 1913 (ab.).

External characters and abdomen: Wingspan ♂ 27–43 mm, females on average slightly smaller. Ground colour pale brown to bluish grey, slightly irrorated with brown or grey. Postmedial and antemedial lines on forewing weak, bent, often angled near costa, at costa stronger, on hindwing usually indistinct or absent. Medial line varying in intensity, usually running through the discal spot, more rarely next to it. Terminal area almost always darker. 400

Discal spot on forewing round or elliptic, on hindwing developed as a weaker dot. Fringes unicolorous dark brown or grey. Frons brownish, smoothly scaled. Vertex, collar, tegula, thorax and abdomen grey, consisting of a mixture of brown, beige and pale grey scales. Antennae partially bipectinate in male (about 4–5 flagellomeres free of branches), branches long. Female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Variation: Highly variable. Ground colour varying from pale brown to beige or bluish grey (violet-grey). Post- and antemedial lines (on forewing only) sometimes obsolete except dark spots on costa. Medial line thicker or thinner, usually running through the discal spot or rarely distal or proximal to it. Terminal area usually darker than the rest of the wings, but rarely concolorous with the rest of the wings. Extreme forms unicolorous brown or beige with discal spots present. Because of the high variability a big number of forms and subspecies have been described (Wehrli 1939–1954; Leraut 2008). Subsp. scandinaviaria from the northern half of Europe including the British Isles with violet-­grey ground colour. Forewing transverse lines more conspicuous, terminal area usually darkened on both wings. Subsp. tyronensis has been described from a bog in the county Tyrone (Northern Ireland) as relatively small (wingspan 30–34 mm), dark marginal shade narrower, dark median shade much fainter (Skinner 2009). Typical characteristics of some taxa earlier validated as subspecies are often found in other regions. Violet-grey specimens are also found in France (Leraut 2008) and Switzerland. Rezbanyai (1981) writes of forms that are ‘locally frequent or predominant within the species’ but he also supposed the existence of mixed populations of ‘prehistoric’ subspecies. Male genitalia: Uncus sclerotized, wide, weakly setose, triangular at base, distally tapering to a rounded tip. Valva partly divided, both parts close together. Dorsal part of valva sclerotized, apex round, setose, with inwardly directed ridge. Apical part of the ridge convex, shallowly curved, margin strongly spinose, in the centre a group of short spines. Basal part of the ridge with a row of spines at the centre (see S. plumaria, S. pyrenaearia). Ventral part of valva basally weakly sclerotized, apical membranous, margin setose. At centre weakly constricted (in S. plumaria and S. pyrenaearia more strongly constricted). Saccus wide, in the centre usually flattened. Juxta a wide plate, lateral margins more or less constricted, apex bilobed. Aedeagus narrow, straight, distal end oblique, flattened, tapered with small pointed process (in S. plumaria rounded without pointed process, in S. pyrenaearia tapered with small pointed process, the latter larger than equivalent in S. brunnearia). For male genitalia of the hybrid ‘parenzani’ see under S. ambustaria. Female genitalia: Papillae anales long, weakly setose. Apophyses posteriores very long (on average 2.65 mm), straight, apophyses anteriores about half length of the former, both ending spatulate. Lamella postvaginalis a sclerotized, conspicuous, trilobed structure, i.e. distally ending in two lateral and one medial projection (tongue). Lateral projections usually narrow, shorter than the medial one, more pointed whilst the medial projection is rounded. Proximal margin straight with rounded projection, usually not jagged (see S. plumaria, S. pyrenaearia), but variable, not always suitable as differential feature for separating species. Anterior end of ductus bursae with distinct lateral projection. Posterior part of corpus bursae long, slender, membranous, striated. Anterior part globular, with signum. Signum sub-quadrate, with diagonal sclerotized ridge (length about 0.3 mm). For female genitalia of the hybrid ‘parenzani’ see under S. ambustaria. 401

Distribution and abundance: European. From Estonia across southern Scandinavia to the British Isles, in the south to northern France, southern Germany and Austria (subsp. scandinaviaria). Nominotypical subspecies in Italy including Sicily, southern Switzerland, south-eastern France and in the Pyrenees. From south-western Finland records of three (supposedly) migrating specimens, the first of them originally reported as Selidosema plumaria (Sihvonen & Mikkola 2002). – Local and not common, now and then found in numbers in its localities. – Outside Europe reported from Turkey to Israel and Jordan (subsp. syriacaria) (Staudinger 1901; Hausmann 1991), but it is doubtful if these records really are S. brunnearia or rather belong to S. plumaria or to a separate species (see genetic data). Also reported from Morocco (Rungs 1981), but after the elevation of S. pyrenaearia to species rank and considering that S. brunnearia does not occur outside the Pyrenees in the Iberian peninsula, the Moroccan records await re-examination and verification (see discussion of taxon mauretania Wehrli, 1953 under S. pyrenaearia).

Phenology: Univoltine. In southern Fennoscandia from late July to late August, in south-­ western Germany from early August to mid-September (Ebert 2003), in Italy from early August until October (Flamigni et al. 2016). Fully grown larvae in north-western Germany in mid- and late May (Wegner 2013). Hibernation as larva. Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous, with slight preference for Fabaceae. Found on Cytisus ­scoparius, Genista anglica and young leaves of Salix aurita in north-western Germany (Wegner 2013), Calluna vulgaris and Lotus corniculatus in Great Britain (Porter 2010), on Sanguisorba ­minor, Lembotropis nigricans, Coronilla varia and Galium mollugo in south-western Germany (Ebert 2003). Recorded on Genista, Vicia, Ononis, Trifolium, Dorycnium, Lotus corniculatus, Hippocrepis and Onobrychis (Flamigni et al. 2016). 402

Habitat: Xerothermophilous. In southern Fennoscandia in heaths and overgrown costal dunes, in Sweden in the so-called ‘alvar’, in north-western Germany also in warm places with mature plants of Cytisus scoparius (Wegner 2013) in southern Europe in a larger ­variety of open habitats. From sea-level up to 1900 m in the Italian Alps (Flamigni et al. 2016). Similar species: This species and the following two (S. plumaria and S. pyrenaearia) are very similar and cannot be separated with certainty without dissection of male genitalia. Identification of females based on genitalia not always reliable. Genetic data: BINs: BOLD:ACE5955 (n=22 from Sweden, Estonia, Denmark, Germany, France, Italy including Sicily); BOLD:AAC3624 (n=3 from Israel). Genetically homo­geneous within the BINs, but populations from southern Italy and Sicily diverging by 0.8%. The cluster from Israel diverging by 1.9% (subsp. syriacaria). Nearest species: S. plumaria (1.4%). The (NGS based) DNA analysis of the holotype of the taxon ‘parenzani’ (see nomenclature above) resulted in a 100% match with S. brunnearia suggesting it to be a hybrid S. ambus­ taria (male) x S. brunnearia (female), the DNA barcode of a paratype resulted in the opposite, i.e. a 100% match with S. ambustaria suggesting it to be the product of hybridization between S. brunnearia (male) x S. ambustaria (female), considering the maternally inherited mitochondrial COI gene. Remarks: Wegner (2013) deals in details with the biology and habitats of S. brunnearia in north-western Germany. The taxonomy and other information about the species pair S. brunnearia and S. plumaria has been confusing until Krampl & Marek (1981) first managed to show the differences of these taxa including their correct names. In the past some entomologists have published detailed results concerning the species-group S. brunnearia, S. plumaria and later also S. pyrenaeata (Fletcher 1949; Urbahn & Urbahn 1978; Rezbanyai 1981; 1983; Krampl & Marek 1981; Sihvonen & Mikkola 2002; Redondo & Gaston 2012).

138. Selidosema plumaria (Denis & Schiffermüller, 1775) Geometra plumaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 106 (Austria: Vienna district). Syntype(s) ♂. Phalaena limbata Rambur, 1829: Annls Sci. Observ. 2: 267, pl. 6, fig. 8 (central France: near Ganges). Holotype ♀. Selidosema plumaria amani Wehrli, 1932: Mitt. münch. ent. Ges. 22 (1): 10 (Turkey: Amanus, Düldül Dagh). Syntypes 5♂. Valid at subspecific rank (Scoble 1999).

External characters and abdomen: Generally as S. brunnearia (see above), but usually differing by more brownish ground colour, rarely with bluish grey forms (locally occurring), medial line usually running proximal to discal spot, straight, slightly angled near costa. Discal spot on forewing elliptic, sometimes lunular. Note: Wing pattern not always reliable for correct determination. Variation: Similar variation as described for S. brunnearia. Male genitalia: Generally as in S. brunnearia and S. pyrenaearia. Valva partly divided, both parts close together. Dorsal part of valva sclerotized, apex round, setose, with inwardly directed ridge. Whole ridge uniformly and more strongly curved, therefore wider than in S. brunnearia, margin strongly spinose, in the centre a group of short spines (see S. brunnearia and S. pyrenaearia). Ventral part of valva basally weakly sclerotized, apically membranous, margin setose. In the centre more strongly constricted (in S. brunnearia less). Juxta a wide plate, lateral margins slightly constricted, apex bilobed, hardly different from juxta 403

of S. brunnearia and S. pyrenaearia. Aedeagus narrow to apex, straight, distal end round (in S. brunnearia oblique, flattened, tapered with small pointed process, in S. pyrenaearia tapered with larger pointed process). Female genitalia: Generally as in S. brunnearia and S. pyrenearia. Anterior end of ductus bursae slightly expanded laterally (Sihvonen & Mikkola 2002). Lamella postvaginalis different from S. brunnearia and S. pyrenaearia, also trilobed, apically all projections rounded. Lateral projections as in S. pyrenaearia, wider than in S. brunnearia, of nearly the same length than the medial one, tip more rounded. Medial tongue rounded. Proximal margin straight with rounded projection, usually not jagged (see S. plumaria, S. pyrenaearia). Note: Female genital features not always suitable for species separation. Distribution and abundance: European-West Asiatic. From southern half of Russia across the Balkan peninsula to southern central Europe, north-eastern Italy and Spain. Erroneously recorded for Finland and Portugal by Leraut (2009), moreover strongly exaggerating the distribution in Italy. – Local, sometimes found in numbers. – Outside Europe (in central and southern Turkey subsp. amani), north-westernmost Syria (DNA barcoded), Georgia, Armenia and Azerbaijan.

Phenology: Univoltine. From early August to late September or early October. Medium-sized larvae in late April with pupation in mid-May (Y. Budashkin, V. Savchuk pers. comm.). Hibernation as larva. Adults active at night, both sexes readily attracted to light. Biology: Larva polyphagous. Found on Asphodeline taurica (Y. Budashkin, V. Savchuk pers. comm.). Recorded on Genista, Lathyrus, Lotus, Onobrychis, Prunus spinosa, Erica, Calluna vulgaris and Artemisia (Flamigni et al. 2016).

404

Habitat: Xerothermophilous. In a large variety of warm, open habitats. From close to sea-level up to 1600 m. Similar species: S. brunnearia and S. pyrenaearia (see there). Genetic data: BINs: BOLD:ACE9860 (n=8 from Greece, Turkey); BOLD:ACE9862 (n=13 from Austria, Spain, Italy, Hungary, Croatia, Turkey and north-western Syria). Both BINs diverging by 1.2%. Divergence requiring further comprehensive, integrative study combining dissections with DNA barcodes. Nearest species: S. brunnearia (1.4%). Remarks: See under S. brunnearia.

139. Selidosema pyrenaearia (Boisduval, 1840) Fidonia pyrenaearia Boisduval, 1840: Genera Index meth. eur. Lepid.: 191 (Pyrenees). Syntype(s). Earlier validated as subspecies of S. brunnearia (Scoble 1999), raised to specific rank by Redondo & Gastón (2012). Selidosema oliveirata Mabille, 1876: Bull. Soc. ent. Fr. 1876: 109 (Portugal, Coimbra). Lectotype ♂ (ZSM/Herbulot). Validated at subspecific rank by Redondo & Gastón (2012), herewith downgraded from subspecies rank to synonymy (see below). Selidosema plumaria mauretania Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 653 (Algeria: Yakouren). Holotype ♂ (ZFMK). Until recently (Rungs 1981; Scoble 1999) retained conspecific with S. brunnearia (in Leraut 2009 as subsp.) but more likely to be attributed to this species, possibly at subspecies rank, see remarks under ‘Distribution’ of S. brunnearia. Requiring further study. Selidosema herbuloti Rungs, 1975: Alexanor 9 (2): 68, figs 1, 2 (Morocco, eastern Middle Atlas: Ain Mellal). Holotype ♂ (MNHN). Validated at species rank by Scoble (1999), downgraded to synonymy of mauretania by Leraut (2009).

External characters and abdomen: Generally as in S. brunnearia and S. plumaria, but with the following differences: Ground colour usually reddish brown to ash grey, distinctly darker. Medial line, if present, usually running through the discal spot, obtusely angled below it or more curved than in S. brunnearia. Discal spot on the forewing elongate-linear. Note: These characteristics are not always reliable for a correct determination. Variation: Variable. Ground colour varying from reddish brown to brownish grey or ash grey, rarely basal and medial areas paler. Medial area can be darkened to blackish brown. Postmedial, medial and antemedial lines of forewing can be absent except dark spots at the costa. Populations from western Iberian peninsula (f. ‘oliveirata’) with predominantly ash grey ground colour, often strongly irrorated with dark grey (Mabille 1876; Redondo et al. 2012), here classified as infrasubspecific, considering the huge variation of S. pyrenaearia in Portugal (M. Corley pers. comm.) and the clinal character transitions. Male genitalia: Generally as in S. brunnearia and S. plumaria. Valva partly divided, both parts close together. Dorsal part of valva sclerotized, apex round, partly setose, with inwardly directed ridge. Apical part of the ridge concave, basal part convex, whole margin strongly spinose but without spines in the centre (see S. brunnearia and S. plumaria). Ventral part of valva basally weakly sclerotized, apically membranous, margin setose. At centre more strongly constricted (in S. brunnearia less). Juxta a wide plate with slightly constricted lateral margins, apex hardly bilobed. Aedeagus straight, distal end with pointed process, larger than in S. brunnearia. Distal end in S. plumaria rounded, in S. brunnnearia oblique, flattened, tapered with small process. 405

Female genitalia: Generally as in S. brunnearia and S. plumaria. Characterizised by the presence of sclerotized lateral folds at the colliculum. Lamella postvaginalis trilobed as in the other two species, distally as in S. plumaria, central ‘tongue’ wider than in S. brunnearia. Proximal margin uneven in a semicircle, usually strongly jagged (see S. brunnearia and S. plumaria). Signum on average slightly larger. Note: Female genitalia features not always suitable for species separation. Distribution and abundance: Verified distribution restricted to the Iberian peninsula (cf. R ­ edondo & Gastón 2012). – Relatively common, usually occurring in limited numbers. – So far not reported from outside Europe, but Moroccan and Algerian records of S. brunnearia may very well prove to be conspecific with S.  pyrenaearia (see nomenclatural remarks above, to mauretania and herbuloti).

Phenology: Univoltine. From early August to late September with single specimens in early October (Redondo & Gaston 2012). Single specimens in June and July perhaps belonging to an additional partial early summer generation (M. Corley pers. comm.; cf. Redondo & Gaston 2012). Adults active at night, readily attracted to light. Biology: Larva oligo- or polyphagous (M. Corley pers. comm.). Found on Lavandula stoechas (Corley 2015). Habitat: Occurring in a variety of open habitats, almost always on acid soils (M. Corley pers. comm.), exceptionally also in woodland. Often in mountain areas up to 1500 m, exceptionally higher (1950 m in Sierra de Durcal, Granada province), sometimes down to sea-level. Similar species: S. brunnearia and S. plumaria (see there).

406

Genetic data: BIN: BOLD:ABZ3718 (n=2 from Portugal). Genetically homogeneous. Nearest species: S. plumaria from Spain (2.2%). Remarks: Recently raised to species rank (Redondo & Gaston 2012), still with limited know­ ledge.

140. Selidosema taeniolaria (Hübner, 1813) Geometra taeniolaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 69, fig. 357 (Europe). Syntype(s). Selidosema taeniolaria extensaria Turati, 1919: Naturalista sicil. 23 (7–12): 323, figs 3, 4 (Italy: Liguria, Pigna (Ventimiglia)). Syntypes 6♂1♀. Validated at subspecific rank by Scoble (1999), herewith downgraded to synonymy based on homogeneity of DNA barcodes, clinal character transitions and absence of constant and significant differential features. Unavailable names (infrasubspecific): unicolor: Leraut (2009) (f.).

External characters and abdomen: Wingspan 25–38 mm. Ground colour brown or greyish, strongly covered with dark grey streaks. All transverse lines of forewing black (if present). Transverse lines of hindwing usually indistinct. Postmedial line angled near costa, between veins CuA2 and A concave. Antemedial line bent, serrate, absent from hindwing. Medial line often reduced or absent. In males medial area sometimes darker. All wings usually with subterminal line edged white, terminal line a row of small black crescent spots. Fringes chequered brown and dark grey, divided by a dark fine line. Discal spots present, sometimes weak, on forewings elongate-linear, on underside more distinct. Frons, vertex, collar, tegula, thorax and abdomen grey, consisting of a mixture of dark and pale grey scales. Abdomen often paler. Antennae partially bipectinate in male (about 5–6 flagellomeres free of branches), branches long. Female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Variation: Variable. Wings can be darker or paler because of varying density of the dark ­ ottling, all lines distinct or reduced. The light shadow of the subterminal line can be dism tinct or reduced, sometimes interrupted. Also strongly varying in size. Specimens from north-western Italy and French Alpes Maritimes (‘f. extensaria’) somewhat larger, on average lighter, but variable individually. Male genitalia: Uncus sclerotized, base triangular, apex beak-shaped, tapered, weakly setose. Valva divided. Dorsal arm of valva slender, bent, apex thickened, setose. On the basal third an inwardly directed, strongly sclerotized, tapered thorn with bent tip. Ventral part of valva nearly membranous, margin weakly setose, in the centre obtusely angled. Saccus wide, rounded. Juxta large, concave between lateral margins and apex. Aedeagus narrow, arcuate, apex tapered. Vesica with microcornuti. Female genitalia: Papillae anales long, weakly setose. Apophyses posteriores very long (on average 3.2 mm), straight, apophyses anteriores about 60% length of the former, spatulate at tip. Lamella postvaginalis a folded plate, distally rounded and more strongly sclerotized. Ductus bursae short, sclerotized. Posterior part of corpus bursae cylindrical, sclerotized, striated. Anterior part larger, round, with signum. Signum triangular with concave margins. Distribution and abundance: (South-west) European. From south-western Switzerland and north-westernmost Italy across southern France to the Iberian peninsula. Distribution in central and southern Italy misleadingly suggested by the map in Leraut (2009). – Rather common, often found in numbers. – Unknown from outside Europe.

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Phenology: Univoltine. From late July to early or mid-October. In southern Spain until Decem­ ber (P. Hale pers. comm.). Larva from mid-April to early June (D. Fritsch pers comm.). Overwintering as small caterpillar (T. Varenne, M. Leipnitz pers. comm.). Adults active at night, readily attracted to light. Biology: Larva polyphagous. Found and reared on Cytisus (P. Skou own data), Ulex, Genista, Artemisia herba-alba, Thymus and Ribes uva-crispa (D.  Fritsch pers. comm.). Recorded on Juniperus, Ephedra, Berberis, Prunus spinosa, P. dulcis, Spartium, Retama, Dorycnium, Buxus, Cistus, Erica, Calluna and Fraxinus (Flamigni et al. 2016). Pupation in a loose cocoon on the ground (T. Varenne pers. comm.). Habitat: Xerothermophilous. In a rather large variety of open scrub habitats, sometimes also in woodland. From sea-level up to 1700 m, in Sierra Navada in southern Spain up to 1900 m. Similar species: None in Europe. S. atraria Powell, 1941 from Morocco very similar, discrimination possible by genitalia dissection. Genetic data: BIN: BOLD:ABA8420 (n=3 from France, Spain). Genetically homogeneous. Nearest species: S. atraria Powell, 1941 from Morocco (2.6%), S. combustaria from Israel (2.7%), S. plumaria (2.8%).

141. Selidosema ambustaria (Geyer, 1831) Geometra ambustaria Geyer, 1831: Samml. Eur. Schmett. 5 Geometrae (1): pl. 108, figs 567, 568 (Italy: Sicily). Syntype(s). Fidonia duponcheliaria Lefèbvre, 1831: Magasin Zool. Paris 1 (2): (Classe IX: Insectes): 32, figured (as Fidonia duponchelii) (Italy: Sicily, Sperlinga). Syntype(s) ♂♀. ‡ Selidosema parenzani: Hausmann, 1993 (partim): Paratype ♂. (ZSM, examined and DNA barcoded). The DNA analysis of the paratype resulted in a 100% match with S. ambustaria

408

suggesting it to be the descendant of a hybrid mating S. brunnearia (male) x S. ambustaria (female), for the holotype see under S. brunnearia. For external characters of the hybrids see under ‘variation’.

External characters and abdomen: Wingspan 27–33 mm. Males: Ground colour beige, covered with streaks, basal and terminal areas of forewing often brown. All transverse lines distinct, blackish, often diffuse. Subterminal line edged whitish. Terminal line a row of small black crescent spots. Fringes brown, slightly chequered, divided by a darker fine line. Hindwings with pale grey lines. Antennae partially bipectinate with long branches, 7–8 apical flagellomeres free of branches. Females: Wings narrower. Ground colour dark grey to beige, transverse lines as in males. Antennae filiform. In both sexes discal spot of forewing large, black, sometimes with a small light centre, on hindwings smaller, greyish brown. Frons, vertex, collar, tegula, thorax and abdomen beige in males, dark grey in females. Hindtibia of both sexes with 2+2 spurs. Variation: Varying in the density of streaks on the wings, in the distinctness of transverse lines and in the extent of darker areas. The hybrid between S. ambustaria and S. brunnearia (named as ‘S. parenzani’) very similar to that of S. ambustaria but with paler ground colour, terminal area darker brown, contrasted, forewing discal spot less developed and connected with costa by a dark streak, postmedial line absent from hindwing (present in S. ambus­ taria), cf. Hausmann (1993a) and Flamigni et al. (2016). For synonymization of this taxon and recognition as hybrid see above, under S. brunnearia. Male genitalia: Uncus wide, apex tapered. Valva divided. Dorsal arm slender, straight, basal half with tapered sclerotized thorn, inwardly directed, shorter than in S. taeniolaria. Apical half oval widened, densely setose. Ventral part basally weakly sclerotized, apically membranous. Constricted in the middle. Margin setose. Saccus wide, round. Juxta large, ending in a structure like a mushroom cap. Aedeagus straight, apex with conspicuous, sclerotized tapered process. Vesica with a group of about 10 spines of different length. Male genitalia of the hybrid between S. ambustaria and S. brunnearia (‘parenzani’) with rather long uncus, apical half of dorsal arm of valva narrow, not oval, densely setose, inwardly directed spine distinctly shorter than that of S. ambustaria; distal process of aedeagus about half length of the equivalent in S. ambustaria, vesica with less and shorter cornuti (see Hausmann 1993a). Female genitalia: Papillae anales longish, weakly setose. Apophyses posteriores long (on average 2.0 mm), straight. Apophyses anteriores less short, about 85% of length of the previous, both ending spatulate, shorter than in females of brunnearia group. Lamella postvaginalis distally rounded and strongly sclerotized (see Hausmann 1993a). Lamella ante­ vaginalis alate. Antrum sclerotized with longitudinally arranged sclerotized slats. Ductus bursae wide, short. Corpus bursae membranous, wide, elongated with signum. Signum triangular, smaller than in brunnearia group (0.20–0.25 mm). Female genitalia of the hybrid between S. ambustaria and S. brunnearia (‘parenzani’) with longer apophyses posteriores (on average 2.2 mm), wider ductus bursae, longer antrum, signum in corpus bursae wider, about 0.4 mm (see Hausmann 1993a). Distribution and abundance: Central Mediterranean (Italian endemic). In Europe only in southern Italy (Calabria) and in Sicily. Distribution in central and northern Italy erroneously suggested by the map in Leraut (2009). – Rather local, usually found in limited numbers. – Unknown from outside Europe. Reputedly in Morocco (Rungs 1981; without locality), likely misidentified.

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Phenology: Univoltine. From late August to late October, hybrids S. ambustaria x S. brunnearia (‘parenzani’) flying rather early, collected in mid-August. Adults active at night, both sexes attracted to light. Biology: Larva monophagous. Recorded on Hypericum, especially H. perforatum. Reared on Artemisia (Flamigni et al. 2016). Habitat: Silvicolous. In open woodland from sea-level up to 1100 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAM0203 (n=3 from Italy: Sicily). Genetically homogeneous. Nearest species: S. combustaria from Israel (1.2%; BOLD:ACT3583), S. picturata from Tuni­sia (1.9%; BOLD:ABY8808). Both aforementioned species in wing pattern (and genitalia?) reminiscent of African members of the genus Xylopteryx, the latter perhaps requiring taxonomic revision at genus level. For the genetic identity of the holotype and a paratype of the taxon ‘parenzani’ (hybrid between S. ambustaria and S. brunnearia) see under genetic data of S. brunnearia. Remarks: See differences in genitalia structures in Hausmann (1993a). In the Madonie mountains of Sicily occasionally hybridizing with S. brunnearia, see under genetic data of S. brunnearia.

142. Selidosema erebaria Oberthür, 1883 Selidosema erebaria Oberthür, 1883: Bull. Soc. ent. Fr. 1883: 49 (western Algeria: Sebdou). Syntype(s). Calamodes postdentaria Rothschild, 1914: Novit. zool. 21: 350 (central Algeria: Guelt-Es-Stel). Syntypes 3♂ (NHMUK). Valid at subspecific rank (Scoble 1999).

410

External characters and abdomen: Wingspan 28–36 mm, hindwing termen undulating, distinctly arched between veins. Males: Ground colour dark grey, heavily irrorated with grey or blackish. Postmedial line distinct, black, on forewing distally often edged whitish grey, proximally edged dark, double-angled near costa, concave near inner margin, on hindwing evenly bent. Antemedial line on forewing bent, often edged whitish grey. Other lines in­distinct or absent. Ground colour of females paler grey, wings irrorated with grey or brownish. Postmedial and antemedial line usually black, bent as in males, on hindwings paler. Medial line usually distinct, more diffuse. In both sexes terminal line distinct, black, fringes grey. Antennae bipectinate in male, about 10 apical flagellomeres free of branches, filiform in female. Frons smoothly scaled. Frons, vertex, collar, tegula, thorax and abdomen concolorous with ground colour. Hindtibia of both sexes with 2+2 spurs. Variation: Variable. Ground colour darker or lighter, being more or less irrorated with grey or black. On forewing proximal dark edging of postmedial line as well as distal whitish edging on both wings can be more distinct or sometimes absent. Subsp. postdentaria with rusty brown-grey ground colour, irrorated with darker scales. Frons, vertex, collar, tegula, thorax and abdomen concolorous (Rothschild 1914). Male genitalia: Uncus wide, triangular, apex rounded with small bent tip. Gnathos present, arms fused ventrally, ring-shaped, ventral element prominent. Valva wide. Costa sclerotized, apical half setose, apex round. Ventral arm of valva wide, sclerotized over half valva length, widely ending in a concave structure with conspicuous group of setae. Saccus wide, round. Juxta weakly sclerotized, rounded, posteriorly with narrow process. Aedeagus short, nearly cylindrical, apex tapered, caecum rounded, without cornutus. Female genitalia: Papillae anales elongated, narrow, weakly setose. Apophyses posteriores long (on average 2.1 mm), straight, widely spatulate at tip. Apophyses anteriores shorter, about 60% of length of the former. Lamella antevaginalis a flat, curved band, lamella postvaginalis an almost rectangular folded plate. Ductus bursae wide, short, sclerotized. Corpus bursae membranous, posterior part elongated, slightly striated, anterior part larger, pearshaped. Signum small, hardly visible, nearly circular. Distribution and abundance: Maghrebian. In Europe only known from the Italian island of Pantelleria (Fischer 2009). – Rather common in Pantelleria. – Outside Europe restricted to North Africa: Nominotypical subspecies from western Algeria to Morocco. In the north of central Algeria and in Pantelleria replaced by subsp. postdentaria. Phenology: Probably univoltine. In Pantelleria in early and mid-October (Fischer 2009). In Morocco recorded from July to September (Rungs 1981). Larval time as well as hibernation stage unknown. Adults active at night, attracted to light. Biology: Larva polyphagous. In Morocco recorded on Pistacia lentiscus, Arbutus unedo and Juniperus oxycedrus (Rungs 1981). Habitat: In Pantelleria in the evergreen forest of ‘Montagna Grande’ where the species was found at 650 m, but not at lower altitudes (Fischer 2009). Genetic data: BIN: BOLD:AAH9876 (n=4 from Italy: Pantelleria and Morocco: Middle Atlas). Pantellerian and Moroccan populations diverging by 1.4% (without differences in wing pattern). Genetically rather isolated and not clustering closely with any other Palaearctic congener. Nearest species: S. brunnearia from Spain (7.2%), but much closer to an unidentified Lassaba from China (5.1%) and Lassaba acribomena (Prout, 1928) from Malaysia and Sumatra (5.5%). 411

Remarks: Earlier, S. erebaria was combined with Menophra (Scoble 1999) or Peribatodes (Leraut 2009). Based on the morphology of genitalia, Fischer (2009) suggests that S. erebaria belongs to the genus Selidosema. Although genetic data confirm a closer relationship to Selidosema than to Peribatodes and Menophra (see above, under genetic data), the generic combination requires further integrative study. Moreover there are significant differences in male genitalia: Gnathos present, ventral part of valva with transverse ridge etc. Hence, the combination of this species name with genus Selidosema is still tentative and requires further studies.

Peribatodes Wehrli, 1943 Peribatodes Wehrli, 1943, (originally described as a subgenus of Boarmia), in Seitz, GrossSchmett. Erde 4 (Suppl.): 478. Type species: Peribatodes rhomboidaria (Denis & Schiffermüller, 1775).

Diversity and distribution: Genus with 11 species, restricted to the western and central Palaearctic region from western Europe to north-western India, and to North Africa. In ­Europe, most species of this genus occurring in central and southern Europe. External characters and abdomen: Wingspan between 26–48 mm (autumn generation slightly smaller; females of the same generation are slightly larger than males). Wings ground colour varying from yellow to dark brown and grey, hindwings slightly lighter. Markings wavy, varying from dark brown to black. Basal line absent from both wings. Antemedial line curved, angled or straight on forewings, present as a faint line or absent from hindwing. Forewing medial and postmedial lines getting closer, or anastomosing towards dorsum. Postmedial line dentate or smooth. Subterminal line absent from upperside, or not visible (but present in P. secundaria and P. powelli groups). Wavy line dentate to irregularly wavy, with white shadow, sometimes partially faint. Hindwing medial line usually present (faint or 412

absent in P. ilicaria); postmedial line curved, dentated. Wavy line usually invisible or faint (but present on the wing underside of the secundaria group). Discal spot in most species present on both wings, sometimes hardly visible on hindwings (absent from hindwings in P. powelli). Terminal line mostly discontinuous (except in the powelli group). Fringes darker at vein endings. Underside slightly lighter than upperside, markings paler; a pale blotch present on underside of forewing terminal area, and additional pale area on forewing apex. Antennae bipectinate in male, filiform in female. Male genitalia: Uncus triangular, beak-shaped, apically curved, acute, basally wide, in most species dorsally extended, setose. Socii absent. Gnathos reduced or absent. Valva basally broad, narrowing towards apex; one or more rows of long and stout setae present on termen and ventral part of valva. Costal part of valva strongly sclerotized, distally separated from apex of valva, apically setose; costa subapically triangularly extended. Editum (cf. Steinmann & Zombori 1985 = subcostal plica of valva in Beljaev 2008) reduced, setose, present at the base of the costa of valva. Juxta weekly sclerotized, elongated. Saccus widely rounded. Aedeagus straight, vesica either without cornuti or with one or two cluster(s) of short spines. Female genitalia: Papillae anales ovally elongated, setose. Apophyses thin, at the end flattened. Lamella postvaginalis wide, slightly sclerotized; lamella antevaginalis well sclerotized, diagnostic for species. Ductus bursae short, well sclerotized. Corpus bursae long, membranous. Signum strongly sclerotized as a transverse fold. Systematics: Albers (1940) regarded the species of this genus as belonging to two species-groups, based on the analysis of male genitalia, and this was followed by Flamigni et al. (2016). Albers & Warnecke (1941) excluded P. powelli (Oberthür, 1913) from the secundaria group, which was confirmed by Wehrli (1943). Flamigni et al. (2016) regarded P. ilicaria (Geyer, 1833) as a separate species-group. Following this system, we validate two additional species-groups (powelli and ilicaria). Diagnostic characters of these four Peri­ batodes species-groups are: rhomboidaria species-group (including: P. rhomboidaria; P. perversaria; P. omalensis sp. n.; P. ebusaria and P. umbraria): –– Male antenna with gradually reduced size of branches towards tip (Text-fig. 197), exception: P. ebusaria. Branches medium-sized (longest branches 1.3 mm) –– Forewing medial areas lighter than basal and terminal area –– Forewing postmedial line inclined inwards near costa and not curved near dorsum (Textfig. 185) –– Hindwing medial line present (Text-fig. 185) –– Terminal line of all wings blackish, narrow, discontinuous (Text-fig. 185) –– In male genitalia, uncus dorsally extremely extended (Text-figs 189–192); costa of valva covered with tiny spines up to the base of its triangular extension; valva without ventral tubercles; aedeagus as long as, or longer than length of valva; vesica with two separated clusters of short spines (exception: P. umbraria with one cluster) secundaria species-group (including: P. secundaria and P. correptaria): –– Male antenna with abruptly reduced size of branches towards tip (Text-fig. 196). Branches large-sized (longest branches 1.7 mm) –– Forewing medial areas lighter than basal and terminal area –– Forewing postmedial line not inclined, deeply curved near dorsum (Text-fig. 186) –– Hindwing medial line present (Text-fig. 186) –– Terminal line of all wings blackish, narrow, discontinuous (Text-fig. 186) 413

–– In male genitalia, uncus dorsally slightly swollen (Text-fig. 193); costa of valva covered with tiny spines on its triangular extension and further towards base of valva; valva with ventral tubercles; aedeagus shorter than length of valva; vesica with one cluster of long spines powelli species-group (including: P. powelli and P. aragonis) –– Male antenna with gradually reduced size of branches towards tip (Text-fig. 197). Branches medium-sized (longest branches 1.2 mm) –– Forewing medial areas lighter than basal and terminal area –– Forewing antemedial line deeply angled outwards; postmedial line deeply curved inwards and angled outwards, then continuing straight to dorsum (Text-fig. 187) –– Hindwing medial line present (Text-fig. 187) –– Terminal line of all wings blackish, narrow, discontinuous (Text-fig. 187) –– In male genitalia uncus short, curved, dorsally not extended (Text-fig. 194); costa of ­valva covered with tiny spines up to the base of its triangular extension; valva without ventral tubercles; aedeagus as long as valva; vesica evenly speckled with tiny sclerotizations ilicaria species-group (including: P. ilicaria) –– Male antenna with gradually reduced size of branches towards tip. Branches shorter than in the other species (longest branches 0.5 mm) –– Forewing medial areas darker than basal and terminal area –– Forewing medial and postmedial lines at centre deeply curved outwards, then inwards inclined near dorsum –– Hindwing medial line absent or faint –– Terminal line of all wings blackish, narrow, continuous –– In male genitalia uncus long, tubular, without any dorsal extension; costa of valva covered with tiny spines up to the base of its triangularly extension; valva without ventral tubercles; aedeagus as long as valva; vesica membranous and without any sclerotizations Phenology: Depending on the climate and altitude, the species of this genus are uni-, bi- or even trivoltine. For the widely distributed species P. rhomboidaria in warmer habitats, even populations with four generations are reported (Busato et al. 2000; Hartig 1931; Flamigni et al. 2016). Accordingly, the flight time of species in warmer habitats is different from those in colder habitats, but generally, the phenology for most species is in the period from early June to late September. Adults are nocturnal, attracted to both light and bait. Biology: Species of this genus are either oligophagous, exclusively associated with conifers (e.g. Pinaceae, Cupressaceae) or, in several species (e.g. P. rhomboidaria and P. ilicaria) polyphagous and feeding on a wider range of plants from different families (e.g. Fabaceae, Rosaceae, Oleaceae, Fagaceae). Habitat: Inhabiting a variety of habitat types like deciduous forests, orchards, gardens, vegetated slopes with conifers, heaths and flood plains (Heinicke & Müller 1976; Leraut 2009), from sea-level up to over 2200 m. Immature stages: Normally eggs are laid separately or in small groups on the leaves or branches of the food-plants. Freshly laid eggs are green or blue, turning to dark beige or brown; ovally elongated with longitudinal ribs. Adult larvae are light to dark brown. Busato et al. (2000) studied the biology and larval chaetotaxy of P. rhomboidaria larvae and showed that the general setae distribution in this species is similar to that of other genera of the subfamily Ennominae. Egg, larva and pupa of P. ilicaria are described by Goméz de Aizpúrua (1997). 414

Text-figs 185–188. Diagnostic external characters (indicated) of four Peribatodes species-groups. Text-fig. 185. Peribatodes rhomboidaria (Denis & Schiffermüller, 1775), Germany: Berlin-­ Rosenthal, 27.v.2002 (coll. B. Müller). Text-fig. 186. Peribatodes secundaria (Denis & Schiffer­ müller, 1775), Austria: Lavanttaler Alpen, Saualpe, 1335 m, 06.vii.2001 (coll. B. Müller). Text-fig. 187. Peribatodes aragonis (Wehrli, 1943), Spain: 4 km NE Albarracin, 1100 m, 4.ix.2008 (coll. B. Müller). Text-fig. 188. Peribatodes ilicaria (Geyer, 1833), Spain: Aledo West of Totano, 640 m, 4.vi.2003 (coll. B. Müller).

415

Text-figs 189–197. Text-figs 189–195. Diagnostic characters of uncus (lateral view, dorsal side is on right) of four Peribatodes species-groups, species names are given under each photo. The lobe is on the dorsal side. Text-figs 196–197. Two types of male antenna in the genus Peri­ batodes. Text-fig. 196. Male antenna with abruptly reduced length of branches towards its tip, present in the secundaria group, and P. ebusaria. Text-fig. 197. Male antenna with gradually reduced length of pectinations towards its tip, present in the rhomboidaria (except P. ebusaria), powelli and ilicaria species-groups.

Parasitoids: Several Hymenopteran parasitoid species are reported, on P. rhomboidaria (e.g. Pimpla rufipes (Miller); Diolcogaster alvearia (Fabricius), Protapanteles triangulator (Wesmael), P. vitripennis (Curtis), Homolobus discolor (Wesmael), Protapanteles trian­ gulator (Wesmael); Baryscapus endemus (Walker), Glyptapanteles vitripennis (Curtis), Glyptapanteles porthetriae (Muesebeck)). – on P. secundaria (e.g. Protapanteles vitri­pennis (Curtis), Casinaria ischnogaster Thomson, Cratichneumon aspratilis Townes, Momoi & Townes, Cratich­neumon luteiventris (Gravenhorst), Hyposoter caedator (Gravenhorst)). – on P. ilicaria (e.g. Hyposoter notatus (Gravenhorst), for more information see: Yu et al. 2006; Papp 1990; Shaw 2010; Noyes 2017). Systematic position: In the multi-gene analysis of Jiang et al. (2017) Peribatodes clusters with Alcis, and in another analysis Peribatodes is sister to Sardocyrnia + Selidosema (Murillo-­ Ramos et al. 2019). 416

143. Peribatodes rhomboidaria (Denis & Schiffermüller, 1775) Geometra rhomboidaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wiener­ gegend: 101 (Austria: Vienna district). Syntype(s). Phalaena Geometra gemmaria Brahm, 1791: Insektenkalender für Sammler und Oekonomen 2: 255 (Germany: near Mainz). Syntype(s). Alcis australaria Curtis, 1826: Br. Ent. 3: 133(a) (Great Britain: Hampshire, new Forest). Syntype(s). Alcis fimbriaria Stephens, 1831: Illust. Br. Ent. (Haustellata) 3: 188, pl. 30, fig. 1 (Great Britain: Suffolk?). Holotype. Boarmia ichnusaria Ghiliani, 1852: Memorie Acad. Sci. Torino (2) 14: 215 (Italy: Sardinia). Syntypes ♂♀. Flamigni et al. (2016) regarded the population of P. rhomboidaria from Sardinia and Corsica as a valid subspecies because of their darker wing pattern. Study of large series collected from these two islands with dark and lighter wing tones convinced us to regard this as a new synonym of P. rhomboidaria (syn. n.). Boarmia perfumaria Newman, 1865: Entomologist 2: 246 (England: London). Syntype(s). Boarmia psoralaria Millière, 1885: Naturalista sicil. 4 (11): 278 (France: Cannes). Syntype(s). Validated at species rank by Scoble (1999), regarded as a synonym of P. rhomboidaria by Leraut (2009), here confirmed. Boarmia gemmaria melas Constantini, 1916: Atti Soc. Nat. Modena (5) 3: 17 (Italy: Mt. Gibbio). Holotype ♂. Boarmia rhomboidaria sublutearia Zerny, 1927: Dt. ent. Z. Iris 41: 123 (Spain: Andalusia, near Algeciras, Barranco del Algarrobo). Syntype(s). Redondo et al. (2009) and Leraut (2009) regarded this population only as a morphological variation and synonym with P. rhomboidaria, here confirmed. Boarmia rhomboidaria defloraria Dannehl, 1928: Ent. Z., Frankft. a.M. 41 (20): 393 (northern Italy, South Tyrol: Terlan, Sigmundskron). Syntypes 3♂4♀. Regarded as synonym of P. rhomboidaria by Hausmann et al. (2004; 2011a), here confirmed. Boarmia rhomboidaria corsicaria Schawerda, 1931: Z. öst. EntVer. 16 (5): 42 (Corsica: Col de Vizzavona). Syntypes 2♀. Synonymy with ichnusaria Ghiliani, 1852 proposed by Flamigni et al. (2016), the latter name regarded as a synonym of P. rhomboidaria here. Boarmia rhomboidaria syritaurica Wehrli, 1931: Mitt. münch. ent. Ges. 21 (2): 44 (Turkey: ­Marasch; Lebanon: Zahle; Syria: Akbès). Syntypes, including at least 2♂2♀ (ZFMK, examined). Regarded as synonym of P. rhomboidaria by Hausmann et al. (2004; 2011a), here confirmed. Peribatodes rhomboidaria saerdabensis Wehrli, 1941: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 479 (N Iran: Elburs Mts., Särdabtal, Tacht i Suleiman, 1900–2200 m). Holotype ♂ (ZFMK, examined). Regarded as synonym of P. rhomboidaria by Okyar & Mironov (2008), here confirmed.

External characters and abdomen. Wingspan ♂ 29–43 mm, ♀ 30–47 mm. Wings light to dark brown, markings dark brown to black. Forewing antemedial line overall curved outwards; medial line angled outwards just after discal spot, in most specimens merged with discal spot and appearing thicker towards costa; postmedial line weakly dentate on veins, characteristically inclined inwards near costa, and then curved outwards, ending at the centre of dorsum; subterminal line absent and replaced by a dark shadow distal to postmedial line; wavy line present, dentate in its apical part, often faint or replaced by white edging towards tornus. Termen of forewing brown, apical area lighter, subapical area dark brown. Hindwing basal, antemedial and subterminal lines absent; medial line straight, faint towards costa; postmedial line dentate on veins, medially curved outwards, near dorsum curved inwards; wavy line dentate, termen whitish. Discal spots on all wings blackish, smaller on hindwings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes light brown, darker at vein endings. Underside of wings light brown, suffuse. Markings are weakly visible. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 13–15 apical 417

segments without pectination), filiform in female. Setal comb present on male abdominal sternite 3. Variation: Highly variable. Ground colour of wings varying from beige, light grey to dark brown, irrorated with brown or dark brown, or even blackish. Forewing markings can be almost absent (in melanistic specimens), partly visible, or blackish and distinct. Medial area can be concolorous with the rest of the wing or distinctly lighter. Wavy line can be continuous, partially visible or nearly absent. Hindwings are often paler than the forewing. Underside lighter than upperside, dull brown or grey. Male genitalia: Uncus eagle-beak form, apically acute, dorsally extremely extended, setose (dorsal extension larger than in all other European congeners). Gnathos reduced, its basal part slightly sclerotized. Apex of valva as narrow and long as costal valva (variable). Costal part of valva strongly sclerotized, setose at apex and at the base of the subapical extension of costa; subapical extension of costa triangular, located at the centre of costa. Juxta elongated, apically concave. Saccus rounded, at centre slightly projecting. Aedeagus straight, very large (as long as length of valva); vesica with two separate clusters of short spines. Female genitalia: Papillae anales ovally elongated, setose. Apophyses posteriores twice as long as apophyses anteriores. Lamella antevaginalis well sclerotized, widely concave. Ductus bursae funnel-shaped, well sclerotized. Corpus bursae membranous, extremely long (6 mm), posteriorly tubular, anteriorly largely pyriform (larger than in all other European Peribatodes species). Signum strongly sclerotized as a short transverse fold (0.3 mm). Distribution and abundance: West Palaearctic. In Europe widely distributed from southern Urals to British Isles and Portugal and from southern Scandinavia to the Mediterranean including all islands. – In Europe common or very common, now and then found in numbers, occasionally recorded as pest (Voigt 1979; Schanowski & Ebert 2003). – Outside Europe in North Africa, Turkey, Cyprus, Middle East, Caucasus, Transcaucasus, Iran, Turkmenistan, Altai Mts., Kazakh­stan and western provinces of Russia (Flamigni et al. 2016; Viidalepp 1996; Sinev 2008).

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Phenology: Uni- or bivoltine (depending on climate and/or altitude). In Denmark from early July to late August (P. Skou own data), in south-western Germany from mid-May to early October (Schanowski & Ebert 2003). In Italy in two generations from April or May to June or July and from August to October, in warm places even three or four generations (Busato et al. 2000; Hartig 1931; Flamigni et al. 2016). In southernmost Spain and Israel in several generations from mid-March to late November (P. Hale pers. comm.; A. Hausmann own data). Overwintering as larva in its third instar between dry leaves, from late August to June (Heinicke & Müller 1976; Busato et al. 2000). Biology: Larva highly polyphagous on a wide range of trees, shrubs, conifers and herbaceous plants of different families. Found on Ulex europaeus, Pinus sp. and Clematis vitalba (Boldt 1929), ornamental Clematis (Hausmann, own data) and Juniperus sabina (Lechner & Ortner 1999). Recorded as a pest species on grape (Vitis vinifera) (Voigt 1979; Schmid et al. 1983; Busato et al. 2000). Recorded on Apocynaceae (Nerium); Araliaceae (­Hedera); Asteraceae (Achillea, Artemisia, Cirsium arvense); Berberidaceae (Berberis thunbergii); Betulaceae (Betula, Corylus); Buxaceae (Buxus); Caprifoliaceae (Lonicera tatarica); Caryo­ phyllaceae (Cerastium tomentosum); Celastraceae (Euonymus europaeus); Cistaceae (Cistus, Helianthemum nummularium); Clusiaceae (Hypericum perforatum), Convolvulaceae (Calystegia); Cupressaceae (Juniperus); Ericaceae (Calluna vulgaris, Vaccinium uliginosum); Fabaceae (Calicotome, Colutea, Cytisus scoparius, Genista tinctoria, Lembotropis nigricans, Medicago, Psoralea, Spartium); Fagaceae (Quercus robur); Lamiaceae (Rosmarinus, Thymus serpyllum); Malvaceae (Malva, Althaea); Oleaceae (Fraxinus, Ligustrum, Syringa); Pinaceae (Picea abies, Pinus); Plantaginaceae (Plantago); Polygonaceae (Rumex); Rhamnaceae (Rhamnus cathartica); Rosaceae (Crataegus, Prunus domestica, P. spinosa, Rosa canina, Rubus idaeus, Pyrus, Malus, Sorbus aucuparia); Rubiaceae (Galium); Salicaceae (Populus, Salix); Taxaceae (Taxus); Ulmaceae (Ulmus) (Vorbrodt & Müller-Rutz 1914; Albers 1940; Csóka 1991; Lechner & Ortner 1999; Flamigni et al. 2016). Habitat: Ubiquitous. Occurring in a variety of habitats like deciduous forests and its surroundings, orchards, gardens, heaths and flood plains (Heinicke & Müller 1976; Leraut 2009). From sea-level up to 1500 m, in Sierra Nevada in southern Spain up to 2000 m (Redondo et al. 2009), in the Aosta Valley in north-western Italy up to 2100 m (Flamigni et al. 2016). Similar species: Most of all, P. rhomboidaria may be confused with P. perversaria, however the latter is generally lighter in colour and its markings are softer. Additionally, the structure of male and female genitalia allows a reliable identification. The setal comb on the third abdominal sternite of male is present in P. rhomboidaria (in European Peribatodes species, this character is present only in P. rhomboidaria, P. umbraria and P. ilicaria). Genetic data: BINs: BOLD:ACE7297 (n=60 from Germany, Austria, Hungary, United Kingdom, France, Italy, Spain, Morocco, Croatia, Greece, Turkey, Lebanon, Jordan, Israel), and a second BIN BOLD:AAA4448 (n=86 from Germany; United Kingdom, France, Netherlands, Spain, Portugal, Morocco, Italy, Turkey, northern Iran), both BINs at a minimum distance of 1.2% from each other, both BINs genetically heterogeneous, showing regional patterns. One DNA barcode from Tunisia (BOLD:ACF0923) diverging by 1.6%. Nearest species: P. perversaria (2.0%), Peribatodes omalensis sp. n. from Crete (3.0%).

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144. Peribatodes perversaria (Boisduval, 1840) Boarmia perversaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 197 (Switzerland: Valesia). Syntype(s). Boarmia abstersaria Boisduval, 1840: Genera Index meth. eur. Lepid.: 197 (Pyrenees). Syntype(s). Most specimens from Pyrenees and eastern Spain show slightly dark brown wings (compared with light brown to yellow specimens from Alpes-Maritimes and ochreous brown specimens from Valais). However, in both Pyrenees and southern France, specimens occur with exactly the same colour combination as those in Valais. Moreover, in the distribution area of abstersaria there are specimens with colour combination exactly matching that of subflavaria and vice versa. Large series of examined specimens from Valais, southern France and eastern Spain could not confirm any diagnostic character on male or female genitalia (in accordance with Leraut 2009 and Flamigni et al. 2016). Additionally these populations do not show any genetic divergence (based on DNA barcoding data). Biological studies on these populations are scarce and not sufficient to definitely confirm their separation. According to this data, we regard abstersaria and subflavaria as junior synonyms of perversaria. Boarmia buxicolaria Mabille, 1873: Annls Soc. ent. Fr. (5) 2 (4): 490, pl. 15, figs 1, 2 (southern France). Lectotype ♂ (a male labelled as paratype, in coll Herbulot, ZSM, examined). Lhomme (1923–35) confirmed high similarity of wing pattern of buxicolaria, subflavaria and perversaria, as well as larval patterns. Wehrli (1943) mentioned also similarity of genitalia with subflavaria and left the question open. Mazel (2008), faced with the same problem, could not easily separate perversaria, abstersaria and subflavaria based on genitalia. ­Leraut (2009) regarded this only as a form of P. perversaria, this synonymy was confirmed by F ­ lamigni et al. (2016), here again confirmed. Boarmia subflavaria Millière, 1876: Cat. Lépid. Alpes-Maritimes (3/4): 404 (France: Alpes Mari­ times, vallée de Thorenc, Pic de l’Aigle). Syntype(s). Junior synonym of perversaria (see taxonomic note under abstersaria). Peribatodes abstersaria montserrata Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 482, pl. 42: i (Spain: Catalonia, Montserrat, 1000–1200 m). Syntypes 5♂2♀ (ZFMK, examined). Leraut (2009) regarded this as a darker form of abstersaria, the latter name regarded here as a junior synonym of perversaria. Boarmia perversaria galliberia Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 480, pl. 42: h (Spain: Prov. Teruel, Albarracín; Andalusia). Syntypes, ♂♀ (ZFMK, externally examined). Leraut (2009) regarded this as a darker form of abstersaria, the latter name here regarded as a junior synonym of perversaria.

External characters and abdomen. Wingspan ♂ 29–39 mm, ♀ 32–40 mm. Ground colour of wings from yellow to ochreous brown and dark creamy, markings dark brown. Forewing ante­medial line angled outwards near costa. Forewing medial line at centre curved outwards; postmedial line weakly dentate on veins, slightly inclined inwards near the costa, and then curved outwards, then continuing to middle of dorsum very close to medial line, even anastomosing at some points; subterminal line absent and replaced by a brown shadow distal to postmedial line. Wavy line faint with white edging, weakly dentate near apex. Termen of forewing covered with darker scales, except a light apical area. Basal, antemedial and subterminal lines absent from hindwing; medial line straight, faint towards costa; postmedial line dentate on veins, at centre curved towards apex, near dorsum slightly curved toward base; wavy line irregularly dentate, termen whitish. Discal spots on all wings blackish, very small on hindwings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes light brown, darker at vein endings. Underside of wings lighter than upperside. Markings are weakly visible. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 10–12

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apical segments without pectination whilst 13–15 in P. rhomboidaria; 15–17 in P. umbraria), filiform in female. Setal comb absent from male abdominal sternite 3. Variation: Extremely high variation in wing colour (this colour variation led to the description of many taxa), but not in genitalia and DNA barcodes. Wings show a wide range of colour tones from yellow to light brown and ochreous brown. The intensity of the forewing markings varies from being partly visible, or distinctly light to dark brown. Medial area can be concolorous with the rest of the wing or distinctly lighter. Wavy line can be continuous, partially visible or nearly absent. Male genitalia: Genital capsule smaller and similar to that in P. rhomboidaria, except the following differences: in P. perversaria uncus shorter, dorsally extended (two-thirds of the extension in P. rhomboidaria); valva basally narrower than in P. rhomboidaria; apex of valva narrower and shorter than costa of valva (in contrast to rhomboidaria); subapical extension of costa closer to apex of costa (at mid costa in P. rhomboidaria); juxta elongated, slightly sclerotized only; saccus rounded. Aedeagus very large (longer than length of valva), straight; vesica with two separate clusters of short spines. Female genitalia: Very similar to those of P. rhomboidaria, except the following differences: in P. perversaria posterior and anterior apophyses slightly shorter than in P. rhomboidaria; ductus bursae cylindrical, smaller than in P. rhomboidaria; corpus bursae extremely long (6 mm), posteriorly tubular but narrower than in the pyriform anterior part, the size of the latter part half of that in P. rhomboidaria; signum twice as large as in P. rhomboidaria (0.6 mm in transverse measurement). Distribution and abundance: (South-west) European: Valais mountains (south-western Switzer­ land, ‘Wallis’), Alpes Maritimes in south-eastern France, eastern and northern Spain (from Pyrenees towards north-western Spain).

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Biology: Larva oligophagous. Recorded on Juniperus sabina and Cupressus (Vorbrodt & Müller-Rutz 1914), on Genista (Lhomme 1923–35), and Buxus sempervirens (Prout 1915). Larva of Iberian populations (‘abstersaria’) feeding on Genista and Quercus (Redondo et al. 2009). Phenology: Univoltine (Ziegler 2019; Robineau 2007). From June to mid-August in the Alps, in Spain from mid-July to late September (P. Skou own data). Overwintering as larva. Habitat: Xeromontane. Warm and dry Alpine slopes (e.g. rocky steppes of Valais) with scattered trees and bushes of Juniperus or Genista. From 400 up to 1700 m, and up to 2200 m in southern French Alps. Similar species: P. perversaria may be confused with some colour variations of P.  rhomboidaria. The former shows smoother patterns and is generally lighter. Additionally, the setal comb on male abdominal sternite 3 is absent in P. perversaria (present in P. rhomboidaria); in male antennae the last 10–12 apical segments have no pectination (13–15 in P. rhomboidaria). Study of the genitalia structure and/or using DNA barcodes allows a reliable identification. Genetic data: BIN: BOLD:ABZ2455 (n=9 from France, Spain, pre-identified as ‘P. abstersaria’ (synonym), as well as n=12 from France, north-western Italy, and n=3 from Switzerland (Valais), pre-identified as ‘P. perversaria’). Genetically homogeneous, perversaria and ‘abstersaria’ (synonym) exactly sequence-sharing. The DNA sequence of the lectotype of buxicolaria (ZSM, coll. Herbulot) proves its synonymy with P. perversaria (100% genetic congruence). Nearest species: P. rhomboidaria (2.0%).

145. Peribatodes omalensis Rajaei, Hausmann & Skou, sp. n. Examined material. Holotype: ♀, Greece, Crete, Nea Roumata [ca. 350 m], [35.397° N, 23.852° E], leg. K. Szeoeke, 09.06.1998 (ZSM, DNA barcoded BC ZSM Lep 09586). Paratypes: 1♂4♀ Greece, western Crete, Omalos Plateau South of Chania, 35°20’39’’N, 23°54’18’’E, 27.06.2012, 1050 m, leg. C. Hviid, K. Larsen; 1♂3♀ with same data, in coll. Skou, 1♀ with same data, in SMNS; 1♀, Greece, Crete, Omalos plateau, road to Kaligeri, 1225 m, 15–20.vi.2014, leg. O. Karsholt, C. Hviid & F. Vilhelmsen. In ZMUC; 2♂ Greece (Hellas), Nom. Chania, 1150 m, 35°21’11” N, 21°54’39” E, at pass, N of Omalos plateau, 07.vi.2004, leg. B. Skule, C. Hviid, E. Vesterhede, coll. Vesterhede.

External characters and abdomen. Wingspan ♂ 26–30 mm, ♀ 30–32 mm. Wings uniformly dark grey, markings dark grey to black, hardly visible. Forewing antemedial line curved, ­angled outwards near costa (in contrast to the other species of the rhomboidaria group), then continuing straight to dorsum; medial line faint and hardly visible as discontinuous blackish dots; postmedial line weakly dentate on veins, inclined inwards near the costa, then continuing straight to middle of dorsum; subterminal line absent; wavy line mostly faint, hardly visible, partly dentated. Termen of forewing slightly darker, except apex. Basal, antemedial and subterminal lines absent from hindwing; medial line hardly visible, straight; postmedial line dentate on veins, at centre curved inwards; wavy line irregular, distally with a thin, light edging. Discal spots of forewings clearly visible, very small or invisible on hindwings, blackish. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes dark grey, darker at vein endings. Underside of wings dark grey, ­basal and medial areas lighter than termen; only the postmedial line is visible. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, 422

­ ipectinate in male (except 10–12 apical segments without pectination), filiform in female. b Setal comb absent from male abdominal sternite 3. Variation: All specimens of the type series are uniformly dark grey and show little variation in colour. Male genitalia: Uncus eagle-beak-shaped, apically acute, subapically slightly inflated, dorsally extremely extended (less than in P. rhomboidaria), setose. Gnathos reduced, its basal part slightly sclerotized. Valva basally broad (but narrower than in P. rhomboidaria), marginally and ventrally setose. Membranous apex of valva as narrow and long as sclerotized costa of valva; valva extended subapically. Costal part of valva strongly sclerotized, apically setose, distally separated from membranous apex of valva. Triangular subapical extension of costa located at two-thirds from base (at mid-costa in P. rhomboidaria). Juxta elongated, apically more extended. Saccus rounded. Aedeagus straight, shorter than valva (same length as valva in P. rhomboidaria); vesica with two separate clusters of short spines. Female genitalia: Papillae anales oval, elongated, setose. Apophyses posteriores twice as long as apophyses anteriores. Lamella antevaginalis slightly sclerotized. Ductus bursae funnel-shaped, well sclerotized (narrower than in P. rhomboidaria). Corpus bursae membranous, long (but less than 5 mm whilst 6 mm in P. rhomboidaria and P. perversaria), tubular, its anterior part slightly pyriform (size of the latter part half that in P. rhomboidaria). Signum strongly sclerotized as a short transverse fold (0.3 mm). Distribution and abundance: Recorded only from Crete, Greece. – Apparently very rare; after checking many collections no other specimens traced than those included into the type series.

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Phenology: Possibly univoltine. So far, scarce records from early June to late June. Biology: Unknown. Habitat: Xeromontane. Recorded from rocky steppes with scattered deciduous trees and shrubs at altitudes from 350 up to 1225 m. Similar species: On Crete two sympatric congeners occur (P. rhomboidaria, P. correptaria). The species can easily be diagnosed based by its comparatively small size, the dark grey wings, the shape of antennal pectination (which is more similar to that of the secundaria group), and the lack of the setal comb on male sternum A3 (the latter character present only in P. rhomboidaria, P. umbraria and P. ilicaria). Additionally, both male and female genitalia show diagnostic characters (uncus, valva, size of aedeagus in male; size and shape of corpus bursae in female). For more reliable identification, DNA barcoding is recommended. Genetic data: BIN: BOLD:AAI2403 (n=5 from Crete). Intraspecific (maximum) variation low (0.3%). Nearest species: P. perversaria (2.5%), P. rhomboidaria (3.0%), the latter also DNA barcoded from Crete (n=3). At a distance of 6.9% from the third Peribatodes species on Crete island, P. correptaria. Etymology. The name of the species refers to the type locality (Omalos Plateau, south of Chania, Crete).

146. Peribatodes umbraria (Hübner, 1809) Geometra umbraria Hübner, 1809: Samml. Eur. Schmett. 5 Geometrae (1), pl. 65, fig. 340 (Europe). Syntype(s). Boarmia umbraria decosteraria Oberthür, 1913: Études Lépid. comp. 7: 264, pl. 167, fig. 1631 (France: Alpes-Maritimes, Menton). Syntype(s), including ♀. Junior synonymy of nominotypical subspecies suggested by Hausmann et al. (2004; 2011a), confirmed by Leraut (2009), here confirmed. ‡ Boarmia umbraria powelli Oberthür, 1913: Études Lépid. comp. 7: 264, pl. 167, figs 1632, 1633 (Algeria: South Oran, Kenchela; Aflou; Géryville). Syntypes ♂♀. Junior secondary homonym of Calamodes powelli Oberthür, 1913: 262. Boarmia umbraria var. vaucheri Thierry-Mieg, 1916: Miscnea ent. 23: 48 (Morocco: Tanger). Syntypes 1♂1♀ (MNHN). Validated at subspecies level by Leraut (2009). Confirmation of this subspecies needs additional research based on larger series. Boarmia umbraria khenchelae Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 482. Replacement name for powelli Oberthür (Algeria, Lambèse, leg. Harold Powell). Syntypes ♂♀, (ZFMK, examined). Junior synonymy of subsp. vaucheri suggested by Leraut (2009), here confirmed. Boarmia umbraria panorma Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 483, pl. 43: a (Italy: Sicily, Palermo). Syntype, ♂ (ZFMK, examined). Junior synonymy of nominotypical subspecies suggested by Leraut (2009), confirmed by Hausmann et al. (2004; 2011a), F ­ lamigni et al. (2016), here confirmed. Boarmia umbraria syrirana Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 483, pl. 43: a, b (SW Iran: Kasrun). Syntypes 6♂1♀ (ZFMK, examined). Colour combination falls within the variation range of the nominotypical subspecies. Validated at subspecies rank in Scoble (1999), here downgraded to synonymy of the nominotypical subspecies (syn. n.). Boarmia umbraria mimeuri Rungs, 1950: Bull. Soc. Sci. nat. phys. Maroc 28: 157, pl. 5, figs 6, 7 (Morocco: Middle Atlas, Ifrane, 1650 m). Junior synonymy of subsp. vaucheri suggested by Leraut (2009), here confirmed.

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Boarmia umbraria matrensis Vojnits, 1970: Acta zool. hung. 16 (3/4): 458, fig. 1a (Hungary: Fot). Holotype ♂ (TMB). Junior synonymy of nominotypical subspecies suggested by Hausmann et al. (2004; 2011a), here confirmed. Peribatodes umbraria perumbraria Leraut, 2009: Moths of Europe, 2: 166, fig. 9, p. 42 (Corsica). Holotype ♀, paratypes 9♂13♀ (MNHN, examined). Colour combination falls within the variation range of the nominotypical subspecies. Validated at subspecies rank in Leraut (2009), here downgraded to synonymy of the nominotypical subspecies (syn. n.).

External characters and abdomen. Large-sized, wingspan ♂ 31–43 mm, ♀ 35–47 mm. Wings from light to dark grey brown, sometimes irrorated with olive-green markings distinctly dark brown to black. Forewing antemedial line angled or curved outwards, sometimes faint and hardly visible; medial line crossing the discal spot, at centre curved outwards, towards costa thicker and in most specimens anastomosing with postmedial line towards dorsum; postmedial line weakly dentate on veins, slightly inclined inwards near costa, and then curved outwards, continuing to middle of dorsum, curved again toward base near dorsum; subterminal line absent. Wavy line distinct, thick and dentate with hardly visible white edging, faint towards tornus. Termen of forewing covered with slightly darker scales, except a light apical area and much darker subapical area. Basal, antemedial and subterminal lines absent from hindwing; medial line thick, straight, faint towards costa; postmedial line dentate on veins, at centre curved towards apex, near dorsum slightly curved toward base. Hindwing wavy line distinct, irregularly dentate, termen light brown, except a darker spot on M-veins. Discal spots on all wings blackish, mostly merged with medial line on forewings and very small on hindwings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes brown, darker at vein endings. Underside of wings lighter than upperside, with pale blotches on apex and terminal area of forewing. Markings present, brownish. Frons, collar and abdomen concolorous with wings. Thorax and tegulae covered with ivory scales. Antennae dorsally concolorous with wings, bipectinate in male (except 15–17 apical segments without pectination; 10–12 in P. perversaria; 13–15 in P. rhomboidaria), filiform in female. Setal comb present on male abdominal sternite 3. Variation: The species shows a high variation in coloration and size. In North Africa (Algeria and Morocco) replaced by subsp. vaucheri, which is also variable, but constantly darker than the nominotypical subspecies, with warmer brown, irrorated with dark olive-green. Male genitalia: Uncus triangular, apically acute, dorsally well extended, rectangular (beakshaped in P. rhomboidaria). Gnathos absent. Valva basally broad, lateral margins convex, setose, at centre with five rows of thin setae (two or three rows in the other species of the rhomboidaria group). Apex of membranous valva as long as costa of valva; the latter apically setose, with a large triangular subapical extension. Saccus rounded. Aedeagus very large (as long as valva), straight, at centre slightly curved; vesica with only one cluster of short spines (in contrast to the other species of the rhomboidaria group). Female genitalia: Smaller, but very similar to those of P. rhomboidaria, except the following differences: in P. umbraria apophyses more stout and clearly longer than in P. rhomboidaria; lamella antevaginalis concave; ductus bursae funnel-shaped, clearly larger and wider than in P. rhomboidaria; corpus bursae (length 4 mm) posteriorly swollen, anteriorly apple-shaped and clearly smaller than in P. rhomboidaria; signum (0.6 mm in transverse measurement) twice as large as that of P. rhomboidaria.

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Distribution and abundance: Mediterranean-Turanian. Southern Europe from west to east. – In southern Europe not rare, sometimes abundant. – Outside Europe from Cyprus, Turkey, Caucasus and Transcaucasus, to the Levant (Syria, Israel, Jordan, Iraq), Iran, Turkmenistan and Afghanistan. In North Africa (Morocco to Tunisia) replaced by subsp. vaucheri.

Phenology: Bivoltine, from early May to early July and from mid-August to late October (P. Skou own data). Sometimes, e.g. in Israel, emerging from mid-April, in southern Italy and Israel until early November (A. Hausmann own data). Biology: Larva oligophagous. Found on Olea europaea (Vorbrodt 1931; Mazel 2008; ­Flamigni et al. 2016), Quercus ilex and Q. suber (Luciano & Roversi 2001) and ­Pistacia (­Nestorova 1998). Recorded also on Juniperus communis, Pinus sylvestris and Picea (­Vorbrodt & MüllerRutz 1914). Habitat: Xerothermophilous. Oak woodlands, vegetated slopes and mountains with coni­ ferous trees. In Europe from sea-level up to 1800 m, up to 1550 m in Cyprus, and up to 2100 m in Morocco (P. Skou own data). Similar species: The species cannot be confused with any other Peribatodes species considering its large size and prominent markings. Genetic data: BINs: BOLD:AAB2950 (n=25 from France, Spain, Italy, Croatia, Greece, Bulgaria, Turkey); BOLD:AAT9436 (n=6 from Morocco). Genetically homogeneous within the BINs, both BINs diverging by 2.4%. Nearest species: P. ebusaria (3.9%), P. perversaria (4.5%), P. rhomboidaria (4.8%).

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147. Peribatodes ebusaria Vallhonrat, 2012 Peribatodes ebusaria Vallhonrat, 2012: Butll.Soc. Cat. Lep. 103: 33–39 (Spain, Baleares: Island of Ibiza). Holotype ♂ (MCNB).

External characters and abdomen. Wingspan first generation ♂♀ 28–33 mm, second generation smaller, ♂♀ 24–27 mm. Wings light brown to grey, markings dark brown. On forewing two parallel antemedial lines visible (characteristic), curved outwards. Forewing medial line not connected to discal spot, at centre curved outwards, towards dorsum anastomosing with postmedial line; postmedial line weakly dentate on veins, curved outwards, then continuing to middle of dorsum, curved again towards base near dorsum. Forewing subterminal and wavy lines hardly visible; apex lighter. Basal, antemedial and subterminal lines absent from hindwing; medial line straight, faint towards costa; postmedial line weakly dentate on veins, slightly curved towards apex; wavy line faint, discontinuous. Discal spot round, black, present on all wings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes pale grey, darker at vein endings. Underside of wings lighter than upperside. Markings present, brownish. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, male antenna bipectinate (except 8–9 apical segments without pectination), shape of pectination similar to that of the species of secundaria group (with abruptly reduced size of branches towards its tip), in contrast to the gradually reduced size of branches towards tip in the species of the rhomboidaria group; female antennae filiform. Setal comb absent from male abdominal sternite 3. Male genitalia: Uncus apically acute, very short (much shorter than in all other species of the rhomboidara group), dorsally extremely extended (nearly of same size as in P. rhomboidaria). Gnathos absent. Valva basally broad with one row of long and stout spines on sacculus, and three rows on the ventral part. Apex of valva broader and slightly shorter than sclerotized costa; the latter with a triangular extension arising from apex, apically setose, distally separated from membranous apex of valva. Juxta elongated, slightly sclerotized. Saccus rounded. Aedeagus considerably shorter than valva, straight; vesica curved, with two separate clusters of short spines. Female genitalia: Judging from the scale bar in Vallhonrat (2012), genitalia smaller than in all other congeners. Papillae anales elongated, narrow, setose. Lamella antevaginalis well sclerotized, widely concave. Ductus bursae short, funnel-shaped, well sclerotized. Corpus bursae membranous, shorter (2 mm) than in the other species, posterior part narrower than anterior part, the latter pyriform. Signum sclerotized as a short transverse fold (0.15 mm). Distribution and abundance: Endemic to the Balearic island of Ibiza. Phenology: Bivoltine. Scarce data from late March, early April, mid-May, late September and early October. Biology: Unknown. Habitat: Warm and dry open habitats with scattered trees. From sea-level up to at least 250 m according to the few collected specimens from Ibiza. Apparently common in Ibiza (P. Skou own data). Similar species: P. ebusaria may be confused with P. perversaria, but the shape of male antenna, wing pattern (e.g. antemedial line and other wing markings), genitalia structures and geographical distribution should provide a certain identification.

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Genetic data: BIN: BOLD:AAL2704 (n=1 from Spain: Balearics). Nearest species: P. perversaria (3.8%), P. umbraria (3.9%).

148. Peribatodes secundaria (Denis & Schiffermüller, 1775) Geometra secundaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (France). Syntype(s). Boarmia gemmaria fallentaria Staudinger, 1879: Horae Soc. ent. ross. 14: 452 (Asia Minor: Siaret). Holotype ♂. Valid at subspecific rank (Scoble 1999), confirmation of the validity of this subspecies requires further study. Boarmia secundaria var. druentiaria Cleu, 1928: Amat. Papillons 4: 90, pl. 1, figs 4, 5 (France: Hautes-Alpes, Vallé de la Durance). Syntypes. Here regarded as a synonym of the nominotypical subspecies based on sympatric occurrence (as previously stated by Scoble 1999). Boarmia secundaria syrisca Wehrli, 1933: Dt. ent. Z. Iris 47 (3): 99, pl. 1, fig. 40 (northern Lebanon). Syntypes 5♂2♀. Boarmia secundaria f. francosuevica Schneider, 1942: Z. wien. EntVer 27 (8): 186, pl. 18, figs 4–9 (Germany: Hessental-Fischachtal). Syntypes ♂♀. Boarmia secundaria f. rometschi Schneider, 1942: Z. wien. EntVer 27 (8): 185, pl. 18, figs 2, 3 (Germany: Baden, Pforzheim). Syntypes ♂♀. Peribatodes secundaria occidentaria Lempke, 1952: Tijdschr. Ent. 95: 233, pl. 53: 1–3 (Netherlands: Puten). Holotype ♂ (ITZA). Here regarded as a synonym of the nominotypical subspecies based on sympatric occurrence. Peribatodes dragone Parenzan & Laever, 1985: Revta Lepid. 13 (3): 178, figured (Italy: Pollino Massif, Colle del Dragone, 1800 m). Holotype. Junior synonymy proposed by Flamigni et al. (2016).

External characters and abdomen. Wingspan ♂ 26–34 mm, ♀ 34–45 mm. Wings beige, yellow-brown to dark brown, medial area slightly lighter than basal and postmedial areas, markings distinctly dark brown. Forewing antemedial line at centre curved outwards 428

(straight in P. correptaria); medial line slightly curved outwards, towards dorsum continuing near postmedial line (rarely anastomosing), slightly curved inwards. Forewing postmedial line sharply dentate on radial veins (in contrast to P. correptaria), slightly bent outwards on M3 vein (straight in P. correptaria), continuing to middle of dorsum, curved deeply inwards (slightly curved inwards in P. correptaria). Forewing subterminal line present, discontinuous, visible only from underside; wavy line discontinuous, sometimes faint, irregularly wavy with a weak white edging. Termen of forewing covered with dark brown scales, with distinct pale blotch near outer margin, and a darker subapical area. Hindwing medial line straight, faint towards costa; postmedial line dentate, slightly curved outwards, near dorsum slightly curved inwards. Subterminal line present, visible from underside only. Hindwing wavy line mostly faint, with a light edging. Discal spots, small, present on all wings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes brown, darker at vein endings. Underside of wings smooth, lighter than upperside, markings light brownish. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 9–10 apical segments without pectination), filiform in female. Setal comb absent from male abdominal sternite 3. Variation: Highly variable. Typical forms are dark beige, shaded dark grey or blackish mixed with lighter scales. There are also paler or darker forms with more homogeneous scaling. Even melanistic forms occur. Markings variable, from dark brown to blackish. In some melanistic specimens markings dull to completely obsolete. Male genitalia: Uncus apically acute, dorsally setose, slightly swollen (narrower in P. correptaria). Gnathos reduced. Valva basally broad with one row of long spines at sacculus valva, and two or three rows at ventral part; centre of valva with ventral tubercles. Membranous apex of valva wide (narrower in P. correptaria), shorter than costa. The latter strongly sclerotized, apically narrow (longer than that in P. correptaria), subapically with a triangular extension (more extended in P. correptaria). Juxta broad, apically triangular (shorter than that in P.  correptaria). Saccus rounded. Aedeagus straight, shorter than length of valva; vesica with one cluster of strongly sclerotized spines (less numerous and with smaller spines in P. correptaria). Female genitalia: Papillae anales oval, setose. Apophyses posteriores twice as long as apo­ physes anteriores. Lamella antevaginalis well sclerotized, concave. Ductus bursae slightly funnel-shaped. Base of ductus seminalis enlarged. Corpus bursae membranous, long (5 mm, clearly larger than that in P. correptaria), tubular. Signum strongly sclerotized as a short transverse fold, of same size as in P. rhomboidaria (0.3 mm). Distribution and abundance: European or European-West Asiatic. The species occurs all over central and northern Europe (from eastern Spain to western Ukraine). In southern ­Europe, from Italy to Balkan countries, and with fragmented populations in eastern Ukraine and southern European Russia. Reported in Corsica by Rungs (1982). – In central Europe often common, towards the south usually rare and very local. – Outside Europe reputedly in Turkey (Staudinger 1879; Viidalepp 1996; cf. genetic data) and northern Lebanon (Wehrli 1933), requiring further confirmation. Phenology: Univoltine, flying from mid-June to late September. Overwintering as larva between dry needles of the food-plants (Schneider 1942; Euroleps 2018).

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Biology: Oligophagous. Larva found on Picea excelsa and Pinus sylvestris (Schneider 1942; Dvorˇáčková & Kulfan 2009) and Ulex europaeus (Boldt 1929). Recorded also on Abies alba (Schanowski & Ebert 2003), and Juniperus communis (Vorbrodt & Müller-Rutz 1914; Heinicke & Müller 1976) Habitat: Silvicolous, mesophilous. Occurring in temperate and mixed coniferous forest (Heinicke & Müller 1976). From sea-level in northern Europe up to 1700 m in Greece. A few records from higher altitudes, e.g. 2300 m in the Alps (P. Skou own data). Similar species: The species may be confused with P. correptaria. These two species can be externally diagnosed based on forewing antemedial line medially curved outwards in P. secundaria (straight in P. correptaria); forewing postmedial line sharply dentate on radial veins, slightly bent on M3 vein outwards in P. secundaria (not dentate and not bent in P. correptaria), continuing to middle of dorsum, curved deeply inwards (slightly curved in P. correptaria). In male genitalia the apical part of valva thicker, apical part of costa longer and subapical costal extension shorter in P. secundaria. Vesica with more and longer spines in P. secundaria. Genetic data: BIN: BOLD:AAC2923 (n=32 from Finland, Germany, Austria, Italy, Macedonia, Turkey). Genetically homogeneous, exactly barcode-sharing with P. correptaria. The barcoded Turkish specimen requiring further study to discriminate it from P. correptaria. Next nearest species: P. umbraria (5.9%).

149. Peribatodes correptaria (Zeller, 1847) Boarmia correptaria Zeller, 1847: Isis 1847 (1): 18 (Turkey: near Istanbul, Pera; Smyrna). Syntypes 2♂ (NHMUK). Boarmia syrilibanoni Wehrli, 1931: Mitt. münch. ent. Ges. 21 (2): 46 (Lebanon: Bescharre). Holotype ♀ (ZSM). Regarded as a synonym of P. correptaria by Okyar & Mironov (2008).

430

Boarmia syrilibani Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 484, pl. 43: d. Emendation of syrilibanoni Wehrli.

External characters and abdomen. Wingspan ♂ 26–32 mm, ♀ 30–34 mm. Wings light chocolate-­brown to dark brown, markings distinctly dark brown. Forewing postbasal area slightly lighter than the rest of the wing (postbasal area concolorous with other areas of the wing in P. secundaria). Forewing antemedial line deeply angled outwards near costa, sometime a second parallel line is visible (medially curved outwards in P. secundaria and without second parallel line); medial line slightly curved outwards, faint towards costa, continuing towards dorsum adjacent to postmedial line (rarely connected); postmedial line smooth (dentate in P. secundaria), slightly inclined near costa, gently curved inwards, continuing to middle of dorsum; subterminal line present as a brown shadow, visible as separated spots on veins from underside; wavy line discontinuous, smooth (dentate in P. secundaria), with a weak white edging. Termen of forewing covered with dark brown scales, with a light apical spot, and a darker subapical area. Hindwing medial line straight, faint towards costa; postmedial line smooth or slightly dentate, gently curved outwards, near dorsum slightly curved inwards (strongly curved inwards near dorsum in P. secundaria); subterminal line present but visible only from underside, near costa connected to postmedial line; wavy line irregularly wavy, with a light edging. Discal spots blackish on forewings and mostly merged with medial line, absent or very small on hindwings. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes brown, darker at vein endings. Under­side of wings lighter than upperside, markings light brownish. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 9–10 apical segments without pectination), filiform in female. Setal comb absent from male abdominal sternite 3. Variation: The species shows variation in colour from light to dark brown or grey. Male genitalia: Very similar to those of P. secundaria, except the following differences: genital capsule smaller, uncus dorsally less swollen. Valva basally broad with one row of long spines on sacculus, and two or three rows on ventral part; membranous apex of valva narrower than in P. secundaria; centre of valva with ventral tubercles. Sclerotized costa of valva shorter than in P. secundaria, subapical extension larger than in the latter. Juxta elongated, apically rounded (shorter in size and apically triangular in P. secundaria). Saccus wide, rounded. Aedeagus straight, shorter than length of valva; vesica with a cluster of strongly sclerotized spines (P. secundaria with more and larger spines). Female genitalia: Very similar to those of P. secundaria, except the following differences: in P. correptaria ductus bursae medially more compressed; corpus bursae slightly shorter (4 mm). Signum strongly sclerotized as a short transverse fold, same size as in P. secundaria (0.3 mm). Distribution and abundance: East Mediterranean. Distributed in Italy (north-westernmost I­taly and Gargano: Flamigni et al. 2016) and Balkan countries to southern Ukraine. – Usually local and not common. – Outside Europe across Turkey and Lebanon to Jordan and Israel (A. Hausmann own data). Phenology: Bivoltine. From early April to early August and from mid-September to mid-October. Biology: Larva oligophagous. Recorded on Cupressus sempervirens (Albers 1941), Thuja, Pinus nigra, P. brutia (Mol 1977). Reared on Picea (E. Friedrich pers. comm.). 431

Habitat: Xerophilous. Open grasslands with bushes and scattered conifers. From sea-level up to 2000 m. Similar species: P. secundaria (see no. 148). Genetic data: BIN: BOLD:AAC2923 (n=14 from Croatia, Bulgaria, Greece including Crete, Cyprus, Lebanon, Jordan). Genetically homogeneous in the main cluster, ­exactly barcode-sharing with P. secundaria. Next nearest species: P. umbraria (5.9%). P. correptaria with isolated, strongly diverging populations in Lebanon and Jordan (n=4, BIN: BOLD:AAE6308), at a distance of 3.4%.

150. Peribatodes aragonis (Wehrli, 1943) Boarmia powelli aragonis Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 484, pl. 43: c (Spain: Aragon). Holotype ♂ (ZFMK, examined). Raised to species rank by Leraut (2009), here confirmed based on DNA barcode data. Peribatodes aragonis gerini Leraut, 2009: Moths of Europe: 169, pl. 42: 17, 18 (Morocco, Middle-­ Atlas, Agulmane, Sidi Ali, 2070 m, 1.–14.vii.1939). Holotype ♂, paratype ♀ (in MNHN, examined), these two old specimens from the distribution range of P. powelli are slightly paler than in the latter species, but genitalia are well matching those of P. aragonis. Subspecific status requiring confirmation and further study.

External characters and abdomen. Wingspan ♂ 29–33 mm, ♀ 30–34 mm. Wings beige to dark brown, markings dark brown. Forewing antemedial lines (two parallel lines) deeply angled outwards near costa, their costal part faint; medial line very thin, straight, hardly visible, not connected to costa; postmedial line starting from costa, straight or with loose waves continuing to dorsum, with a second dark brown line parallel to postmedial line; subterminal line straight, apically interrupted by the creamy apical spot; wavy line subapically connected to a dark brown streak, towards dorsum faint, with an external light edging. Termen of forewing light (except a 432

dark brown subapical streak), apical area creamy. Hindwing medial line faint towards costa; postmedial line at centre curved inwards; subterminal and wavy lines faint towards apex. Discal spots present on all wings. Terminal line of all wings blackish, narrow, continuous. Fringes light brown, unicolorous on all parts of the wings. Underside of wings lighter than upperside. Markings are weakly visible. Frons, collar, thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 6–8 apical segments without pectination), filiform in female. Setal comb absent from male abdominal sternite 3. Variation: The species shows some colour variation from light to dark beige-brown. Male genitalia: Uncus curved, apically acute, dorsally not extended, but setose. Gnathos absent. Valva basally broad with one row of long spines on sacculus, and one or two rows on ventral part; centre of valva with a few ventral tubercles. Membranous apex of valva narrower and shorter than costa of valva. The latter strongly sclerotized, apically curved, subapically with a large triangular extension, strongly setose. Juxta vase-shaped, slightly sclerotized. Saccus wide, rounded. Aedeagus slightly curved; everted vesica covered with evenly distributed tiny sclerotized dots. Female genitalia: Papillae anales thin (thicker in P. powelli), setose. Apophyses posteriores twice length of apophyses anteriores. Lamella antevaginalis widely v-shaped; ductus bursae funnel-shaped, much wider than in all other European congeners (clearly narrower in P. powelli); corpus bursae membranous, very long (5.7 mm), posteriorly tubular, very thin, anteriorly wide (corpus bursae considerably shorter in P. powelli); signum strongly sclerotized (less sclerotized in P. powelli) as a short transverse fold (0.2 mm). Distribution and abundance: West Mediterranean. Central to eastern Spain, in the north to the Pyrenees. – Local, usually in limited numbers. – Outside Europe with an isolated population in Morocco, Middle Atlas (Leraut 2009, regarding it as a separate subsp. gerini), apparently also in the High Atlas (see under genetic data).

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Phenology: Uni- or bivoltine. From August to October (Wehrli 1943), according to Redondo et al. (2009) from late July to early October. Larva found in July (Wehrli 1943). Biology: Larva possibly monophagous on Juniperus. Found on Juniperus oxycedrus, as stated on a specimen label (Wehrli 1943). Reared also on Juniperus (D. Stadie pers. comm.). Habitat: Xeromontane. Mountain slopes and open habitats with scattered Juniperus stands (D. Stadie pers. comm.). From 700 up to 1300 m (Redondo et al. 2009), in Morocco reputedly at 2070 m (Leraut 2009). Similar species: The most similar species is P. powelli. Wing patterns of both species are highly similar, but female genitalia (e.g. size of ductus bursae) allow a certain identification. Both Peribatodes aragonis and P. powelli may be confused with Menophra thuriferaria (Zerny, 1927) and M. harterti (Rothschild, 1912). See diagnostic characters of Ecleora solieraria (Rambur, 1834), M. thuriferaria and P. aragonis as outlined in Text-figs 169–171. Ekboarmia atlanticaria is also similar, see Text-fig. 161. Genetic data: BIN: BOLD:AAV8028 (n=2 from Spain). In Morocco with a separate BIN (BOLD:AAL8979; n=2; distance from P. a. aragonis 3.4%, however collected on High Atlas and not necessarily belonging to P. aragonis gerini, the latter described from the Middle Atlas). Nearest species: P. powelli (BOLD:AAL2705; n=8; distance 7.5%). Previously in literature, P. aragonis was treated as a subspecies of P. powelli, however, genetic data suggests species status of the former.

151. Peribatodes ilicaria (Geyer, 1833) Geometra ilicaria Geyer, 1833: Samml. Eur. Schmett. 5 Geometrae (1), pl. 111, figs 582, 583 (France: Provence). Syntype(s). Boarmia manuelaria Herrich-Schäffer, 1852: Syst. Bearb. Schmett. Eur. 6 (55): 73; ibidem (1853), 3: pl. 88, fig. 541 (non binominal) (France). Syntype(s). Junior synonym. Boarmia manuelaria maghrebica Le Cerf, 1923: Bull. Soc. ent. Fr. 1923 (15): 198 (Morocco: Tedders). Holotype ♂ (MNHN). Regarded as a valid subspecies (P. ilicaria maghrebica) by Leraut (2009). Boarmia ilicaria var. marinaria Cleu, 1928: Revue fr. Lépidopt. 4: 89, pl. 1, figs 1, 2 (France: Hautes-Alpes). Syntype(s). Boarmia jacobiaria Fernández, 1931: Eos, Madr. 7 (2): 218, pl. 2, fig. 14 (Spain: Prov. Cuenca, Uclés). Syntype(s). Regarded as a synonym of P. ilicaria by Redondo et al. (2009), here confirmed.

External characters and abdomen. Wingspan ♂ 27–35 mm, ♀ 30–38 mm. Wings from light beige to dark grey and dark brown, slightly irrorated with dark olive-green, exceptionally also ivory-yellow; markings dark brown to black. Forewing basal area and hindwing basal and medial areas slightly lighter than the rest of the wings. Forewing antemedial line at centre curved outwards; medial and postmedial lines deeply curved outwards, near dorsum deeply curved inwards; postmedial line dentate on veins towards costa; subterminal line absent; wavy line dentate, discontinuous, sometimes faint, partially edged white distally. Termen of forewing beige, apical spot light brown, subapically darker. Basal, antemedial and subterminal lines absent on hindwing; medial line faint or completely absent; post­medial line dentate on veins, at centre deeply curved outwards; wavy line irregularly dentate, sometimes faint. Discal spots tiny, but visible. Terminal line of all wings blackish, narrow, discontinuous, stronger between vein endings. Fringes brown, darker at vein endings. Under­side of wings lighter than upperside. Markings are weakly visible. Frons, collar, 434

thorax and abdomen concolorous with wings. Antennae dorsally concolorous with wings, bipectinate in male (except 6–7 apical segments without pectination), filiform in female. The setal comb present on male abdominal sternite 3. Variation: Highly variable, from light to dark brown. Specimens from Morocco have light yellow-brown wings without olive-green tones and are regarded as a different subspecies P. ilicaria maghrebica. For genetic variation see Genetic data. Male genitalia: Uncus elongated, apically acute, dorsally setose, not extended. Gnathos present but vestigial. Valva basally and apically broad with one row of long spines on sacculus, and two rows on ventral part; editum (cf. Steinmann & Zombori 1985 = subcostal plica of valva in Beljaev 2008) arch-shaped, elongated, strongly setose. Membranous apex of valva much broader (but shorter) than costa of valva. The latter strongly sclerotized, setose and apically covered with stout spines, apically with a triangular extension. Juxta vase-shaped, apically v-shaped, slightly sclerotized. Saccus broad. Aedeagus straight, same length as ­valva, apically covered with tiny spines; vesica membranous without any sclerotization. Female genitalia: Papillae anales ovally elongated, setose. Lamella antevaginalis well sclerotized, deeply concave. Ductus bursae shortly funnel-shaped, well sclerotized. Corpus bursae membranous, cylindrical, moderately long (3 mm). Signum strongly sclerotized as a short transverse fold (0.3 mm). Distribution and abundance: West Mediterranean element with Atlanto-Mediterranean distribution pattern, ranging from North Africa and Iberian peninsula to France, southern Belgium and some scattered records in Germany (Warnecke 1940; Schanowski & Ebert 2003; Gelbrecht & Müller 1987). Rather common, usually in limited numbers. Records in southern Denmark, northern Germany and in Poland may origin from temporary populations after migration out of the regular distribution area (Karsholt & Skou 2012). Additional

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records far from the known range of P. ilicaria (Greece, Bulgaria, Italy, Slovakia, Romania and Turkey) are considered erroneous due to misidentification (Karsholt & Skou 2012). – Outside Europe in North A ­ frica (Morocco, Algeria, Tunisia), here showing constantly lighter wing pattern and regarded as a different subspecies (P. ilicaria maghrebica). Phenology: Bi- or trivoltine, usually from early April to early October. In Portugal and southernmost Spain from mid-March to mid-November (P. Hale, M. Corley pers. comm.). Biology: Larva polyphagous on various trees. Found on Prunus spinosa (Warnecke 1940) and Pinus (Sønderup 1919) and Pinus sylvestris (Goméz de Aizpúrua 1997). Reared on Ligus­ trum sp. (Warnecke 1940). Recorded also on Quercus ilex (France: Culot 1919; Lhomme 1923–35), Malus communis, Juniperus oxycedrus, Cedrus (Goméz de Aizpúrua 1997), Hedera helix, Syringa, Clematis, and Rosa (Lhomme 1923–35). Habitat: Habitats with scattered trees and open forest, from sea-level up to 2000 m (­Redondo et al. 2009; P. Skou own data). Similar species: The species can easily be identified based on wing pattern and genitalia structure. The setal comb on the male third abdominal sternite is only present in P. ilicaria, P. umbraria and P. rhomboidaria. Genetic data: Genetically heterogeneous, diverging into two BINs: BOLD:AAE6311 (n=5 from France, Spain, Portugal); BOLD:AAB2951 (n=2 from France, Spain), distance between the two BINs 3.6%, requiring further taxonomic analysis. Nearest species: P. perversaria (7.9%).

Alcis Curtis, 1826 Alcis Curtis, 1826, Br. Ent. 3: 113. Type species: Phalaena repandata Linnaeus, 1758. Genus synonyms after Scoble (1999). Poecilalcis Warren, 1893, Proc. zool. Soc. Lond. 1893: 427. Type species: Cleora nigridorsaria Guenée, 1858. Dictyodes Wehrli, 1934, Int. ent. Z. 27: 509. Type species: Arichanna maculata Moore, 1868. Alcisca Wehrli, 1943, in Seitz, Gross-Schmett. Erde 4 (Suppl.): 511. Type species: Boarmia fredi Wehrli, 1941.

Diversity and distribution: Alcis contains approximately 140 species, which occur mainly in the Palaearctic (mostly East Palaearctic) and Oriental regions; one species in the Neotropical region. The genus needs to be analysed critically in a modern context. Only three species in Europe. External characters and abdomen: Large to medium-sized, usually greyish or brownish moths. Forewing upperside with fovea. Veins R1 and R2 arise separately from the cell. Anten­nae bipectinate in male, filiform in female. Hindlegs with two proximal and distal spurs, in male with pencil. Male genitalia: Uncus and gnathos fully developed. Valva rounded, costa sclerotized, apical part broader. Apical half of ventral margin membranous. Centre of valva with distinct sclerotized, digitiform flap, arising subcostally, usually directed ventrad, sometimes double or setose (Holloway 1994). Juxta with long posterior processes. Female genitalia: Ovipositor narrow, apophyses posteriores very long, apophyses anteriores shorter, ending spatulate. Corpus bursae tubular, hyaline, several times as long as wide. 436

Remarks: In multi-gene analysis Alcis is included in a large assemblage of genera including among others Sardocyrnia, Selidosema and Peribatodes (Murillo-Ramos et al. 2019). In the multi-­gene analysis of Jiang et al. (2017) in the same genetic cluster as Peribatodes.

152. Alcis repandata (Linnaeus, 1758) Phalaena (Geometra) repandata Linnaeus, 1758: Syst. Nat. (Ed. 10) 1: 524 (Europe). Syntype(s) (LSL). Phal[aena] (Geometra) consobrinaria Scriba, 1790: Beitr. Ins. Gesch. 1: 25, pl. 2, figs 2a, 2b (Italy). Syntype(s). Phalaena (Geometra) repandaria Borkhausen, 1794: Natur. eur. Schmett. 5: 154. Emendation of repandata Linnaeus. Phalaena bistortae Vallot, 1802: Concordance systématique, servant de table de matières à l’ouvrage de Réaumur, intitulé: Mémoires pour servir à l’histoire des insectes: 20, 192. Syntype(s). Deileptenia conversaria Hübner, 1809: Samml. Eur. Schmett. 5 Geometrae (1 & 2), pl. 62, fig. 321; ibidem. Geometrae (1), (1813), pl. 76, fig. 393 (Europe). Syntype(s). Geometra destrigaria Haworth, 1809: Lepid. Br. (2): 276 (United Kingdom). Syntype(s). Alcis muraria Curtis, 1826: Br. Ent. 3: folio 113 (Scotland: Isle of Arran). Syntype(s). Listed as synonym in Scoble (1999), validated at subspecies rank by Leraut (2009). Boarmia dealbata Turati & Verity, 1911: Boll. Soc. ent. ital. 43: 210 (Italy: Alpi Marittime, Valle del Gresso). Syntype(s). Boarmia repandata var. sodorensium Weir, 1881: Entomologist 14: 220, pl. 1, figs 12, 13 (Scotland: Outer Hebrides). Syntype(s). Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): albescens: Leraut (2009) (f.); brunneata: Heydemann (1938) (f.); contrasta: Lempke (1952) (f.); decorata: Leraut (2009) (f.); fusca: Lempke (1952) (f.); fuscomarginata: Lempke (1952) (f.); marginata: Lempke (1970) (f.); nigra: Tutt (1896) (ab.); nigricata: Fuchs (1875) (ab.); obscura: Lempke (1970) (f.); pallescens: Leraut (2009) (f.); pseudobastelbergeri: Folten (1942) (ab.); tangens: Lempke (1970) (f.).

External characters and abdomen: Wingspan 31–46 mm. Ground colour whitish grey to brownish, wings with irregularly distributed dark scales. Wavy line serrate, conspicuous especially in darker specimens. On forewing post- and antemedial lines and linear discal spot distinct. On hindwings teeth of wavy line on M2 often not reaching terminal line. Fringes with longitudinal line, chequered. Antennae partially bipectinate to 4/5 of length in male, filiform in female. Frons, collar, thorax, and abdomen concolorous with wings, however first (anterior) abdominal segment silver grey, following segments with double row of dark spots. Hindtibia of both sexes with 2+2 spurs, in male with hair pencil. Variation: Variable species. Ground colour varies from pale whitish grey to grey or brown. Extreme form with dark medial area (“ab. conversaria”). The dark spot between subterminal and wavy line is variable, on average less distinct than in A. deversata, sometimes missing. Melanic forms with black ground colour (in older collections specimens fading to dark brown), often scattered with remainder of ground colour of nominotypical form). Ante­ medial line, postmedial line, discal spot and wavy line of this form usually clearly visible. First abdominal segment also black. Subsp. muraria from moorland in Scotland and Northern Ireland with ground colour whitish grey and often weakly marked (see Skinner 2009). Subsp. sodorensium from the Outer Hebrides (Isle of Lewis) is a small lead-grey coloured race (see Skinner 2009). 437

Male genitalia: Uncus ending with indicated tip. Gnathos arms wide, apex rounded. ­Valva rounded, costa sclerotized, broader towards apex. At centre a pair of ventrally directed digitiform processes separated by a distance of approximately length of process (see A.  dever­sata). Above ventral process a small, setose, curved lamella (larger in A. deversata). Separation from A. deversata is easy by the shape of juxta: juxta arms of A. repandata long, pointed, fused for over 1/4 of the juxta length from its anterior end; saccus rounded. Aedeagus with long rectangular hook and a sclerite, tapering towards tip. Very similar to that of A. deversata. Female genitalia: Very similar to those of A. deversata. Papillae anales membranous, slightly setose. Apophyses posteriores long (on average 5.0 mm), straight. Apophyses anteriores shorter, on average 1.87 mm, ending spatulate (n=3). Lamella postvaginalis similar to that of A. deversata, but less sclerotized. Ductus bursae partly sclerotized forming ring with anteriorly directed tongue. Corpus bursae very long, hyaline with two slightly sclerotized blotches, one quarter moon-shaped, the other oval, both granulate. Distribution and abundance: European-Turanian. In Europe widely distributed from the Urals to Ireland and northern Portugal and from central Fennoscandia to the northern Iberian peninsula, France including Corsica, Italy including Sicily and the Balkan peninsula to northern Greece. The distribution map in Leraut (2009) shows a too large area in the Iberian peninsula, Greece and in northern Europe. – Common or very common, now and then occurring in numbers. – Outside Europe from Turkey, Georgia, Armenia, Azerbaijan, to northern Kazakhstan and south-western Siberia.

Phenology: Univoltine. In southern Scandinavia as well as in south-western Germany from mid-June to mid-August. In Spain from April to mid-August (Redondo et al. 2009). Under good conditions to the south in Europe a partial second generation in September and October. Larva overwintering from August to June. Adults readily attracted to light and sugar bait. 438

Biology: Larva polyphagous. Found on Pteridium aquilinum, Dryopteris, Abies alba, Picea excelsa, Pinus, Populus tremula, Salix cinerea, S. caprea, Corylus avellana, Betula pendula, Quercus, Urtica dioica, Rumex obtusifolius, Silene dioica, Clematis vitalba, Berberis thunbergii, Sorbus aucuparia, Crataegus, Rubus, Agrimonia eupatoria, Sanguisorba officinalis, Rosa canina, Prunus padus, P. spinosa, Cytisus scoparius, Genista tinctoria, Vicia, Acer campestre, Impatiens noli-tangere, Frangula alnus, Tilia cordata, Vaccinium myrtillus, V. uliginosum, Calluna vulgaris, Galium, Galeopsis tetrahit, Sambucus nigra, Lonicera xylosteum, Senecio fuchsii (Germany: Ebert 2003). Found on Corylus avellana also in north-western Italy (King & González-Estébanez 2015). In wintertime reared with Hedera helix (Bergmann 1955). Pupation in a loose cocoon among plant litter (Porter 2010). Habitat: Hygrophilous to mesophilous, silvicolous. Occurring in a wide variety of habitat types, such as coniferous, deciduous and mixed forests, open forest areas, scrubby places, tall herb communities, bog-land, ruderal places, gardens, commons. From 0 up to 1500 m above sea-level, up to 1700 m in the Alps and up to 1400 m in the central Apennine mountains in Italy. Similar species: Alcis deversata (see no. 153) on average with more contrasted wing pattern. Sometimes separation from A. deversata is only possible through examination of the genitalia. Differences in genitalia, see above. Genetic data: In Europe with two deeply diverging and often sympatric genetic clusters (separate BINs) at a minimum distance of 4.5%: BOLD:AAA8484 (n=48 from Finland, United Kingdom, Germany, Austria, Italy, Croatia, Macedonia, Bulgaria) and BOLD:AAA8482 (n=79 from United Kingdom, Germany, Austria, France, Italy, in the east to Mongolia, Far East Russia and Far East China). The latter BIN genetically identical with specimens from Altai mountains, currently being identified as A. extinctaria (Eversmann, 1851), described from Russia: Irkutsk. Either synonymy or genetic introgression may explain this result. Both BINs genetically homogeneous. Nearest species: A. deversata (4.8% from BOLD:AAA8482, 5.5% from BOLD:AAA8484).

153. Alcis deversata (Staudinger, 1892) Boarmia repandata deversata Staudinger, 1892: Dt. ent. Z. Iris 5: 377 (Russia: Altai; Mongolia: Kentei). Syntype(s) (MNHU). ‡ Boarmia repandata var./ab. maculata Staudinger, 1901: in Staudinger & Rebel, Cat. Lepid. palaearct. Faunengeb. 1: 341 (Dauria; Amur; Ussuri; Switzerland; Altai; Siberia). Syntype(s). Junior secondary homonym of maculata Moore. Boarmia repandata bastelbergeri Hirschke, 1908: Jber. wien. ent. Ver. 18: 107, pl. 1, figs 2, 3; pl. 2, figs 2, 2a-c (Austria: Steiermark). Syntypes ♂♀ (ZFMK). Boarmia kaibatonis Matsumura, 1931: 6000 illustr. Insects Japan-Empire: 876, figured (Japan?). Syntype(s), including ♂. Boarmia maculata f. sachalinensis Matsumura, 1931: 6000 illustr. Insects Japan-Empire: 877 (Sachalin). Syntype(s). Valid at subspecific rank (Scoble 1999). Alcis bastelbergeri hasegawai Inoue, 1942: Trans. Kansai ent. Soc. 12 (1): 20, pl. 4, figs 15, 16 (Japan: Iwate Prefecture, Morioka). Syntypes 2♂1♀. Alcis conversariodes Inoue, 1942: Trans. Kansai ent. Soc. 12 (1): 23 (Japan: Iwate Prefecture, Morioka). Types ♂. Boarmia maculata sinimaculata Wehrli, 1943: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 504 (China: Shensi, Tapai-shan, Tsinling). Syntype(s) (ZFMK).

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Boarmia repandata kukunorensis Albers, 1949: Mitt. münch. ent. Ges. 35–9: 256, pl. 5, fig. 1 (China: Kuku-Nor). Holotype. Valid at subspecific rank of (Scoble 1999). Boarmia (Cleora) repandata cioccolatina Bryk, 1949: Ark. Zool. 41A (1): 204, pl. 6, fig. 5 (Korea: Motojondo). Holotype ♂ (NHRS).

External characters and abdomen: Wingspan 31–40 mm. Ground colour whitish grey to brownish, pattern of European specimens more homogeneous, i.e. with less irregularly distributed dark scales than in A. repandata. The dark spot between subterminal and wavy line is more distinct than in A. repandata. Basal area of hindwings often darker, medial line conspicuous, nearly straight, diffuse (in A. repandata not present or weak). Nominotypical form has a dark medial area. Fringes, antennae, frons, collar, thorax, and hindtibia as in A. repandata. The anterior abdominal segment silver-grey, the segments 9 to 8 (7) dorsally with two darker spots on each segment, which can also form a ring. Sometimes separation from A. repandata requires examination of the genitalia. Variation: Variable species. Ground colour varies from pale whitish grey to grey or brown. Extreme forms as in Alcis repandata. Male genitalia: Uncus, gnathos and valvae rounded. Costa of valvae sclerotized, broader towards apex. Pair of ventrally directed digitiform processes (at centre of valva) separated

Text-figs 198–199. Diagnostic characters (indicated) of similar European Alcis species. Text-fig. 198. Alcis repandata (Linnaeus, 1758), Germany, Brandenburg, Oder-Spree, 16.vi.2005. Textfig. 199. Alcis deversata (­Staudinger, 1892), Slovakia: Trenčin, Mníchova-Lehota, 02.v.1986 (all coll. B. Müller).

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by a shorter distance than length of processes (see A. repandata). Above ventral process a setose, curved lamella (smaller in A. repandata). Separation from A. repandata is easy by the shape of the juxta: juxta arms fused for 1/3 of juxta length, reminiscent of the head of a horned bull. Saccus rounded. Aedeagus very similar to that of A. repandata. Female genitalia: Very similar to those of A. repandata. Papillae anales membranous, slightly setose. Apophyses posteriores long (on average 4.0 mm), straight. Apophyses anteriores shorter, on average 1.73 mm, ending spatulate (n=3). Both shorter than in A. repandata. Lamella postvaginalis similar to that of A. repandata, but more strongly sclerotized. More characteristics see under A. repandata. Distribution and abundance: Eurasiatic. In Europe from the Ural mountains to eastern France, and from Latvia, Poland and northern Germany to the Alps and the northern Balkan peninsula. In Germany expanding towards north-east. The distribution map in Leraut (2009) shows a too large area in northern Germany, in Poland and in Italy. – Local, usually in single specimens. – Outside Europe from the Urals through southern Siberia, Altai, Mongolia, northern China (subsp. kukunorensis) and Korea to Far East Russia, Sakhalin and Japan. According to Viidalepp (1996) the eastern populations should be subsp. sachalinensis, but it remains unclear how far this subspecies is distributed to the west.

Phenology: Univoltine. From mid-July to late August in south-western Germany (Ebert 2003). Caterpillar found in September (Kerschbaum & Pöll 2010). Hibernation as larva (observation in rearing). Adults active at night, both sexes attracted to light. Nectar-feeding observed on Knautia, Eupatorium cannabinum and Senecio (Ebert 2003). Biology: Larva polyphagous. Found on Rosa (wild species) (D. Fritsch pers. comm.), Cle­ ma­tis vitalba and Rubus ideaus (Ebert 2003). Reared on Lonicera xylosteum, L. ­tatarica, 441

Rubus caesius (Ebert 2003), Rosa and Salix x sepulcralis (Lenek 1961). Pupation in the ground (T. Drechsel pers. comm.). Habitat: Silvicolous. In submontane and montane forests, deciduous as well as mixed forests and edges of coniferous forests. In Upper Austria from 200 up to 1000 m, most common between 700 and 800 m (Kerschbaum & Pöll 2010). Similar species: Alcis repandata (see no. 152). Genetic data: BIN: BOLD:AAC1009 (n=29 from Finland, Germany, Austria, Slovakia, in the east to Mongolia and Russian Far East), genetically rather homogeneous. From north-western to south-western China BIN-sharing populations at 0.6% distance (n=22), probably conspecific. Nearest species: A. repandata (4.8%). Remarks: In older publications named as Alcis maculata (Staudinger, 1892) or A. bastel­ bergeri (Hirschke, 1908). Because of shifting Arichanna maculata (Moore, 1868) to the genus Alcis by Inoue (1987), Alcis maculata (Staudinger, 1892) became a secondary homonym, and Alcis deversata (Staudinger, 1892) must be used as the valid name (see Mironov et al. 2008).

154. Alcis jubata (Thunberg, 1788) Phalaena jubata Thunberg, 1788: D. D. Mus. nat. Acad. Upsal. 6: 75, figured (Germany). Lecto­ type ♀ (UZIU). Phalaena dilatata Hufnagel, 1767: Berlin Mag. 4 (5): 610 (Germany: Berlin). Syntype(s). Unused senior synonym. Geometra glabraria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1), pl. 31, fig. 162 (­Europe). Syntype(s). Geometra teneraria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1), pl. 67, fig. 351 (­Europe). Syntype(s). Deileptenia plancaria Hübner, 1825: Verz. bekannter Schmett. Unnecessary replacement name for glabraria Hübner (Scoble 1999). Boarmia semiothisata Lenz, 1917: Mitt. münch. ent. Ges. 8 (5–10): 47 (France: Corsica, Vizzavona, Col di Sorba). Holotype. Alcis jubata melanonota Prout, 1930: Novit. zool. 35: 330 (Japan: Japanese Alps). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): obcura: Fuchs (1875) (ab.).

External characters and abdomen: Wingspan ♂ 20–25 mm, ♀ 23–27 mm. All wings white with irregularly distributed dark scales and pale grey terminal line. Forewings with large black discal spots, medial lines at costa blackish and thickened, terminal area darker with whitish wavy line. Hindwings with smaller discal spot, postmedial line distinct. Fringes che­ quered whitish and grey. Frons and collar whitish with single dark scales. Hindtibia of both sexes with 2+2 spurs. Abdomen with two rows of black dots. Male genitalia: Uncus long, tapered. Gnathos arms fused ventrally, elongated. Costa of valva sclerotized, broader towards apex and base, medially narrow. Centre of valva with a sclerite bearing small, ventrally directed projections of different length and size. Juxta arms fused over 3/4 of juxta length. Saccus rounded. Aedeagus with one sclerotized tooth. Female genitalia: Papillae anales membranous, slightly setose. Apophyses posteriores straight, length on average 1.45 mm. Apophyses anteriores shorter, on average 0.86 mm. Corpus bursae very long (length/diameter ≈ 6.7), hyaline, weakly striated, without signum. 442

Ductus bursae wide, short, sclerotized, smaller than corpus bursae. Lamella postvaginalis and lamella antevaginalis membranous. Distribution and abundance: Eurasiatic. In Europe from the Ural mountains to the British Isles and from central or northern Fennoscandia to the Pyrenees, Corsica, the Alps and the Balkan peninsula except for Greece. – Occurring locally, uncommon, usually found as single or few specimens. The species has declined in the past hundred years, due to decline of the food-plants which have disappeared in many regions (e.g. most of Germany), probably due to acid rain. – Outside Europe eastwards from the Urals through southern Siberia and Mongolia to Far East Russia. In Japan, southern Kuriles Islands and Sakhalin replaced by subsp. melanolota.

Phenology: Univoltine. In Fennoscandia as well in south-western Germany from late June to mid-August. Larva from August, hibernating to June. Adults active at night, readily attracted to light and sugar bait. Biology: Larva oligophagous. Found on Usnea barbata, Evernia prunastri (Prochnow 1905), recorded on Usnea dasypoga, Alectoria spec. (Skou 1984). Usnea barbata, U. dasypoga (Usnaeceae), Alectoria, Jungermannia (Scoble 1999), the last mentioned, however, drawn into doubt by M. Corley (pers. comm.). According to Ebert (2003), the genus Usnea is the main food-plant. Hibernation between branches of food-plant (Prochnow 1905). Habitat: Silvicolous. In older forests, mainly but not solely with conifers, in areas with relatively high air humidity and with abundant occurrence of the lichen food-plants. Locally abundant in oak woods in high rainfall areas in western Britain (M. Corley pers. comm.). In Fennoscandia from sea-level, in south-western Germany mainly around 900–1000 m

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(Ebert 2003), in the Alps from 300 up to 1400 m (Kerschbaum & Pöll 2010) and in the Pyrenees from 900 up to 1800 m (Redondo et al. 2009). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAD8767 (n=11 from Finland, Austria, Russia). Genetically hetero­geneous, maximum variation 2.5%. Genetically very isolated, nearest species: Alcis variegata (Moore, 1888) from India to Taiwan and Indonesia (9.3%).

Deileptenia Hübner, 1825 Deileptenia Hübner, [1825] 1816, Verz. bekannter Schmett.: 316. Type species: [Phalaena] ribeata Clerck, 1759. Genus synonym after Fletcher (1979). Diloptenia Agassiz, 1847, Nomencl. zool. (Index univl.): 118, 124.

Diversity and distribution: The genus Deileptenia contains four species in the Palaearctic and Oriental regions, only one in Europe. External characters and abdomen: Larger moths, wingspan 32–45 mm. Typical boarmiine habitus. Wings brownish grey with whitish irroration, ante- and postmedial lines distinct. Medial lines on forewing forming four thicker spots on costa. Medial area lighter. Antennae partially bipectinate in male, filiform in female. Fovea absent. Male and female genitalia: For the single European species see description below. Remarks: In multi-gene analysis Deileptenia is included in an assemblage of non-European genera, which is sister to a lineage that includes among others Alcis, Sardocyrnia, Selido­ sema and Peribatodes (Murillo-Ramos et al. 2019). In the multi-gene analysis of Jiang et al. (2017) in the large genetic cluster including among others Alcis and Peribatodes.

155. Deileptenia ribeata (Clerck, 1759) Phalaena ribeata Clerck, 1759: Icones Insectorum Rariorum 1: pl. 6, fig. 5 (Sweden). Syntype(s). Phalaena (Geometra) ribearia Linnaeus, 1761: Fauna Suecica (Edn 2): 326. Emendation of ribeata Clerck, 1759. Geometra abietaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s) (lost). Alcis sericearia Curtis, 1826: Br. Ent. 3: folio 113, pl. 113 (England: Hampshire, New Forest, near Lyndhurst). Syntype(s). Boarmia (Cleora) daisetsuzana Matsumura, 1927: Insecta matsum. 1 (4): 185 (Japan: Hokkaido, Mt Daisetsu, Kurodake). Holotype ♀. Boarmia subdisplicens Bryk, 1949: Ark. Zool. 41A (1): 206, pl. 7, fig. 11 (Korea: Shuotsu). Holotype ♂ (NHRS). Boarmia ribeata coreiphila Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 497, pl. 43i (North Korea: Poktussan, 2500 m). Syntype(s). Unavailable names (infrasubspecific): illineata: Lempke (1952) (f.); nigra: Cockayne (1948) (ab.); sericearia: Curtis (1826) (ab.).

External characters and abdomen: Wingspan ♂ 32–40 mm, ♀ 35–41 mm. Ground colour of forewings brownish grey, East Asian populations more whitish grey with dark irroration. Transverse lines forming four thicker spots on costa. Postmedial line with right angle near costa. Medial and postmedial lines approaching or anastomosing at inner margin. Terminal line blackish, discontinuous. Hindwings with serrate postmedial line, paler. Fringes with 444

indistinct longitudinal line, chequered beige and dark brown, beige segments of fringes correspond with black dots of terminal line. Abdomen, thorax, frons and collar concolorous with ground colour, frons smoothly scaled. Antennae partially bipectinate to 6/7 of length (about 12 flagellomeres free of branches) in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Medial line often ending before reaching the inner margin. Density of irroration varies. Ground colour of some Asian populations ash-grey to white-grey. Male genitalia: Uncus hooked, beak-shaped. Gnathos arms fused, spatulate. Valva rounded, costa sclerotized, setose. Ventrally with sclerotized ridge, bearing one larger and one smaller process with dorsally directed group of setae. Ventral margin granulate. Juxta plateshaped, without bifurcation. Saccus broad, rounded. Aedeagus short, thick, subapically with acute tooth, vesica with long, large cornutus. Female genitalia: Papillae anales slightly sclerotized, slightly setose, elongate. Apophyses posteriores very long, straight. Apophyses anteriores shorter, ending spatulate. Lamella antevaginalis sclerotized, broad, curved. Lamella postvaginalis narrow, spatulate, sclerotized. Ductus bursae with elongate reticulation, posterior part wider. Corpus bursae long. Anterior part membranous and striated sac with signum, posterior part sclerotized, 4/5 striated. Distribution and abundance: Eurasiatic. In Europe from the Urals to Ireland and from central Fennoscandia to the Pyrenees, southern France, northern Italy and the northern part of the Balkan peninsula. Erroneously recorded for Greece in Leraut (2009), moreover showing strongly exaggerated distributions in northern Europe and Italy, and too limited distribution in Ireland. – In Europe somewhat local, sometimes rather abundant in its localities. – Outside Europe through southern Siberia to Far East Russia, Sakhalin, the Kuriles, Korea and Japan.

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Phenology: Univoltine. In southern Scandinavia from late June to early August, in south-western Germany until late August (Ebert 2003), in northern Italy from mid-June to late August (Flamigni et al. 2016) and in Spain from early July to early August (Redondo et al. 2009). Larva from late July to late June, overwintering. Biology: Larva polyphagous. Found on Abies alba, Picea excelsa, Carpinus betulus, Betula pendula, Quercus, Prunus padus, P. spinosa, Vaccinium uliginosum and Lonicera xylos­ teum (Ebert 2003). Also recorded on Larix, Pinus, Pseudotsuga, Taxus, Salix, Alnus, Fagus, Ribes, Rubus, Rhododendron and Syringa (Flamigni et al. 2016). Reared on Coronilla varia (Flamigni et al. 2016). Pupation in a light silken web on the ground. Habitat: Silvicolous. In conifer forests with a mixture of deciduous trees. In northern E ­ urope from sea-level, in the Pyrenees from 600 up to 1400 m (Redondo et al. 2009) and in north-eastern Italy up to more than 1500 m (Flamigni et al. 2016). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC3800 (n=33 from Finland, United Kingdom, Germany, Austria, Italy, Russia (Altai), northern China, Japan). Genetically homogeneous in Europe, but populations from Japan diverging by 1.1%. Nearest species: Xandrames dholaria Moore, 1868 from India (5.7%). Nearest species in Europe: Calamodes melanaria (= subscudularia) from Italy (6.3%). Nearest congener: Asian D. mandshuriaria (Bremer, 1864) (8.9%).

Paraboarmia Krampl, 1994 Paraboarmia Krampl, 1994, Folia Heydrovskyana 2 (7): 89. Type species: Boarmia viertlii ­Bohatsch, 1883. There are no genus synonyms.

Diversity and distribution: Paraboarmia is a monotypic genus, the only species occurs in the West Palaearctic region. External characters, abdomen and Genitalia: See species description below. Remarks: Paraboarmia has not been included in a multi-gene analysis so far. Tentative ­position, requiring further research. Phthonandria combination (as in Viidalepp 1996) not supported by structural morphology (our observation) nor by molecules: type species of Phthonandria included in multi-gene analysis and grouping in the Gnophini (Murillo-Ramos et al. 2019).

156. Paraboarmia viertlii (Bohatsch, 1883) Boarmia viertlii Bohatsch, 1883: Wien. ent. Ztg. 2: 111, figured (Hungary). Syntypes syntype at ZSM examined and DNA barcoded).

♂♀ (1♂

External characters and abdomen: Wingspan 25−28 mm. Ground colour of wings whitish grey, irrorated with brown scales. Postmedial, medial and antemedial lines on forewing black, antemedial line absent from hindwing. On forewing, postmedial line bent near costa and before reaching inner margin. Whitish wavy line on forewing usually distinct, on hindwing often indistinct and diffuse. Fringes chequered whitish and brownish grey, with weak longitudinal line. Black discal spots present on all wings, also on underside. Frons, vertex, collar, tegula, and thorax concolorous with ground colour, irrorated more or less with dark scales. Antennae bipectinate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. 446

Variation: Darker or lighter due to varying irroration with dark scales. Male genitalia: Uncus and gnathos very large. Both on lateral margins concave, ­apically wide. Uncus nearly flat ending, gnathos more round. Valva setose, wide, simple. Apex rounded, costa and base weakly sclerotized. In the centre a small short ridge with group of short setae. Juxta developed as a triangular sclerite, tip round. Aedeagus cylindrical, straight. Vesica with three cornuti of different lengths and a weakly sclerotized granulated part. Female genitalia: Papillae anales membranous, soft, slightly setose. Apophyses posteriores very long (on average 3.1 mm), apophyses anteriores shorter (on average 1.4 mm), both spatulate at tip. Ductus bursae short, sclerotized. Corpus bursae membranous, slightly striated, anteriorly wider, rounded. Signum medium-sized, stellate. Lamella antevaginalis alate, with two small and narrow, tapered ‘wings’. Lamella postvaginalis a sclerotized triangular plate, anteriorly bifurcate. Distribution and abundance: European-West Asiatic. Scattered occurrence from south-­eastern France, Italy including Sicily and southern central Europe to the Balkan peninsula and Lesbos. – In Europe very local and rather rare, usually found in single specimens. – Outside Europe only in Turkey and Georgia.

Phenology: Univoltine. From early June to early August. Larvae from August to May, overwintering (Fajčik 2003). Adults active at night, attracted to light. Biology: Larva monophagous. Found on Quercus pubescens (M. Leipnitz pers. comm.). Recorded on Quercus (Fajčik 2003). Pupation among moss or in the ground (M. Leipnitz pers. comm.).

447

Habitat: Xerothermophilous. In warm places with Quercus. From close to sea-level up to 950 m in southern Italy. In Turkey up to 1500 m. Similar species: Differing from similar boarmiine moths by the smaller size and broader wings. Genetic data: BIN not yet assigned (n=2, 458bp and 408bp, from Hungary and Turkey, including a syntype). Hungarian and Turkish specimens exactly barcode-sharing (sequence overlap: 408bp). Nearest species: North American Hulstina wrightiaria (Hulst, 1888) at a very short distance of 3.4%, suggesting true phylogenetic relationships, possibly even synonymy at genus level.

Adactylotis Hübner, 1823 Adactylotis Hübner, [1823] 1816, Verz. bekannter Schmett.: 304. Type species: Geometra gesti­ cularia Hübner, 1817. Genus synonyms after Scoble (1999). Adactylotus Agassiz, 1847, Nomencl. zool. (Index univl.): 7, [Emendation of Adactylotis]

Diversity Europe.

and distribution:

Two species occurring in the West Palaearctic region, including

External characters and abdomen: Medium-sized, wingspan 22–34 mm. Wings pale sandy yellowish brown with medial and postmedial lines. Discal spot on hindwings and fovea on forewing base present. Veins R1 and R2 shortly stalked, arising from the cell. Antennae partially bipectinate in male, filiform in female. Hindtibia of both sexes with 2 + 2 spurs. Male genitalia: Uncus and gnathos broad. Uncus weakly setose, laterally concave, ending in a bent triangular tip. Gnathos arms ventrally fused, broadly rounded. Valvae undivided, apex rounded, weakly setose. Costa sclerotized, setose. Ventral margin sclerotized over half valva length, at centre with oval lobe, covered with inwardly directed spines. Centre of ­valva with a group of long setae. Juxta on both ends rounded, at centre weakly constricted. Saccus wide, rounded. Female genitalia: Papillae anales distinctly elongated, setose. Apophyses posteriores very long, apophyses anteriores shorter, both spatulate at tip. Lamella antevaginalis membranous. Lamella postvaginalis sclerotized. Ductus bursae narrow, posterior part sclerotized, anterior part membranous. Corpus bursae long, membranous. Posterior part narrow, with dilatation. Anterior part large, round. Signum densely stellate. Remarks: Adactylotis has not been included in a multi-gene analysis so far. Position tentative, based on similarity of male genitalia morphology between Adactylotis, Pseudocoremia and Deileptenia.

157. Adactylotis gesticularia (Hübner, 1817) Adactylotis gesticularia Hübner, 1817: Samml. Eur. Schmett. 5 Geometrae (1): pl. 92, figs 472, 473, (Europe). Syntype(s). Cabera graellsiaria Feisthamel, 1835: Annls Soc. ent. Fr. 4 (1): 134, pl 1A, fig. 3 (Mont-Joui). Syntype(s). Cabera inquinataria Boisduval, 1840: Genera Index meth. eur. Lepid.: 219 (Spain). Syntype(s). Itame ungemachi Lucas, 1933: Bull. Soc. ent. Fr. 38 (13): 196 (Morocco: Harcha (Zemmour); Aïn Leuh). Syntypes. Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): unistrigata: Leraut (2009) (f.).

448

External characters and abdomen: Wingspan 23–36 mm. Fore- and hindwings pale sandy yellowish brown, light grey speckled, speckling often denser in the terminal area. Wings appear homogeneously coloured. Postmedial line distinct, on veins sometimes more strongly dotted, on forewing angled close to costa, on hindwing curved; medial line rather straight, thicker. Antemedial line on forewings often indistinct or missing. Discal spot on hindwings small, grey, sometimes difficult to see, no discal spot on forewing. Frons, collar, thorax, abdomen and fringes concolorous with wings, however, fringes on forewings towards the apex often darkened. Antennae bipectinate in male, filiform in female, each segment half pale and half brownish. Legs pale brown, irregularly mixed with brownish scales, partially forming rings. Variation: Very little. Ground colour of the wings ranges from light beige to whitish grey. Degree of speckling varies. Subsp. ungemachi from Morocco with postmedial line reduced to dots on the veins. Forewing underside dark grey, costal area distinctly lighter. Male genitalia: Very similar to A. contaminaria (see genus description). However, ventral margin equally sclerotized, in the centre a smaller group of spines than in A. contaminaria. Aedeagus cylindrical, apex with outwards bent teeth (somewhat shorter than in A. contaminaria), at the opposite side a small sclerotized, outwards directed tip. Female genitalia: Very similar to A. contaminaria (see genus description). Posterior part of corpus bursae with one weakly sclerotized dilatation. Distribution and abundance: West Mediterranean. In Europe restricted to southernmost France and Iberian peninsula. – Rather common, sometimes found in numbers, in southern France local. – Outside Europe only in Morocco (subsp. ungemachi).

449

Phenology: Bivoltine. In the Iberian peninsula from early April to late September, depending on latitude and altitude; in southern Spain from March to July and in September and October (P. Hale pers. comm.); in southern France in two generations from April to May and from August to September (Robineau et al. 2007). Caterpillar thoughout the year, hiber­nating (Gómez de Aizpúrua 1989); one adult larva in southern Portugal in early April (D. Fritsch pers. comm.). Active at night, readily attracted to light. Biology: Larva monophagous on Quercus. Found on Quercus ilex and Q. coccifera (D. Fritsch pers. comm.). Recorded on Quercus, mainly Q. ilex, Q. coccifera, Q. pyrenaica, Q. faginea (Redondo et al. 2009) and Q. robur (Robineau et al. 2007). Habitat: Basically silvicolous. In or near forests and other places with the food-plant. Often occurring in places where food-plants only appear as low-growing shrubbery. Up to at least 1500 m altitude. Genetic data: BIN: BOLD:AAJ5596 (n=4 from Spain). Genetically homogeneous. Nearest species: A. contaminaria (3.6%). Remarks: The taxon inquinataria was described by Boisduval because of different line-pattern. But this feature is within the normal variation.

158. Adactylotis contaminaria (Hübner, 1813) Geometra contaminaria Hübner, 1813: Samml. Eur. Schmett. 5 Geometrae (1): pl. 68, fig 356, (Europe). Syntype(s). Phalaena numerata Fabricius, 1798: Syst. Ent. (Suppl.): 455 (Paris). Syntype(s) lost. Unavailable names (infrasubspecific): purata: Leraut (2009) (f.).

External characters and abdomen: Wingspan 24–36 mm. Ground colour of wings sandy yellowish brown, unevenly reddish brown speckled, compressed in forewing centre into a poorly defined spot. Forewings with antemedial, medial and postmedial lines, postmedial line curved inwards near costa, outwards near inner margin, antemedial line thin, medial line thicker, both curved. Hindwings with curved medial and postmedial line. All transverse lines reddish brown. Discal spots on hindwings indistinct, grey. Discal spots on forewings on medial line. Fringes concolorous with wings, chequered with grey, on forewings dark near costa. Frons, collar, thorax, legs and abdomen concolorous with wings, with reddish brown scales. Antennae bipectinate in male, filiform in female, each segment half pale and half reddish brown. Variation: Very little. Male genitalia: Very similar to those of A. gesticularia, ventral margin of valva equally sclerotized (see genus description), however, lobe with a larger group of spines than in A. contaminaria. Aedeagus cylindrical, apex with outwards bent teeth (longer than in A. gesticularia), at the opposite side a small sclerotized, outwards directed tip. Female genitalia: Very similar to those of A. gesticularia (see genus description). Posterior part of corpus bursae with two dilatations, the larger one sclerotized, weakly striated with very small teeth. The smaller one round with a very small tooth. Distribution and abundance: South-west European. In northern Spain, France, and Italy including Sicily. Erroneously recorded for Belgium and Switzerland in Fauna Europaea (­Hausmann et al. 2004; 2011a). Erroneously recorded for Portugal, central and southern Spain by Leraut (2009). – Rather local, usually occurring in limited numbers. 450

Phenology: Bivoltine. From mid-April to late June and from July to September (Flamigni et al. 2016). Generations possibly overlapping. Penultimate instar larva in early October in north-eastern Spain (D. Fritsch pers. comm.). Overwintering as pupa (D. Fritsch pers. comm.). Biology: Larva monophagous on Quercus. Found on Quercus robur (D.  Fritsch pers. comm.). Recorded on Q. robur and other Quercus species (Flamigni et al. 2016). Active at night, attracted to light. Habitat: Silvicolous. In open woodland. From close to sea-level, up to 1900 m in Italy (Flamigni et al. 2016). Genetic data: BIN: BOLD:AAC9850 (n=10 from France, Italy, including Sicily). Genetically homogeneous. Nearest species: A. gesticularia (3.6%).

Pseudocoremia Butler, 1877 Pseudocoremia Butler, 1877, Proc. zool. Soc. Lond. 1877: 394. Type species: Selidosema fragosata Felder & Rogenhofer, 1875. [Junior synonym of Pseudocoremia (Larentia) productata (Walker, 1862)]. Zylobara Butler, 1879, Cistula ent. 2: 498. Type species: Rhyparia fenerata Felder & Rogenhofer, 1875.

Diversity and distribution: About 30 species (Scoble 1999; Stephens & Gibbs 2003; ­Stephens et al. 2007) in New Zealand including the Chathams, and Norfolk Island. One species in Europe, as a result of anthropogenic transport (James 2008). External characters and abdomen: Medium-sized, wingspan 25–30 mm, few species with brachypterous females. Wings very variable, even within single species, dominant colours

451

of forewings often being various shades of brown, yellowish brown and grey. Hindwings often with weak pattern, paler, greyish, yellowish or whitish. Forewing postmedial line often most distinct, deeply concave before inner margin. Forewing medial area can be distinctly darker than rest of wing. Male antennae bipectinate with long pectinations, female antennae filiform. Fore- and midlegs striped in many species. Hindtibia of both sexes with 2+2 spurs. Male 3rd sternite with setal comb (only P. suavis examined), 8th segment elongated (only P. suavis examined). Male genitalia: Uncus triangular, apex weakly bifid. Socii absent. Gnathos arms often fused ventrally, medial element upturned. Base of valva costa sclerotized, apical portion setose. Ventral margin of costa sclerotized, apical portion plate- or hook-shaped. Medial area of valva rather membranous, with distinct setose ampulla at halfway. Juxta base long, apex often bifid. Posterior arms of juxta absent. Saccus often weakly upturned. Vesica with or without cornuti. Female genitalia: Terminal segments 9+10 often elongated. Apophyses posteriores very long. Lamella antevaginalis and lamella postvaginalis often membranous or weakly sclerotized. Ductus bursae short. Corpus bursae often greatly elongated, narrow, anterior part weakly expanded. Signum absent or small, weakly stellate. Biology, habitat, phenology: P. suavis polyphagous on a wide range of trees and shrubs in New Zealand, including Nothofagus spp, Podocarpus spp., and it is a serious defoliator in some exotic forest plantations, particularly with Pinus radiata and Pseudotsuga menziesii (Scott 1984; Berndt et al. 2004). It is found commonly in New Zealand on exotic Ulex europaeus, which is very common also in its European locality in Cornwall in Great Britain (James 2008). P. suavis inhabits lowlands up to 900 m in New Zealand. Several Pseudocoremia species have been found in New Zealand in the low alpine zone and the larvae have been found to feed on Hebe sp. (Scrophulariaceae), including Hebe odora, H. hectorii, H. cockayneana and H. crawii (Fox 1973; Stephens et al. 2007). The Chatham Island endemic P. ombrodes is polyphagous, recorded feeding on a variety of trees and shrubs (Holloway 1977; Dugdale & Emberson 1996). Adults of several species fly between mid-November and April in New Zealand, whereas P. suavis in Europe has been found between April and October, at least in two generations (James 2008). Adults are nocturnal and attracted to light, few short-winged females may be incapable of flying. Immature stages: Larvae of P. suavis variable in colour, but often brown-grey, or green, with whitish irregular markings, strongest dorsally (Brown & Barron 2016). Larvae of P. hollyae brown, with orange, green and yellow variegations over the dorsal and lateral surface, well camouflaged on the Hebe stems (currently classified in Veronica, Plantaginaceae) (Stephens et al. 2007). Larvae of P. suavis dropping to the ground using silken thread. Pupation in the litter layer, or under stones or twigs (Alma 1977). Remarks: Pseudocoremia has been listed under Angeronini (Hausmann et al. 2004; 2011a), but other literature sources have classified it in the Boarmiini (for instance Stephens & Gibbs 2003), and the Boarmiini position has also been supported by molecular phylogenetics (Young 2006). Among the European genera Pseudocoremia are morphologically similar to numerous Boarmiini genera, particularly Hypomecis, sharing with it numerous structures including the setose ampulla in the male valva. Systematic position: Boarmiini.

452

159. Pseudocoremia suavis Butler, 1879 Pseudocoremia suavis Butler, 1879: Cistula ent. 2 (22): 497 (New Zealand: Mid Canterbury; Christchurch). Holotype ♂ (NHMUK, examined externally). Pachycnemia usitata Butler, 1879: Cistula ent. 2 (22): 501 (New Zealand: Dunedin; Otago). Holotype ♀ (NHMUK, examined externally).

External characters and abdomen: Wingspan 28–30 mm. Forewings various shades of brown or grey, often with moss-green hues particularly in females, irrorated with blackish. Antemedial and postmedial lines blackish, latter is characteristically evenly curved inwards before costa and outwards before inner margin. Postmedial line distally often bordered with whitish, particularly at tornus. Subterminal line blackish, undulating, often bordered with whitish. Terminal line weak, blackish. Fringes concolorous with wings. Hindwings uniform pale yellow, terminal area irrorated with blackish brown. Forewing discal spots weak, hindwing spots often absent. Wings below suffused brown to pale yellow, weakly marked. Frons, collar, thorax and abdomen concolorous with wings. Male antennae bipectinate with long pectinations, female antennae filiform. Fore- and midlegs striped blackish and pale yellow. Hindtibia of both sexes with 2+2 spurs. Tympanal organs medium-sized. Male 3rd sternite with setal comb, 8th segment elongated. Other abdominal segments without modifications.

Variation: Very variable. Forewing coloration often with various shades of brown or grey, but sometimes blackish or even whitish. Forewing markings almost absent, partly visible, or blackish and distinct. Medial area concolorous with rest of wing or distinctly darker. Terminal line continuous, broken into spots or nearly absent. Hindwings often pale yellow, but can be dull brown or grey. In New Zealand P. suavis is among the most variable species, whereas the specimens in Great Britain (Cornwall) are more uniform, blackish brown being the dominant colour. This could suggest that the initial importation of P. suavis into Great Britain was based on a limited number of specimens, the genetic diversity of the population being narrow after going through a ‘genetic bottleneck’. Male genitalia: Uncus triangular, weakly setose, apex bifid. Socii absent. Gnathos arms fused ventrally, medial element upturned, weakly dentate. Subscaphium (ventral margin of tuba analis) sclerotized. Valvae upturned, narrow. Base of valva costa sclerotized, apical portion setose. Basal margin of costa sclerotized, with setose plate, its distal part elongated. Medial area of valva membranous, with distinct setose ampulla at halfway. Juxta sclerotized, base long, apex bifid, inner margin round. Posterior arms of juxta absent. Saccus elongated, apex weakly upturned. Aedeagus long, narrow, apex weakly spoon-shaped. Vesica opens at about 135 degree angle, narrow, base sparsely covered with microcornuti, small lateral diverticulum at halfway, bearing a single cornutus. Row of small cornuti at vesica base near apex of aedeagus. Female genitalia: Papillae anales weakly sclerotized, elongated, membrane between them striated. Segments 9+10 elongated. Apophyses posteriores very long. Lamella antevaginalis membranous, rectangular. Lamella postvaginalis membranous or with small sclerotization medially. Ductus bursae short, sclerotized laterally. Corpus bursae greatly elongated, narrow, posterior part grooved and weakly coiled. Signum very small, partly stellate, located at centre of corpus bursae. Distribution and abundance: Accidentally introduced to Great Britain: Cornwall, Helston (James 2008). – In Europe very local, currently well established, sometimes with numerous specimens found in a single night. – Outside Europe in New Zealand. 453

Phenology: Plurivoltine. Recorded in April, July, September and October. Numerous specimens collected at the end of September in 2016 (J. Clarke, pers. comm.). Adults starting to fly at dusk, active also at night, both sexes readily attracted to light (J. Clarke, pers. comm.). In New Zealand the larvae undergo the development (from egg hatch to adult) in around 60 days, but they can have either five or six larval instars (Bernd et al. 2004), a phenomenon the authors call developmental polymorphism. In New Zealand the adults are observed mostly between September and April (peaking in December and January), i.e. during local spring and summer, but there are no clearly synchronized generations (Berndt et al. 2004). Individual specimens may be found throughout the year (Alma 1975). Biology: In Great Britain the eggs are laid in groups, and larvae were successfully reared in captivity on Pinus sylvestris and Cupressocyparis leylandii needles (Brown & Barron 2016). In New Zealand P. suavis is polyphagous on a wide range of trees and shrubs, including Nothofagus spp., Podocarpus spp., Kunzea ericoides and it has caused serious defoliation on some exotic forest plantations, particularly with Pinus radiata and Pseudotsuga menziesii (Scott 1984; Berndt et al. 2004). It is also found commonly in New Zealand on exotic Ulex europaeus (James 2008). Habitat: Heathland, forests and gardens near coast, at sea-level, potentially more abundant amongst the trees under dense tree cover (J. Clarke, pers. comm.; Brown & Barron 2016). In New Zealand P. suavis inhabits lowlands up to 900 m. Similar species: No similar species in Europe. Mottled specimens may resemble Hydriomena Hübner, 1825 (Larentiinae), but the bipectinate male antennae and elongated abdomen are distinct in P. suavis. Genetic data: BIN: BOLD:AAZ0625 (n=7 from United Kingdom, New Zealand). Genetically homogeneous, maximum variation 1.1%, involving both European and New Zealandian populations. Nearest species: Arichanna jaguararia (Guenée, 1858) from China (7.2%), 454

Peribatodes umbraria (7.3%), probably not indicating true phylogenetic relationship but long branch attraction and indicating a long and isolated evolutionary history of P. suavis. Remarks: P. suavis may include a species complex in New Zealand, based on male response to components of the female sex pheromone (Gibb et al. 2006).

Parectropis Sato, 1980 Parectropis Sato, 1980, Tyô to Ga 31 (1/2): 41. Type species: Geometra extersaria Hübner, 1799. There are no genus synonyms.

Diversity and distribution: Twelve species in the Palaearctic and Oriental regions. External characters and abdomen: Medium-sized moths, of typical boarmiine type, wingspan up to 40 mm. Ground colour of wings grey or brownish grey. Postmedial line distinct, discal spots usually present. Male hindtibia with hair pencil, in female lacking. Venation of male and female similar. Forewing vein Sc separate, R1 and R2 free, arising from before upper angle of cell. Fovea present in males, absent in females. Male antennae ciliate-­ fasciculate. Abdomen ventrally without long hairy scales (Sato 1980). Male genitalia: Uncus short, stout, apex bluntly pointed. Gnathos arms fused. Valva apically narrower, half covered with numerous setae. Harpe long, with spines. Juxta without process. Aedeagus with many spines and single cornutus, ribbed, pointed, base corrugate (Sato 1980). Female genitalia: Papillae anales, setose, elongate. Ductus bursae sclerotized. Corpus bursae long, posteriorly ribbed with diverticulum. Anterior part membranous, partly striated, with single folded signum (Sato 1980). Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Aethalura.

160. Parectropis similaria (Hufnagel, 1767) Phalaena similaria Hufnagel, 1767: Berlin Mag. 4 (5): 512 (Germany: Berlin region). Syntype(s). Geometra corticaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 100 (Austria: Vienna district). Syntype(s). Phalaena Geometra luridata Borkhausen, 1794: Natur. eur. Schmett. 5: 235 (Europe). Syntype(s). Geometra extersaria Hübner, 1799: Samml. Eur. Schmett. 5 Geometrae (1): pl. 30, fig. 159 (Europe). Syntype(s). Boarmia extersaria var. obscurior Staudinger, 1897: Dt. ent. Z. Iris 10: 60 (Russia: Amur, ­Raddefka, Askold, Sutschan). Syntypes 12♂♀. Valid at subspecific rank (Scoble 1999). Boarmia extersaria grisescens Djakonov, 1926: Ezheg. gosud. Muz. N. M. Martyanova 4 (1): 74 (Russia: Minusinsk, Sajan Mountains). Syntypes 1♂1♀. Valid at subspecific rank (Scoble 1999). Parectropis extersaria japonica Sato, 1980: Tyô to Ga 31 (1/2): 41, figs 31, 32, 52, 64, 65 (Japan: Niigata Prefecture, Myôko, Tsubame). Holotype ♂ (NSMT). Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): cornelseni: Hoffmann (1908) (ab.); deumbrata: Lempke (1952) (ab.); heckeli: Pfau (1928) (ab.); intermediata: Rebel (1930) (ab.); obsoleta: Lempke (1952) (ab.); pallida: Lempke (1952) (ab.); strandi: Pfau (1929) (ab.).

455

External characters and abdomen: Wingspan 29–35 mm. Ground colour beige with dark grey mottling. Postmedial line less distinct or reduced near inner margin, outwardly edged whitish. Medial and antemedial lines thick, strong. Medial line on forewings running distal to discal spot or through it. Postmedial, medial and antemedial lines dilated to thicker spots at forewing costa. Terminal line broken into chain of crescent dots between veins. Wavy line distinct, on forewings with large whitish blotch in centre. Discal spot black, elongated on forewings, indistinct or absent on hindwings. On underside lines and discal spots paler, lines on forewing usually ending before inner margin. Whitish blotch on wavy line distinct. Fringes chequered beige and brownish grey, the beige segments of fringes corresponding with black dots of terminal line, inside a line, parallel to termen. Abdomen, thorax, frons and collar concolorous with ground colour. First abdominal segments usually with black rings. Antennae ciliate-fasciculate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Wing pattern darker or lighter depending on the degree of mottling. Melanic specimens are reported, ranging from intermediate forms, black with remnants of whitish lines, to nearly unicolorous black forms (Rebel 1930; Pfau 1928). The whitish blotch on wavy line usually remaining present. Melanic forms very local. Male genitalia: Uncus sclerotized, base trapezoid, apex laterally concave, bluntly pointed. Gnathos arms fused ventrally, upturned, ending in a median process. Valva apically narrower, setose, apex rounded. Costa sclerotized. Sacculus inwards with pointed harpe bearing 8–10 spines. Saccus wide, rounded. Juxta large without process. Aedeagus short, thick, vesica with a group of spines and a long cornutus, ribbed, pointed, base corrugate. Female genitalia: Papillae anales slightly sclerotized, setose, elongated. Apophyses posteriores of medium length, straight, apophyses anteriores shorter, both ending spatulate. Lamella antevaginalis desmoid, partly sclerotized. Lamella postvaginalis strongly sclerotized, with posterior projection rounded at apex. Ductus bursae wide, short, sclerotized. Posterior part of corpus bursae long and striated, with weakly sclerotized diverticulum. Anterior part membranous, partly striated. Signum elongated, folded. Distribution and abundance: Eurasiatic. In Europe from the Urals to southern England and France and from southernmost Scandinavia to northern Spain, southern France, central Italy and the northern Balkan peninsula. – Local and uncommon, but sometimes found in numbers. – Outside Europe in Georgia and Armenia (nominotypical subspecies), subsp. grisescens across southern Siberia, Altai to Dahuria, subsp. obscurior from Far East Russia to Sakhalin, subsp. japonica in Japan. Phenology: Univoltine, in the south bivoltine. In southern Scandinavia from late May to early July, in south-western Germany from mid-May to late July and with a partial second generation through August (Ebert 2003) and in the Alps from early May to mid-July and with a partial second generation from late July to late August (Kerschbaum & Pöll 2010). Larva from late June to September (Porter 2010). Overwintering as pupa. Adults active at night, attracted to light. Biology: Larva oligophagous. Found on Corylus avellana, Betula pendula, Alnus glutinosa, Quercus and Acer pseudoplatanus (Ebert 2003). Recorded on Fagus sylvatica and Tilia (Skou 1986). Pupation just below the surface of the ground (Porter 2010).

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Habitat: Silvicolous. In deciduous forests and forest edges, often on slightly moist ground. From sea-level in Scandinavia up to 900 m in south-western Germany (Ebert 2003) and in the Alps (Kerschbaum & Pöll 2010), in northern Italy up to 1400 m (Flamigni et al. 2016). Similar species: Differing from other similar Boarmiinae by the whitish blotch on the wavy line of the forewings. Sometimes, similar blotches occur in Peribatodes ilicaria but in the latter the postmedial lines of all wings are dentate, conspicuous (broken into dots as in Parectropis similaria), s-shaped on forewing and close to termen on hindwing. Genetic data: Diverging into three clearly separated BINs: BOLD:AAD5817 (n=18 from Latvia, United Kingdom, Czech Republic, Austria, France, Italy, Croatia), BOLD:AAD5816 (n=16 from United Kingdom, Germany, Czech Republic, China), BOLD:ACF5599 (n=6 from Latvia, Netherlands, Germany, Czech Republic, Austria, China). BINs diverging by minimal distances of 1.7–3.8%, requiring further taxonomic attention. Nearest congener: P. disjuncticilia (Herbulot, 1991) from Nepal (6.4%). South African “Ectropis” spoliataria (Walker, 1860) at a distance of 7.5%, several other species appearing closer due to long branch attraction phenomena.

Aethalura McDunnough, 1920 Aethalura McDunnough, 1920, Bull. Dep. Agric. Can. ent. Branch 18: 36. Type species: Boarmia intertexta Walker, 1860 [replacement name]. Genus synonyms after Scoble (1999). ‡ Aethaloptera Hulst, 1896, Trans. Am. ent. Soc. 23: 321, 358. Type species: Boarmia intertexta Walker, 1860. [Junior homonym of Aethaloptera Brauer, 1875 (Trichoptera)].

Diversity and distribution: Aethalura contains three species each in the Palaearctic and Oriental regions. Only one species in the Nearctic region.

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External characters and abdomen (Palaearctic and Nearctic species): Small or medium-sized moths, wingspan 19–31 mm. External characters as described below for Aethalura punctulata. Antennae ciliate-fasciculate in ♂, filiform in ♀. Fovea present. Forewing veins R1 and R2 fused, anastomosing with Sc. Origin of M1 separate from R3–R5. Male genitalia: Uncus rounded, short. Gnathos absent. Valva divided into dorsal and ventral arm, harpe connected with costa and sacculus, apically rounded with bunch of setae. Valva distally wider. Female genitalia: Apophyses posteriores long, straight. Apophyses anteriores shorter, spatulate at tip. Corpus bursae long, anterior part membranous, sack-shaped, with signum. Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Parectropis.

161. Aethalura punctulata (Denis & Schiffermüller, 1775) Geometra punctulata Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 106. Syntype(s) lost. Phalaena (Geometra) tigris Walchs, 1779: Naturforscher, Halle 13: 31, pl. 3 figs. 8a, 9b [8b] (Austria). Syntype(s). Geometra scopularia Thunberg, 1784: Diss. Ent. sistens Insecta Suecica 1: 7. Lectotype ♂ (UZIU). Selidosema punctularia Hübner, 1823: Verz. bekannter Schmett.: 299. Emendation of punctulata Denis & Schiffermüller (Scoble 1999). Unavailable names (infrasubspecific): albescens: Prout 1915 (ab.); anastomosaria: Lempke (1970) (f.); cinerea: Leraut (2009) (f.); costijuncta: Lempke (1970) (f.); intermedia: Lempke (1952) (f.); marginata: Lempke (1952) (f.); obscuraria: Paux (1901) (ab.); trilineata: Bru˚ čková (1945) (ab.).

External characters and abdomen: Wingspan 22–27 mm, ground colour greyish white, speckled with fine dark scales. Forewings with distinct antemedial, medial, postmedial and reduced subterminal lines, all more distinct at costa. Wavy line and discal spots indistinct. Hindwings with antemedial and postmedial line. Fringes chequered, divided by a longitudinal line. Frons, collar, thorax, abdomen and fringes concolorous with wings. Antennae ciliate-fasciculate in ♂, filiform in ♀. Hindtibia with 2+2 spurs.

Variation: Low. Ground colour lighter or darker. Line-pattern varies, particularly shape of medial and wavy line. Monochrome black (melanic) form mentioned by Bergmann (1955).

Male genitalia: Uncus rounded, short, gnathos absent. Valva deeply divided with weakly setose membrane between arms. Distal half of dorsal arm setose, in the centre with setose semi-spherical bulge. Medial part with sclerotized harpe, connected to costa and sacculus, apically rounded with bunch of setae. Saccus divided. Aedeagus thick, anteriorly thinner. Apically with slightly concave ridge. Vesica with elongated granulate plate. Female genitalia: Papillae anales elongated, soft, slightly setose. Apophyses posteriores long, straight. Apophyses anteriores about half length of apophyses posteriores, curved, ending spatulate. Ductus bursae short, sclerotized. Lamella antevaginalis hyaline, forming a ring around ostium bursae. Lamella postvaginalis small, curved, sclerotized. Corpus bursae long. Posterior part twice bent at ductus bursae. Ductus bursae bent twice anteriorly of colliculum. Posterior third of corpus bursae sclerotized, partly granular. Anterior part membranous, pear-shaped. Signum prominent, flattened, discoid. 458

Distribution and abundance: Euro-Siberian. In Europe from the Ural mountains to Ireland and northern Portugal and from central Fennoscandia to the northern Iberian peninsula, southern France, northern Italy and the northern Balkan peninsula. – Outside Europe in Caucasus, Georgia and in southern Siberia.

Phenology: Uni- or bivoltine. In Denmark from mid-April to mid-June, in south-western Germany from early April to early June and a partial second generation from late June to late July (Ebert 2003), in northern Italy from March to April with second generation in July (Flamigni et al. 2016) and in the Alps from early April to mid-June (Kerschbaum & Pöll 2010). Hibernating as pupa. Active at night, attracted to light. Biology: Larva oligophagous. Found on Betula pendula and Alnus (Ebert 2003); according to Porter (2010), Betula is the main food-plant. Recorded on Salix and Ledum palustre (Flamigni et al. 2016). Pupation in a cocoon in the surface of the ground. Habitat: Silvicolous. In mixed forests on moist ground, in moors and along rivers. From sea-level up to 800 m in south-western Germany (Ebert 2003) and up to 1300 m in northern Italy (Flamigni et al. 2016). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC5352 (n=29 from Finland, United Kingdom, Germany, Austria, France, Italy, Russia). Genetically slightly heterogeneous, maximum variation 1.1%. Nearest species: North American A. intertexta (Walker, 1860) (3.8%). Remarks: Nearctic sister species Aethalura intertexta (Walker, 1860) widespread in North America, on average smaller (wingspan 21–25 mm).

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Ascotis Hübner, 1825 Ascotis Hübner, [1825] 1816, Verz. bekannter Schmett.: 313. Type species: Geometra selenaria Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Hypopalpis Guenée, 1862, in Maillard, Notes sur l’île de la Réunion (Bourbon) (Annexe G): 29. Type species: Hypopalpis terebraria Guenée, 1862. Burichura Moore, 1888, Descr. new Indian lepid. Insects Colln late Mr W. S. Atkinson: 245. Type species: Boarmia imparata Walker, 1860. Trigonomelea Warren, 1904, Novit. zool. 11: 475. Type species: Trigonomelea semifusca W ­ arren, 1904.

Diversity and distribution: The genus Ascotis contains 11 species in the Palaearctic and Oriental regions and in Africa. Only one species in Europe. External characters and abdomen: Large-sized moths, wingspan up to 47 mm. Palaearctic species with crescent-shaped discal spots. Ground colour greyish to whitish, dark speckled. Thorax and abdomen robust. Fovea present. Forewing veins R1 and R2 arising separately from cell. R2 remains free (in ♀) or distally fused with R3 and R4 or R3–R5 (in ♂). Male and female genitalia: Palaearctic species, see under A. selenaria.

Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Cleora and Ectropis.

162. Ascotis selenaria (Denis & Schiffermüller, 1775) Geometra selenaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s). Phalaena furcaria Fabricius, 1794: Ent. Syst. 3 (2): 141 (Germany). Syntype(s). Geometra dianaria Hübner, 1817: Samml. Eur. Schmett., 5, Geometrae (1): pl. 94, fig. 483 (­Europe). Syntype(s). Herewith downgraded from subspecific rank (Scoble 1999) to synonymy, based on absence of significant and constant differential features. Boarmia selenata Herrich-Schäffer, 1863: Syst. Verz. Eur. Schmett. (Edn 3): 12. Emendation of selenaria Denis & Schiffermüller. Ophthalmodes cretacea Butler, 1879: Ann. Mag. nat. Hist. (5) 4: 373 (Japan: Yokohama). Syntype(s) (NHMUK). Valid at subspecific rank (Scoble 1999). Boarmia selenaria var. artemis Staudinger, 1897: Dt. ent. Z. Iris 10: 58 (Russia: Amur). Syntype(s). Valid at subspecific rank (Scoble 1999). Boarmia selenaria var. lutescens Wagner, 1923: Z. öst. ent. Ver. 8 (5/6): 43 (Romania: Prahora district, Hatcarau, Cornul). Syntype(s). Boarmia (Ascotis) selenaria kaolina Bryk, 1949: Ark. Zool. 41A (1): 209 (Korea: Shuotsu). Holotype ♂ (NHRS). Valid at subspecific rank (Scoble 1999). Ascotis selenaria ijimai Inoue, 1955: Tinea 2 (1/2): 85 (Japan: South Hokkaido, Shiraoi). Holotype ♂ (NHMUK). Valid at subspecific rank (Scoble 1999).

External characters and abdomen: Wingspan 29–47 mm. Ground colour whitish to grey, dark speckled. Postmedial line and antemedial line serrate. Medial line blurred, running through the discal spot or next to it, on forewings sometimes missing. Postmedial and medial line approaching each other towards inner margin. Basal line often blurred and unclear. Postmedial line sub-apically with shadow, sometimes missing. Terminal line reduced to black dots on each vein. All discal spots crescent-shaped, conspicuous, pale centered. Fringes concolorous with wings, hardly chequered. Underside much brighter, discal spots strikingly black, often also pale centered. Abdomen, thorax, frons and collar concolorous with ground 460

colour. Thorax and abdomen robust. Abdomen with two rows of black dots. Antennae ciliate-fasciculate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variable species. Ground colour varies from darker grey to whitish with touch of beige depending on local conditions. Density of speckling is also variable. Extreme form with medial line on both wings black. Some of the forms occur in certain areas more frequently or exclusively, e.g. dark (not melanic!) forms in sandy areas of eastern Germany or pale yellowish forms in western Hungary. However, all these forms are also more or less frequently observed elsewhere in Europe. Therefore we retain the validation of subspecies in Europe to be unnecessary. Male genitalia: Uncus hooked, beak-shaped. Gnathos arms fused, wide and flat ending, both sclerotized. Valva long and narrow over whole length with sclerotized ridge near costa, and second narrow medial ridge, irregularly shaped, with spinose apex. Saccus rounded, shortly projecting. Aedeagus thick, as long as valva length, apically ending in a finger-shaped process, tip rounded. Vesica with elongated, curved serrate plate and curved sclerite. Female genitalia: Papillae anales elongated, membranous, soft, slightly setose. ­Apophyses posteriores very long, straight. Apophyses anteriores shorter, ending spatulate. Ductus bursae short, sclerotized. Corpus bursae about nine times as long as anteriorly wide, sacshaped, membranous, slightly striated. Anterior part with striated diverticulum. Signum prominent, stellate. Distribution and abundance: Eurasiatic. In Europe from the Urals through central Europe to France and the Iberian peninsula and from central Germany to northern Spain, southern France including Corsica, Italy including Sardinia and Sicily, Malta and the Balkan pen­ insula including Greece and Crete. – Rather common, usually occurring in limited numbers. – Outside Europe in Israel, Cyprus, Turkey, Georgia, Armenia, Azerbaijan and through

461

southern Siberia to Altai. In the east to Far East Russia (subsp. artemis), Korea (subsp. kaolina), China and Japan (subsp. cretacea and ijimai). Phenology: Bi- or trivoltine. Generations apparently overlapping, from late March in southern­ most Europe to mid-September. Larva from June to May, hibernating. Adults active at night, readily attracted to light. Biology: Larva polyphagous. Recorded on Populus, Betula, Castanea, Quercus, Ulmus, Ficus, Clematis, Laurus, Persea, Rubus, Rosa, Pyrus, Malus, Amelanchier, Mespilus, Crataegus, Cytisus, Genista, Robinia, Acacia, Mimosa, Citrus, Pistacia, Schinus, Rhamnus frangula, Hippophae, Eucalyptus, Arbutus, Ligustrum, Plumeria, Sambucus, Viburnum, Cannabis, Rumex, Melilotus, Medicago, Trifolium, Glycine, Euphorbia, Foeniculum, Daucus, Pimpinella, Salvia, Solanum, Arctium, Aster, Artemisia, Taraxacum, Dahlia and Aspa­ ragus, reared on Convolvulus (Flamigni et al. 2016). Pupation on or in the surface of the ground (Flamigni et al. 2016). Habitat: In southern Europe occurring in nearly any kind of habitat while, in central Europe, preferring deciduous forests on relatively dry soil, in eastern Germany also in Pinus forests (J. Gelbrecht pers. comm.). From sea-level (Greece) up to 1600 m in Spain. Similar species: Hypomecis punctinalis (Scopoli, 1763) with male antennae partially bipectinate, smaller thorax and smaller discal spots. Genetic data: All European populations belonging to one single (but very heterogeneous) BIN: BOLD:AAC0403 (n=61 from 11 countries from Germany to Sicily, Malta and Crete in the south and from France (Corsica) to Far East Russia and China); maximum variation in Europe 3.0%, requiring further investigation. Populations from Cyprus and southern Turkey to Israel and Jordan with separate BIN AAC0406 (n=9) at a distance of 5.0%, possibly requiring to be treated as a separate taxon. In central and southern Asia many populations, often identified as A. selenaria or A. selenaria dianaria belonging to other BINs and probably not conspecific with European A. selenaria. Nearest species: Congeners from China (Yunnan) currently (mis)identified as A. selenaria dianaria (5.3%).

Cleora Curtis, 1825 Cleora Curtis, 1825, Br. Ent. 2: 88. Type species: Geometra cinctaria Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Cerotricha Guenée, 1858, in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 284. Type species: Cerotrichia licornaria Guenée, 1858. ‡ Barsine Meyrick, 1883, N. F. Jl Sci. Dunedin 1: 530. Type species: Scotosia panagrata Walker, 1862. [Junior homonym of Barsine Walker, 1854 (Lepidoptera, Arctiidae)]. Meyrickia Butler, 1884, Entomologist’s mon. Mag. 21: 133. Type species: Scotosia panagrata Walker, 1862. [Unnecessary replacement name for Barsine Meyrick, 1883.] Aegitrichus Butler, 1886, Trans. ent. Soc. Lond. 1886: 434. Type species: Aegitrichus lanaris Butler, 1886. Chogada Moore, [1887] 1884–7, Lepid. Ceylon 3: 415. Type species: Boarmia alienaria Walker, 1860. Carecomotis Warren, 1896, Novit. zool. 3: 402. Type species: Carecomotis perfumosa Warren, 1896. Neocleora Janse, 1932, The Moths of South Africa 1: 119 (key), 266. Type species: Boarmia tulbaghata Felder & Rogenhofer, 1875.

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Diversity and distribution: Approximately 185 species in the Palaearctic, Nearctic and Oriental regions, in Africa and Madagascar, and in Australia (Scoble 1999). Mainly in the Old World, Hodges (1983) including only two Nearctic species, one species only in Europe. Further research is likely to lead to changes in the species list. External characters and abdomen: Medium-sized moths. Ground colour grey to white marked with black and dark brown spots. Male forewing with prominent fovea. Discal spots conspicuous, irregularly formed, with white centre. Male antennae partially bipectinate over basal half to two thirds or (in some species) quadripectinate (Holloway 1994). Female antennae filiform. Hindtibia of both sexes with 2+2 spurs. Male genitalia: Sacculus bearing stout ornamentation, i.e. robust spine(s) or other sclerotized processes. Vesica of aedeagus bifurcate, cornuti often very large (Holloway 1994). Female genitalia: Papillae anales elongate, slender as in Ascotis. Corpus bursae long, usually extensively sclerotized, fluted over a long basal section. Typical ennomine signum in the bulb (Holloway 1994). Remarks: The Afrotropical and Asian species of the genus Cleora have been in the focus of several studies (Fletcher 1953; 1967; Sato 1989; see also Holloway 1994). The above given genus description is mainly based on Holloway (1994) covering a number of mainly Asian species.

163. Cleora cinctaria (Denis & Schiffermüller, 1775) Geometra cinctaria Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wienergegend: 101 (Austria: Vienna district). Syntype(s) lost. Phal[aena] Geometra pascuaria Brahm, 1791: Insektenkalender für Sammler und Oekonomen 2: 120 (Germany: near Mainz). Syntype(s). Boarmia cinctaria f. duponcheli Wagner, 1923: Z. öst. EntVer 8 (5/6): 43 (locality not stated). Syntype(s). Valid at subspecific rank (Scoble 1999). Cleora cinctaria bowesi Richardson, 1952: Entomologist’s Rec. J. Var. 64: 269, pl. 11, figs 2, 7 (Scotland: Perthshire, Struan). Holotype ♂. Valid at subspecific rank (Scoble 1999). ‡ Cleora cinctaria fumata Inoue, 1968: Tinea 7 (2): 158, pl. 30, fig. 3, text-fig. 1c (Japan: Nagano Prefecture, Sugadaira, 1300 m). Holotype ♂. Junior secondary homonym of Carecomotis tenebrata fumata Fletcher, 1953. Cleora cinctaria superfumata Inoue, 1972: Tinea 9 (1): 235. Replacement name for fumata ­Inoue, 1968. Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): caminariata: Fuchs (1882) (ab.); consimilaria: Duponchel (1829) (ab.); fuscaria: Cockayne (1949) (ab.).

External characters and abdomen: Wingspan ♂ 25–36 mm, ♀ 26–33 mm. All medial lines usually conspicuous. Basal and antemedial lines curved, sometimes broken, basal line often dilated to small band. Postmedial line associated distally with pale brown band. Medial band either running through discal spot or proximal to it. Wavy line quite straight. Terminal line dotted. Fringes with indistinct longitudinal line, chequered dark and pale grey. Paler segments of fringes corresponding with black dots of terminal line. Hindwings slightly paler. Frons and collar concolorous with ground colour. Antennae partially bipectinate to 3/4 of length (about 18 flagellomeres free of branches) in male, filiform in female. Abdomen with double row of black dots, first segment anteriorly silver-grey.

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Variation: Forewing medial area and hindwing costal area vary in ground colour from whitish to grey and brownish grey. Specimens with more white are significantly more contrasted. Subsp. bowesi from Scotland with silvery white ground colour and blackish grey markings (Skinner 2009). Male genitalia: Uncus long, narrow, tapered, beak-shaped. Gnathos broad, rounded, arms fused. Valva broad, apically becoming narrower. Costa sclerotized, setose subapically. Sacculus with thorn. Harpe sclerotized, nearly square plate with rounded corners and narrow pointed process. Juxta narrow, centrally forming sclerotized line. Saccus rounded with upward tip medially. Aedeagus straight, rounded, 5–6 times as long as wide. Vesica with one or two teeth and a group of smaller cornuti, which are positioned mostly outside the ­uneverted vesica. Number of cornuti varies. Female genitalia: Papillae anales membranous, striated, slightly setose. Apophyses posteriores long, straight. Apophyses anteriores shorter, ending spatulate. Lamella antevaginalis wide, sclerotized, double lateral arms, one wide, rounded, the other short, tapered. Corpus bursae membranous, slightly striated. Signum discoid, irregularly shaped. Distribution and abundance: Eurasiatic. In Europe widespread from the Ural mountains to Ireland and Portugal and from central or southern Fennoscandia to Spain, France including Corsica, Italy including Sardinia and the Balkan peninsula including northern Greece. In Scotland replaced by subsp. bowesi. The distribution map in Leraut (2009) shows a much too large distribution in northern Europe and in the Iberian peninsula while the occurrence in Greece is missing. – In Europe relatively common though local. Usually found in limited numbers. – Outside Europe widely distributed from Turkey, Armenia and central Asia through southern Siberia to Far East Russia, northern China and Japan (subsp. super­ fumata).

464

Phenology: Univoltine or bivoltine. In southern Scandinavia from late April to late May, in south-western Germany from early April to late May (Ebert 2003), in northern Spain from mid-April to early June (Redondo et al. 2009). By Lake Garda, Lake Iseo and in Sardinia with a second generation from late August to October. Caterpillars from May to July or ­August (Seppänen 1970). Hibernation as pupa. Adults active at night, attracted to light. Biology: Larva polyphagous. Found on Genista anglica, Vaccinium myrtillus, Galium verum, Dianthus carthusianorum, Cytisus scoparius, Ledum palustre (Wegner 2013), Pinus silvestris, Betula pendula, Prunus spinosa, Genista sagittalis, Coronilla varia and Thymus pulegioides (Ebert 2003). Recorded on Frangula alnus, Quercus, Myrica gale (Skou 1986), Salix, Populus, Alnus, Rubus, Sorbus, Cytisus, Spartium, Cistus, Erica, Calluna, ­Vaccinium,Carex, Rumex, Biscutella, Lotus, Vicia, Psoralea, Malva malacoides, Hypericum, Peucedanum, Lysi­ machia, Galium, Scabiosa, Anthemis, Achillea, Artemisia and Tanacetum (Flamigni et al. 2016). Pupation in the ground. Habitat: Silvicolous. In a very wide variety of habitats such as deciduous and mixed forests on both sandy and moist soil, in wet and dry meadows, in hedges in open cultivated land, in scrubland, gardens and parks. From sea-level in northern Europe and up to 1000 m in south-western Germany (Ebert 2003), up to 1500 m in Italy (Flamigni et al. 2016) and in the Pyrenees (Redondo et al. 2009). Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC4786 (n=28 from Finland, United Kingdom, Germany, Austria, France, Italy, Russia (Urals), Kyrgyzstan). Genetically slightly heterogeneous, maximum variation 1.5%. Nearest species: South Asian Cleora injectaria (Walker, 1860) (3.9%). Remarks: Wegner (2013) deals in details with the biology and habitat in north-western Germany.

Ectropis Hübner, 1825 Ectropis Hübner, [1825] 1816, Verz. bekannter Schmett.: 316. Type species: Geometra crepuscularia Denis & Schiffermüller, 1775. Genus synonyms after Scoble (1999). Boarmia Stephens, 1829, Nom. Br. Insects: 43. Type species: Geometra crepuscularia Denis & Schiffermüller, 1775. [Junior homonym of Boarmia Treitschke, 1825 (Geometridae, Enno­ minae), and junior objective synonym of Ectropis Hübner, [1825] 1816]. Tephrosia Boisduval, 1840, Genera index meth. eur. Lepid.: 198. Type species: Geometra crepuscularia Denis & Schiffermüller, 1775. ‡ Coenobita Gistl, 1848, Naturg. Thierreichs: xi. [Unnecessary replacement name for Tephrosia Boisduval, and junior homonym of Coenobita Latreille, 1829 (Crustacea)].

Diversity and distribution: Scoble (1999) lists 100 species in Ectropis and four other species tentatively attributed to this genus. External characters and abdomen: Medium to larger-sized moths with ground colour fawn, pale yellow or white. Wings with multiple, fine, dark fasciae. Fasciae mostly crenulate, often interrupted at veins near inner margin. The inner component of the double postmedial lines is darker, more dentate where it interacts with veins M3 and CuA1 (Holloway 1994). Fovea present but small. Antennae ciliate-fasciculate in male, filiform in female. Veins Sc and R1 separate, R1 and R2 stalked in male, entirely coincident in female (Sato 1986).

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Male genitalia: Uncus developed, triangular, tip long, tapered. Gnathos absent or much reduced. Valva narrow, simple, tapering. Juxta constricted in central part, distally extending into sclerotized scobinate ring. Aedeagus with tubular vesica, in some species with sclerotized spine or lobe developed from near base (Holloway 1994). Female genitalia: Papillae anales distinctly elongate. Apophyses anteriores often very long (see also Holloway 1994). Lamella antevaginalis with a pair of triangular (alate) extensions. Ductus bursae short. Corpus bursae ovate or pear-shaped, with large, stellate signum (Sato 1986). Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Cleora and Ectropis.

164. Ectropis crepuscularia (Denis & Schiffermüller, 1775) Geometra crepuscularia Denis & Schiffermüller, 1775: Ank. syst. Werkes Schmett. Wiener­ gegend: 101 (Austria: Vienna district). Neotype (NHMW). Phalaena Geometra virginiaria Cramer, 1780: Uitlandsche Kapellen (Papillons exot.) 3: 149, 176 (index), pl. 275, fig.G (U.S.A.: Virginia). Syntypes ♂♀. Phalaena Geometra bistortata Goeze, 1781: Ent. Beytr. 3 (3): 438. Syntype(s). Phalaena Geometra biundulata Villers, 1789: Linn. ent. 2: 337 (Italy: Brescia). Syntype(s). Phalaena Geometra biundularia Borkhausen, 1794: Natur. eur. Schmett. 5: 162 (Europe). Syntype(s). Phalaena Geometra baeticaria Scharfenberg, 1805: Vollständige Naturgeschichte der schädlichen Forstinsekten 3: 638 (Europe). Syntype(s). Boarmia strigularia Stephens, 1831: Illust. Br. Ent. (Haustellata) 3: 192 (England: Kent, Darenth Wood). Syntype(s). Tephrosia laricaria Doubleday, 1847: A synonymic list of British Lepidoptera: Sheet B, 16 (­United Kingdom). Syntype(s). Validated at subspecific rank by Scoble (1999), not mentioned in ­Leraut (2009), herewith downgraded to synonymy based on absence of constant and significant differential features. Putative differences in phenology not justifying separate taxonomic status unless correlated with differential features in morphology or DNA. Boarmia defessaria Freyer, 1847: Neuere Beitr. Schmettkde 6 (85): 46, pl. 510, fig. 1 (Europe). Syntypes. Tephrosia occiduaria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. ­Lepid.) 9: 266 (North America). Syntypes 2♂. Tephrosia abraxaria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 21: 403 (­Canada: Nova Scotia). Syntypes 4♂ (NHMUK). Boarmia cineraria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 21: 488 (Canada: Nova Scotia). Holotype ♂ (CNCI). Boarmia divisaria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 21: 489 (Canada). Syntype(s) ♀. Tephrosia intrataria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 21: 403 (North America). Holotype ♂. Boarmia signaria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 20: 346 (Canada: Nova Scotia). Holotype ♀ (NHMUK). Tephrosia spatiosaria Walker, 1860: List Specimens lepid. Insects Colln Br. Mus. 21: 403 (U.S.A.: New York). Syntypes 4♂♀ (NHMUK). Boarmia crepusculata Herrich-Schäffer, 1863: Syst. Verz. Eur. Schmett. (Edn 3): 12. Emendation of crepuscularia Denis & Schiffermüller. Tephrosia fumataria Minot, 1869: Proc. Boston Soc. nat. Hist. 13: 84 (U.S.A.: Massachusetts, West Roxbury). Syntype(s).

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Boarmia fernaldaria Hulst, 1888: Entomologica am. 3 (11): 215 (U.S.A.: Maine). Syntypes 2♂1♀. Boarmia lutamentaria Graeser, 1889: Berl. ent. Z. 32 (2): 401 (Russia: Amurland, Chabarofka). Syntypes 4♂7♀. Valid at subspecific rank (Scoble 1999). Boarmia crepuscularia var. tristis Riesen, 1897: Stettin. ent. Ztg. 58: 322 (Courbière). Syntypes 2, including at least 1♂. Boarmia crepuscularia pallidaria Krulikovski, 1910: Ent. Obozr. 9 (3): 313 (eastern Russia: Viatka). Syntype(s). Valid at subspecific rank (Scoble 1999). Boarmia (Ectropis) crepuscularia bergmanaria Bryk, 1949: Ark. Zool. 41A (1): 210 (Korea: Shuotsu). Holotype ♂ (NHRS). Valid at subspecific rank (Scoble 1999). Unavailable names (infrasubspecific): albolimbata: Alberti (1954) (ab.); cinerea: Lempke (1952) (f.); defessaria: Freyer (1848) (ab.); extrema: Rebel (1930) (ab.); intermedia: Lempke (1952) (f.); marginata: Lempke (1970) (f.); obscurata: Heinrich (1917) (ab.); obsoleta: Lempke (1952) (f.); passetii: Thierry-Mieg (1886) (ab.); unicolor: Lempke (1970) (f.).

External characters and abdomen: Wingspan 30–38 mm, second (or third) generation smaller, 25–30 mm. Ground colour of forewings whitish grey to brownish grey with dark mottling. Postmedial line double, dentate, both with two distinct teeth on veins M3 and CuA1. Wavy line whitish. Terminal line black, dotted. Medial line weak, not sharply defined. Antemedial line distinct, dark, bent. Postbasal line brown, blurred, straight. Discal spots indistinct or absent. Fringes, abdomen, thorax, frons and collar concolorous with ground colour, fringes often with weak longitudinal line. Lighter specimens with double dots on the first abdominal segments. Antennae ciliate-fasciculate in male, filiform in female. Hindtibia of both sexes with 2+2 spurs. Variation: Variable species. Ground colour and degree of irroration on all wings variable in a wide range from greyish to brownish, darker or lighter. Transverse lines more or less distinct. Blackish and intermediate specimens with a conspicuous whitish wavy line on fore- and hindwings are common mainly in Central and Western Europe, there often dominating. Totally black (melanic) specimens are very rarely observed (Bergmann 1955). Male genitalia: Simple structure. Uncus sclerotized, tip narrow, long, tapered. Gnathos absent. Valva narrow, membranous, at base wider. Costa weakly sclerotized. Apical area ­setose. Base of valva with long coremata. Juxta developed as flat sclerite, constricted at centre. Saccus rounded. Aedeagus long, cylindrical. Vesica with spinose area and thumbshaped cornutus, apex of vesica very long, narrow, membranous. Female genitalia: Papillae anales long tube, distally membranous, setose. Apophyses posteriores very long (on average 8.2 mm including ovipositor), thin, straight, at tip not dilated. Apophyses anteriores shorter, spatulate at tip. Lamella antevaginalis alate, weakly sclerotized, lamella postvaginalis round sclerite. Antrum funnel-shaped, membranous, colliculum and anterior part of ductus bursae narrow, sclerotized. Corpus bursae pear-shaped, membranous, concentrically striated with large, stellate signum at centre. Distribution and abundance: Holarctic. In Europe from the Ural mountains to the British Isles and from northern Fennoscandia to northern Iberian peninsula, southern France including Corsica, Italy and the northern Balkan peninsula. – Common, sometimes found in numbers. – Outside Europe widely distributed from Turkey and Transcaucasus through northern Kazakhstan and southern Siberia to Yakutia, Mongolia, China, Korea, Far East Russia, Sakhalin, southern Kurile Islands and Japan (various subspecies). Also in North America.

467

Phenology: Uni- or bivoltine. In the north of the European distribution area in only one generation from mid-April to early or mid-June. In Denmark from early April to early June and from late June to early August. In south-western Germany from early March to late May and from mid-June to early August (Ebert 2003). In northern Spain from mid-March to early June and from mid-July to early September (Redondo et al. 2009). Occasional specimens of an incomplete third generation from late August to mid-October recorded in England, Denmark and Germany (M. Corley pers. comm.; P. Skou, A. Hausmann own observation). Caterpillars generally from mid- or late May to mid-July and from early August to early September. Hibernating as pupa. Adults active at night, males readily attracted to light, females only rarely. Biology: Larva polyphagous. Found on Picea excelsa, Salix, Betula pendula, Alnus ­glutinosa, Fagus sylvatica, Quercus, Rumex, Berberis vulgaris, Malus domestica, Sorbus aucuparia, ­Rubus, Rosa canina, Prunus cerasifera, P. padus, P. spinosa, Cytisus scoparius, Robinia pseudo­acacia, Acer pseudoplatanus, Frangula alnus, Pimpinella saxifraga, Daucus carota, Vaccinium myrtillus, Fraxinus excelsior, Ligustrum vulgare, Galeopsis tetrahit, Stachys sylvatica, Atropa belladonna, Scrophularia nodosa, Sambucus nigra, Viburnum opulus, Achillea millefolium, Artemisia, Cirsium and Prenanthes purpurea (Ebert 2003). Pupation in the soil. Habitat: Occurring in a wide variety of habitat types, due the the large number of foodplants. Apparently most common in open forests. From sea-level up to 1500 m in the Alps (Kerschbaum & Pöll 2010) and to 1700 m in the French Pyrenees. Similar species: Paradarisa consonaria (see no. 111). Genetic data: Species showing, in Europe, a deep genetic split (6.4%) into two often sympatric, and very homogeneous BINs: BOLD:AAA2076 (n=117 from Finland, Netherlands, ­Germany, Austria, Czech Republic, France, Italy, Macedonia, Ukraine, western to eastern Russia, Kazakhstan, Iran) and BOLD:ACE6053 (n=44 from Norway, Finland, United 468

­ ingdom, Nether­lands, Germany, westernmost Austria, France, north-eastern and southern K Italy, Croatia, China, Japan). In central and eastern Austria (including type locality), Czech Republic, Russia, Ukraine, Kazakhstan and Iran, so far, only BIN BOLD:AAA2076 was recorded, whilst being almost absent in most parts of northern Italy (Alto Adige, Friuli). Sequencing of nuclear genes did not reveal indications of a beginning speciation event (Mutanen 2015). A possible explanation may be a postglacial merging of two species with different mitochondrial evolution during certain glacial periods. North American populations of E. crepuscularia with a similar split, but separate BINs BOLD:AAA2073 (n=87) at a distance of 2.3% from BOLD:ACE6053 and BOLD:AAA2074/2075 (n=35) at a distance of 5.3–5.6% from BOLD:ACE6053. Nearest species: several Asian congeners, e.g. E. bhurmitra (­Walker, 1860), E. dentilineata (Moore, 1868), E. obliqua (Prout, 1915) (6.8–8.5%).

Arichanna Moore, 1868 Arichanna Moore, 1868, Proc. zool. Soc. Lond. 1867: 658. Type species: Scotosia plagifera Walker, 1866. Genus synonyms after Scoble (1999). ‡ Rhyparia Hübner, [1825] 1816, Verz. bekannter Schmett.: 305. Type species: Phalaena melanaria Linnaeus, 1758. [Junior homonym of Rhyparia Hübner, [1825] 1816, (Lepidoptera: Arctiidae)]. Icterodes Butler, 1878, Illustr. typical Specimens Lepid. Heterocera Colln Br. Mus. 2: ix. Type species: Rhyparia fraterna Butler, 1878. Paricterodes Warren, 1893, Proc. ent. Soc. Lond. 1893: 389. Type species: Abraxas tenebraria Moore, 1868. Hemipyrrha Grote, 1896, Proc. ent. Soc. Lond. 1896: xv. Type species: Phalaena melanaria Linnaeus, 1758. [Unnecessary replacement name for Rhyparia Hübner, [1825] 1816]. Phyllabraxas Leech, 1897, Ann. Mag. nat. Hist. (6) 19: 441. Type species: Phyllabraxas curvaria Leech, 1897. Epicterodes Wehrli, 1933, Ent. Z., Frankf. a. M. 47: 29, 41, 47, 51. Type species: Arichanna flavomacularia Leech, 1897.

Diversity and distribution: The genus Arichanna currently contains 74 species. Species are found in the Palaearctic (mostly East Palaearctic) and Oriental regions (most species between India and China). Only one species in Europe. External characters and abdomen: Medium-sized to large moths, wingspan 32–55 mm. Most species with conspicuous colours, forewings often brown, mixed with grey, orange or/ and white, and with dark lines and blotches. Hindwing grey/orange/white with dark blotches, discal spot usually distinct, in some species absent. Wing margins smooth. Forewing veins R1 and R2 separately arising from cell. Male genitalia: Valva of the sub-group named Icterodes Butler, 1878 (see remarks) with sclerotized costa and subcostal setose ridge, at base with setose projection. Aedeagus apically tapered, vesica with one or two cornuti. Female genitalia: Papillae anales soft, setose. Apophyses posteriores very long, apophyses anteriores shorter. Anterior end of corpus bursae membranous, striated. Signum stellate. Ductus bursae wide, sclerotized. Lamella postvaginalis a sclerotized plate. Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Bupalus. Wehrli (1939–1954) splits the genus into six ‘groups’ (sic!). A. melanaria belongs to his second ‘group’ (Icterodes Butler, 1878).

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165. Arichanna melanaria (Linnaeus, 1758) Phalaena (Geometra) melanaria Linnaeus, 1758: Syst. Nat. (Ed. 10): 521 (Europe). Syntype(s) (LSL). Icterodes fraterna Butler, 1878: Illustr. typical Specimens Lepid. Heterocera Colln Br. Mus. 2: ix, 53, pl. 37, fig. 9 (Japan: Yokohama). Syntype(s) (NHMUK). Originally as species. Valid at subspecific rank (Scoble 1999). Rhyparia melanaria var. askoldinaria Oberthür, 1880: Études ent. 5: 52, pl. 9, fig. 11 (Russia: Askold Island). Syntypes 2. Valid at subspecific rank (Scoble 1999). Icterodes sordida Butler, 1883: Ann. Mag. nat. Hist. (5) 11: 116 (W Korea: Jinchuen). Syntype(s) ♀ (NHMUK). Rhyparia melanaria var. decolorata Staudinger, 1892): Dt. ent. Z. Iris 5 (2): 374 (Mongolia: Kentei region). Syntypes including ♀. Valid at subspecific rank (Scoble 1999). Icterodes melanaria praeolivina Wehrli, 1933: Ent. Z., Frankf. a. M. 47 (4): 31, fig. 6 (Korea). Syntypes 3♂1♀. Unavailable names (infrasubspecific): hanseni: Hedemann (1881) (ab.).

External characters and abdomen: Wingspan ♂ 32–36 mm, ♀ 32–40 mm. Forewings: Ground colour whitish grey, towards base becoming more or less darker. Costa orange-grey with scattered black scales. All transverse lines developed as rows of blackish spots of different size. Hindwings: Ground colour yellow-orange, medial lines present as larger or smaller black spots. All discal spots large, black. Fringes of ground colour, chequered with dark grey, in hindwings conspicuous only near tornus. Frons, collar and thorax brownish grey to yellow-­grey, thorax sometimes with indistinct orange and black pattern. Abdomen yellow-­ orange, dorsally with black dots. Antennae partially bipectinate to 9/10 of length (about 7–8 flagellomeres free of branches) in male, filiform in female. Hindtibia of both sexes with 2+2 spurs, in male with hair pencil. Variation: Size of black spots variable. Forewings can be darker or lighter. Male genitalia: Uncus triangular, apex sclerotized, concave. Gnathos arms narrow, fused ventrally, forming a rounded structure with small process in the centre. Valvae wide, costa sclerotized, setose, subcostal setose ridge, at base with setose, rounded projection. Saccus rounded, laterally concave. Aedeagus thick, cylindrical, apex elongate, sclerotized, projecting. Vesica with two cornuti, one tapered, the other shorter, granular. Female genitalia: Papillae anales wide, soft, setose. Apophyses posteriores long, weakly spatulate at tip. Apophyses anteriores short, 3/4 of the length of tergum A8, weakly spatulate at tip. Ductus bursae wide, short. Colliculum wide, sclerotized. Corpus bursae large, posterior part striated, sclerotized, granular. Anterior end very large, oval, membranous. Signum stellate, two distinct teeth on one side, positioned in a concentrically striated area. Lamella postvaginalis a sclerotized triangular plate with rounded corners. Distribution and abundance: Eurasiatic. In Europe from the Urals to Norway, Denmark, Germany and the Alps. The distribution map in Leraut (2009) shows no distribution in Ukraine. – A local species depending on habitat and food-plant, but sometimes found in numbers. In Denmark observed as migrant. – Outside Europe widely distributed in Siberia, Transbaikalia, Mongolia, northern China, Korea, Far East Russia, Sakhalin, the Kuriles and Japan (various subspecies).

470

Phenology: Univoltine. In Denmark from mid-June to mid-August and in south-western Germany from late June to mid-August (Ebert 2003). Caterpillars from July-August to early June, hibernating. In southern Bavaria apparently overwintering as egg (W. Hundhammer pers. comm.; repeated rearing observations). Adults active at night, attracted to light. Biology: Larva oligophagous. Found on Vaccinium uliginosum (Ebert 2003), recorded on Ledum palustre (Skou 1984). Reared on Rhododendron (M. Leipnitz pers. comm.). Pupation on the ground. Habitat: Hygrophilous. In moors and in open forests on moor-like soils. In northern Europe from sea-level, in southern Germany from 400 up to 1100 m. Similar species: No similar species in Europe. Genetic data: BIN: BOLD:AAC8717 (n=17 from Finland, Germany, Austria, Russia, J­apan), one exactly sequence-sharing specimen from eastern Russia/Vladivostok identified as A. askoldinaria. The specimen from Japan diverging by 0.9% (subsp. fraterna). North-easternmost Chinese populations, currently identified as A. melanaria with another BIN BOLD:AAC8718 (n=8), diverging by 3.3% requiring further taxonomical study. Nearest species: A. sinica (Wehrli, 1933) from China and A. sparsa (Butler, 1890) from Nepal and India (both 3.4%), on BOLD eight further unidentified Arichanna species from China at similar distances.

Bupalus Leach, 1815 Bupalus Leach, [1815] 1830, Brewster’s Edinburgh Encyclopaedia 9: 134. Type species: Phalaena piniaria Linnaeus, 1758. Phaophyga Billberg, 1820, Enumeratio Insect. Mus. G. J. Billberg: 89. Type species: Phalaena piniaria Linnaeus, 1758.

471

Chleuastes Hübner, 1822, Syst.-alphab. Verz.: 38–52. Type species: Phalaena piniaria Linnaeus, 1758. Catograpta Hübner, [1823] 1816, Verz. bekannter Schmett.: 297. Type species: Phalaena piniaria Linnaeus, 1758. ‡ Bupala Snellen, 1867, Vlinders Nederland: 514 (key), 602. Type species: Phalaena piniaria Linnaeus, 1758. [Junior homonym of Bupala Pascoe, 1863 (Coleoptera)].

Diversity and distribution: According to Scoble (1999) Bupalus contains four species in the Palaearctic region. Leraut (2009) transferred Bupalus michaellarius Rungs, 1941 to a monotypic genus Ifrania Leraut, 2009. This is acceptable because of the completely different genitalia structure (see Leraut 2009). After Beljaev (2016) the validity of the taxon Bupalus cembraria (Motschulsky, 1866) remains doubtful, type specimens are unknown, the original diagnosis is reminiscent of dark coloured Siberian forms of Angerona prunaria. With that the number of Bupalus species decreases to three or two. External characters and abdomen: Medium-sized moths with ground colour whitish or yellow-­brown in male, whitish or reddish brown in female. Apical and costal areas brownish or grey darkened. Detailed description of the European species see below. Genitalia of European species: see species description below. Remarks: In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Arichanna. According to Pierce (1914) and Wehrli (1939–1954) the male genitalia show no similarity to any other genus of the family Geometridae, whilst being somewhat reminiscent of Ochropacha duplaris (Linnaeus, 1761) (Drepanidae).

166. Bupalus piniaria (Linnaeus, 1758) Phalaena (Geometra) piniaria Linnaeus, 1758: Syst. Nat. (Ed. 10) 1: 520. Syntype(s) (LSL). Phalaena (Geometra) tiliaria Linnaeus, 1758: Syst. Nat. (Ed. 10) 1: 520. Fidonia iberaria Kolenati, 1846: Meletemata Entomologica 5: 106 (Iberia). Syntype(s). Bupalus piniaria bernieri de Lajonquiére, 1958: Bull. mens. Soc. linn. Lyon 27 (10): 304 (France: Gironde, Villenave d’Ornon. Holotype ♂. Validated at subspecific rank (Scoble 1999). Downgraded to synonymy by Leraut (2009). Bupalus piniarius espagnolus Eitschberger & Steiniger, 1975: Ent. Z., Frankfurt a. M. 85 (21): 241, figs 1–5 (Spain: Andalusia, Lapeza, Cortijo de Carbonales (Granada)). Holotype ♂. Validated at subspecific rank (Scoble 1999). Downgraded to synonymy by Leraut (2009). Unavailable names (infrasubspecific): albida: Lempke (1970) (f.); costimaculata: Lempke (1952) (f.); dziurzynskii: Koller (1912) (ab.); flava: Lempke (1970) (f.); flavescens: White (1876) (ab.); ­ rulikovski fulvaria: Dziurzynski (1912) (ab.); funebris: Cockayne (1939) (ab.); fuscantaria: K (1908) (ab.); hirschkei: Dziurzynski (1912) (ab.); mughosaria: Gumppenberg (1887) (var.); nigricans: Dziurzynski (1912) (ab.); nigricarius: Backhaus (1889) (ab.); strigata: Dziurzynski (1912) (ab.); unicolora: Strand (1901) (ab.).

External characters and abdomen: Wingspan 30–35 (38) mm. Distinct sexual dimorphism. Male: Ground colour dark brown with whitish or yellow-brown medial area on the upper side. Veins, medial and antemedial lines more or less dark brown, diffuse, dividing the pale area into blotches. Hindwing underside with whitish or yellow-brown diagonal streak. Female: Ground colour varying from orange to brown. Postmedial and medial line diffuse, dark or pale brown. Apical third of forewing and terminal area of the hindwing concolorous with transverse lines. Hindwing underside more contrasting than upper side, also with whitish dia­ gonal streak, apical area of forewing and hindwing underside freckled with whitish blotches. 472

Fringes of both sexes chequered. Antennae bipectinate in ♂, filiform in ♀. Frons smoothscaled, in ♂ with mixed brown and pale sandy scales, in ♀ more reddish brown. Tegulae greyish to brown. Abdomen covered with brown and beige scales. Hindtibia with 2+2 spurs. Variation: Highly variable. Extent of dark brown and whitish or yellow-brown areas in male and of orange and pale brown areas in females varying considerably. Extreme forms are uniformly brown in male and nearly brown in female. Leraut (2009) examined a large number of specimens from different regions and came to the conclusion that no population can ‘justify a status of separate subspecies’. We can follow his opinion. Male genitalia: Uncus sclerotized, tip round, lateral margins with one rounded process each. Gnathos arms fused, ending in long tapered, slightly bent spur, on inner side serrate. Valva short, thick, apex round, membranous, setose. Costa bears a sclerotized, setose plate. Sacculus wide, with group of spines directed towards apex. Saccus long, rounded. Aedeagus slender, with obtuse-angled bend. Proximal ending spherical. Without cornuti. Female genitalia: Papillae anales membranous, oval, setose. Apophyses posteriores long, slightly bent, near papillae anales thickened. Apophyses anteriores short, less than 1/3 of the length of the latter. Tergum 8 very large. Ostium bursae stoutly sclerotized. Ductus bursae slender, membranous. Corpus bursae membranous, slightly striated. Without signum. Distribution and abundance: Eurasiatic. In Europe from the Urals to westernmost Europe and from northern Fennoscandia to Spain, southern France, the Alps and the Balkan peninsula including northern Greece. The map in Leraut (2009) erroneously shows a distribution all over the Iberian peninsula, in Portugal, all over Italy and in northern Fennoscandia. – Common, now and then occurring in larger numbers, sometimes even with mass occurrences, known as a potential serious pest in forestry. – Outside Europe from Turkey and Armenia through northern Kazakhstan, southern Siberia to Buryatia and Transbaikalia. In Morocco a single record from the Rif Mountains (Rungs 1981).

473

Phenology: Univoltine. In southern Scandinavia from mid- or late May to late July, in south-­ western Germany from early May to late July (Ebert 2003), in Spain from early April to early August at the highest altitudes and in the Alps from mid-April to mid-July (Kerschbaum & Pöll 2010). Larva from late May to early September (Ebert 2003). Overwintering as pupa. Both sexes mainly active at night, readily attracted to light; males flying also by day in sunshine. Biology: Larva oligophagous. Found on Pinus strobus, Pinus sylvestris and Pseudotsuga menziesii (Ebert 2003). Pupation below the surface of the soil. Habitat: Silvicolous. Natural and anthropogenic pine forests on sand and limestone, peat bogs, moors, gardens and parks with pines. From sea-level, in the Pyrenees up to 2000 m. Similar species: No similar species in Europe. Females of the Moroccan Ifrania michaellarius (Rungs, 1941) sometimes similar, the latter with obtusely angled postmedial line. Genetic data: BIN: BOLD:AAC7293 (n=36 from Finland, United Kingdom, Netherlands, Germany, Austria, France, Italy). Genetically homogeneous. Nearest species: B. vestalis Staudinger, 1897 from China (1.6%).

Megametopon Alphéraky, 1892 Megametopon Alphéraky, 1892, in Romanoff Mém. Lépid. 6: 58, figs. Type species: Megametopon piperatum Alphéraky, 1892. Contropis Wiltshire, 1967, Beitr. Naturk. Forsch. SüdwDtl. 26: 164, pl. 3, fig. 32, pl. 15, figs 67, 67a. Type species: Contropis tagana Wiltshire, 1967. New synonym (see Remarks).

Diversity and distribution: Three species in the Central and Eastern Palaearctic region. External characters and abdomen/Genitalia: See species description below. Remarks: Megametopon has not been included in a multi-gene analysis so far. Systematic position within Boarmiini tentative, awaiting further study. In Viidalepp (1996) placed next to Tephronia + Eumannia. The peculiar structure on the male frons (circular sclerite with unscaled central projection) and the supposedly brachypterous females clearly suggest synonymy between the central Asian genera Megametopon and Contropis. László (2001) in his study on the genus Contropis suggests relationships with Agriopis bajaria, based on male genitalia, not supported, however, by COI barcodes (distance 9.3%). Some examined genitalia slides not even excluding synonymy of the type species at species level, but preliminarily we still validate Megametopon modestaria (Püngeler, 1914), comb. n. and Megametopon modestaria tagana (Wiltshire, 1967), comb. n.

167. Megametopon grisolaria (Eversmann, 1848) Fidonia grisolaria Eversmann, 1848: Bull. Soc. imp. Nat. Moscou 21 (3): 225 (Russia: Steppes between the lower Volga and the Ural rivers). Syntypes 8♂. Unavailable names (incorrect subsequent spelling): griseolaria: Viidalepp (1988, 1996).

External characters and abdomen: Male: Wingspan ♂ 20−26 mm. Forewing ground colour grey to brownish grey, more or less speckled dark grey, pattern poor in contrast. Post­ medial line dark grey, usually dotted, curved, diffuse. Antemedial line black, often dotted, obtusely angled near costa. Medial line indistinct or absent. Subterminal line reduced to diffuse blotches. Terminal line grey, fringes of ground colour. Discal spot distinct. Hindwing: 474

Ground colour and fringes as in forewing, less speckled, without lines, discal spot indistinct. Antennae bipectinate. Collar, thorax and abdomen of ground colour. Frons of males smoothly scaled, with circular sclerite with central projection as in Dasypteroma thaumasia and Central Asian Megametopon modestaria (Püngeler, 1914). Margin of circular sclerite and projecting centre almost without scales, brown. Hindtibia with 2+2 spurs. Female brachypterous (Viidalepp 1988a). Variation: Very little. Male genitalia: Relatively small. Uncus sclerotized, triangular, slightly setose, apex ventrally bent, with very small double tip. Gnathos ventrally fused, apex wide, rounded, sclerotized. Valva simple, narrow, slightly setose, medially constricted. Apex of costa partly overlapped with apex of sacculus. Juxta developed as plate with concave lateral margins, apically bilobed. Saccus weakly concave. Aedeagus nearly cylindrical, straight, short, apex round, caecum blunt. Vesica with slender, weakly sclerotized cornutus. Female genitalia: Unknown. Distribution and abundance: West Turanian. In Europe restricted to south-eastern European Russia. – In Europe apparently rare, however, recently recorded in numbers in southern and western Kazakhstan, there more common (Gorbunov 2011). – Outside Europe only known from Georgia, Kazakhstan and Uzbekistan. Records from (mountainous) distribution areas in Uzbekistan and north-eastern Kazakhstan perhaps referring to the sister species M. modestaria.

Phenology: Univoltine. From early September to early October. Flight period apparently short, perhaps only lasting two weeks (Gorbunov 2011; K. Nupponen pers. comm.). Males active at night, attracted to light, wingless females probably sitting in low vegetation at night. 475

Biology: Larva probably feeding on wormwood, Artemisia absinthium (Gorbunov 2011). Habitat: Xerothermophilous. Inhabiting lowland steppes, semi-deserts and deserts. Similar species: Males of Dasypteroma thaumasia with similar wing pattern and structure of frons, but pattern with more contrast. Males of Central Asian Megametopon modestaria with similar wing pattern and structure of frons, see Remarks to genus. Genetic data: BIN: BOLD:ACP2974 (n=4 from south-western Kazakhstan and Turkmeni­ stan). Genetically heterogeneous, populations from south-western Kazakhstan and Turkmenistan ­diverging by 1.4%. Nearest species: Calamodes subscudularia (5.4%).

The ‘Tephronia group of genera’ Diversity and distribution: The ‘Tephronia group of genera’ consists of three closely related genera (Tephronia, Eumannia, Phyllometra), including 27 species in the western Palaearctic region, two further species are distributed in the transitional zones towards the Afrotropical region (Ethiopia, Djibouti, and Yemen). In addition to the above mentioned three European genera, there is a group of central Asian species which apparently is closely related, currently combined with the two genus names Autotrichia Prout, 1934 (four species described) and Pleotrichia Vasilenko, 2005 (monotypic). Relationship between Autotrichia and Phyllo­ metra was earlier suggested by Prout (1934) and Wehrli (1953). For more details see under genus Phyllometra. Material studied: More than 2000 Tephronia/Eumannia/Phyllometra specimens were examined for this study, the analysis was furthermore based on more than 170 genitalia slides, as well as on 190 DNA barcodes. Previous taxonomy: Prout’s (1915) diagnosis for Tephronia highlights the “smoothly scaled frons, palpi small, tender, proboscis vestigial, sometimes absent, ♂ antennae bipectinate, hindtibia only with terminal spurs ([accidentally: see under T. sepiaria ‘variation’] absent in fingalata), wings long, with long cell. Forewing in both sexes with long fovea, veins R1 and R2 of forewing fused, R3 and R4 diverging close to apex, sometimes fused, too. Hindwing with Sc close to Rs+M1 at the distal part of cell, Rs and M1 stalked, M2 weak, but present. Larva thick, tapering at both ends. Head small, flat, posteriorly rounded, warts prominent. On lichens, exclusively palaearctic. Incertae sedis.” Prout’s (1915) diagnosis for Mannia (= Eumannia) states: “hindtibia with all spurs, costalis of hindwing anastomosing, cell sometimes less long, R1 on shorter stalk (in oxygonaria separate).” Note: the taxon oxygonaria was transferred later to genus Habermania Viidalepp, 1988. Prout’s (1915) diagnosis for ‘Egea’ – now Phyllometra – includes the following characters: “small moths, wings – especially hindwing – long, in female narrow and poorly developed … palpi short, antennae of male bipectinate, with long branches. Hindtibia with four spurs, discal cell of forewing long, second subcostalis [R2] absent. On hindwing the second radiale [M2] weak, as a transition to the subfamily Geometrinae [Prout refers to what today is validated as Ennominae]”. Note: Prout (1915) still (mis)placed this genus in the subfamily Oenochrominae, transferred to Ennominae in Djakonov (1927). Prout’s discrimination between Tephronia and Eumannia needs to be modified and supplemented since the presence of just the terminal spurs on the male hindtibia is restricted to 476

T. sepiaria and T. nuragica and since of all the other species (with four male hindtibial spurs) just two are showing the venation characters mentioned by Prout (l.c.) for Eumannia. Even in the last 20 years the taxonomy and nomenclature of the European species of the ‘Tephronia group’ (s.l.) was treated quite different by various authors (cf. Tab. 6): Table 6: Nomenclature and taxonomy of European species and subspecies of the ‘Tephronia genus-group’ as validated and presented in recent literature covering the whole fauna of Europe; only those names are presented which were temporarily or continuously regarded as valid at species or subspecies level. * = supposedly extralimital; FaEu = Fauna Europaea (Hausmann et al. 2004; 2011a); GME6 2019 = present volume. Müller 1996

Scoble 1999

FaEu 2004

Leraut 2009

GME6 2019

Tephronia sepiaria

Tephronia sepiaria

Tephronia sepiaria

Tephronia sepiaria

Tephronia sepiaria

subsp. carieraria

(syn. carieraria)

(syn. teriolica)

(syn. teriolica)

subsp. teriolica

Tephronia cremiaria

subsp. chapmani

subsp. chapmani

(syn. chapmani)

(syn. chapmani)

subsp. cremiaria

(syn. cremiaria)

(syn. cremiaria)

(syn. cremiaria) Tephronia nuragica

Tephronia codetaria

Tephronia codetaria

Tephronia codetaria

[Tephronia codetaria*]

Tephronia codetaria*

subsp. sicula

Tephronia sicula

subsp. sicula

subsp. sicula

subsp. lhommaria

subsp. lhommaria

Tephronia lhommaria

Tephronia lhommaria

subsp. cleui

subsp. cleui

(syn. cleui)

subsp. cleui

subsp. diniensis

(syn. diniensis)

Tephronia cyrnea

Tephronia cyrnea

(syn. cyrnea)

Tephronia cyrnea

(syn. tonnara) Eumannia lepraria

Eumannia lepraria

Tephronia lepraria (par- Eumannia lepraria tim)

Tephronia lepraria

Eumannia psyloritaria

Eumannia psyloritaria

Eumannia psyloritaria

Tephronia psylo­ritaria

Eumannia psyloritaria

Tephronia theo­philaria

Tephronia lepraria (partim) Tephronia espaniola

Tephronia espaniola

Tephronia espaniola

Tephronia espaniola

Tephronia espaniola

Tephronia oranaria

Tephronia oranaria

Tephronia oranaria

Tephronia oranaria

Eumannia oranaria

subsp. castiliaria

subsp. castiliaria

subsp. castiliaria

subsp. castiliaria

subsp. cebennaria

subsp. cebennaria

subsp. hubertcleui

(syn. hubertcleui)

subsp. alpinaria

subsp. alpinaria

subsp. cebennaria

Eumannia oppositaria

Eumannia oppositaria

Eumannia oppositaria

Eumannia oppositaria

Eumannia opposi­taria

Phyllometra culminaria

Phyllometra culminaria

Phyllometra culminaria

Tephronia culminaria

Phyllometra culminaria

Tephronia gracilaria

Phyllometra gracilaria

Tephronia gracilaria

Tephronia gracilaria

Phyllometra gracilaria

477

Extensive integrative taxonomic studies performed in the preparation of this volume showed that the three genera Tephronia, Eumannia and Phyllometra not only diverge into different genetic clusters (COI, cf. Text-fig. 200) but can also be discriminated morphologically as presented in Tab. 7: Table 7: Differential features in morphology and activity pattern allowing discrimination of the genera Tephronia, Eumannia and Phyllometra.

hindwing coloration fringe wingspan in ♀ hindwing veins Sc and Rs+M1 ♂ antennal branches ♂ genitalia, valva process ♀ genitalia, signum main activity

Tephronia

Eumannia

Phyllometra

pale unchequered normal separate 0.4–0.6 mm at centre spinose ring or absent nocturnal

pale unchequered normal fused 0.2–0.4 mm near apex bipartite sclerite nocturnal

darker chequered reduced separate 0.7–0.8 mm at centre very small or absent diurnal

Text-fig. 200. Maximum Likelihood tree for the COI 5’ gene fragment (>500bp) for 126 specimens of the European species of the ‘Tephronia genus-group’ (Tephronia, Eumannia, Phyllo­ metra) and Ithysia barcoded from Europe and adjacent countries, with significant bootstraps (>50) indicated on the branches (MEGA6; 500 replicates; Kimura-2–parameter model; uniform rates). Clusters for species and diverging populations collapsed to triangles (width = sample size; depth = genetic variation).

478

Differential external characters and abdomen: Common characters for the ‘Tephronia genus-­group’: Frons flat. Eyes small. Palpi of both sexes very short. Antennae of ♂ bipectinate, of ♀ filiform. Proboscis vestigial (short and weak) or absent. Sternum A8 of male simple, without modifications. Tympanum in both sexes very small, ansa a narrow stick. Forewing R1 and R2 fused, Sc and R1 usually not reaching costa. Hindwing often with weak, but tubular M2, a very unususal character in Ennominae. Genitalia: Common characters in ♂ genitalia for the ‘Tephronia genus-group’: spinulose projection from centre of valva; uncus sub-truncate; aedeagus with one cornutus (exception: T. espaniola). The very similar build of male genitalia seem to support the subordination of Eumannia and Phyllometra under Tephronia. However, external morphological features (e.g. venation, length of male antennal branches), female genitalia and phylogenetic hypothesis based on DNA barcode data reveal a clear pattern of separable groups classified here at genus level. Genetic data: DNA analysis of the 190 DNA barcodes (COI 5’) of the ‘Tephronia genus-group’ (cf. Fig. 200) reveals comparatively deep divergences between the three genera Tephronia, Eumannia and Phyllometra as defined here. The analysis included 126 DNA barcodes referring to European species and 89 specimens from the territory of Europe.

Tephronia Hübner, 1825 Tephronia Hübner, [1825] 1816, Verz. bekannter Schmett.: 318. Type species: Geometra cineraria Denis & Schiffermüller, 1775, by subsequent designation by Fletcher (1979). G. cineraria is a junior synonym of Tephronia sepiaria (Hufnagel, 1767). ‡ Mniophila Boisduval, 1840, Genera Index meth. eur. Lepid.: 200. Type species: Geo­metra cineraria Denis & Schiffermüller, 1775, by subsequent designation by Guenée (1858). G. cineraria is a junior synonym of Tephronia sepiaria (Hufnagel, 1767). Junior homonym of Mniophila Stephens, 1831 [Coleoptera]. Gymnospile Gumppenberg, 1887, Nova Acta Acad. Caesar. Leop. Carol. 49: 324 (key), made available in nr 59: 373. Type species: Geometra cremiaria Freyer, 1838, by subsequent monotypy. G. cremiaria is a junior synonym of Tephronia sepiaria (Hufnagel, 1767). Unavailable names (incorrect subsequent spellings): Tephrina: Wardikjan (1985); Trephonia: Wolfsberger (1966); Tephrosia: Didmanidze (2002).

Diversity and distribution: Genus Tephronia with 18 species in the western Palaearctic region, two further species are distributed in the transitional zones towards the Afrotropical region (Ethiopia, Djibouti, and Yemen). Sister genera Eumannia and Phyllometra closely related and showing similar male genitalia. One species was recently described from Sardinia by Fiumi et al. (2013) (T. nuragica), another by Tautel (2015) (T. tonnara, here recognized as a synonym of T. cyrnea). For potential synonymy of Autotrichia Prout, 1934 and Pleotrichia Vasilenko, 2005 see above under ‘The Tephronia genus group’. Order of species: The genus Tephronia in the current concept includes nine European species. Two species groups and three more isolated species can be distinguished when considering feature combinations involving the number of male hindtibial spurs, shape of signum in female genitalia, DNA barcodes and venation: The sepiaria species-group: Differential characters: female signum developed as a single spinose ring, ♂ hindtibia with two spurs only, except for T. codetaria and T. bytinskii. Euro­ pean species included: T. sepiaria, T. nuragica. Outside Europe, this group includes one undescribed taxon from Turkey. Genitalia of T. bytinskii (Wehrli, 1939), comb. n. (in ­Scoble 479

1999 combined with Eumannia) from Canary Islands and Morocco (with melloulica ­Leraut, 2009, syn.n. which has to be established as a new synonym, because it has identical genitalia and overlapping DNA barcodes; cf. remarks to T. sepiaria) are similar to those of T. sepiaria but it is characterized by four male hindtibial spurs and COI barcodes closer to the lhommaria species-group. T. codetaria (in Europe subsp. sicula) in female genitalia shows the single spinose ring of the previous group, but in the number of male hindtibial spurs corresponds to the following species and species groups. The lhommaria species-group: Very similar in their DNA barcodes. Differential characters in morphology: Postmedial line of forewing deeply concave. ♂ hindtibia with four spurs, female ductus bursae sclerotized, corpus bursae usually without signum but in T. psylori­ taria with a spinose ring as in the sepiaria group. Species included: T. lhommaria, T. cyrnea, T. lepraria, T. psyloritaria. T. theophilaria Hausmann sp. n.: female genitalia with very long female corpus bursae, signum absent, male hindtibia with four spurs, in the DNA barcode rather isolated. T. espaniola: male (e.g. large aedeagus) and female genitalia (e.g. long female corpus bursae, signum absent) unique within the genus, male hindtibia with four spurs, in the DNA barcode rather isolated. Differential external characters and abdomen: Common characters for the genus as defined here: Frons flat. Eyes small. Palpi of both sexes very short. Antennae of ♂ distinctly bipectinate in basal half, shortly bipectinate in distal half, longest branches 0.4–0.6 mm. Antennae of ♀ filiform. Proboscis absent, rarely short and weak. Male hindtibia with two or four spurs. Sternum A8 of male simple, without modifications. Tympanum in both sexes very small, ansa narrow stick. Forewing venation with long cell, Sc and R1 not reaching costa, R2 absent, R3 and R4 diverging close to apex, or R3 sometimes absent. Hindwing with Sc close to Rs+M1 but not fused, Rs and M1 usually distinctly stalked. M2 weak, usually present. Male genitalia: ♂ genitalia with little differentiation at species level, the conspicuous projection at the centre of valva subject to intraspecific variation and only in a few cases relevant for species discrimination; the most valuable differential characters are in the shape of the (single) cornutus, length of aedeagus and shape of saccus. See also characters presented under ‘Order of species’ (above). Female genitalia: ♀ genitalia with more significant differential traits (at species-level) than in males, diagnostic features mainly in the length of corpus bursae, presence and shape of signum, as well as shape, sclerotization and appendices of ductus bursae. See also characters presented under ‘Order of species’ (above). Biology lichens.

and immature stages:

Larva lichenophagous, with well camouflaged coloration on

Genetic data: DNA barcode data suggest that the North African species T. duercki Reisser, 1933, T. fatimaria Bang-Haas, 1906 (with synonym ismailaria Oberthür, 1913; in Scoble 1999 combined with ‘Eumannia’), T. praerecta Wehrli, 1933 and T. nigrolineata Zerny, 1934 evolved on isolated lineages from the European species and species-groups, whilst T. aethiopica Herbulot, 1983 from Ethiopia, and T. yemenitica Hausmann, 2006 from ­Yemen are closer to the European clusters. T. fatimaria is characterized by a double spinose ring in female genitalia, T. duercki with scattered spinules, whilst T. praerecta and T. nigrolineata 480

show a very long corpus bursae without signum, though male hindtibiae with two spurs suggest certain relationships of these two species with T. sepiaria.

168. Tephronia sepiaria (Hufnagel, 1767) Phalaena sepiaria Hufnagel, 1767: Berlin. Mag. 4 (5): 516 (Germany: Berlin). Syntypes (lost). Geometra cineraria Denis & Schiffermüller, 1775: Ankündung syst. Werkes Schmett. Wiener­ gegend: 100 (Austria: Vienna). Syntypes (lost). Junior synonym. ‡ Boarmia corticaria: Duponchel (1829): in Godart & Duponchel: Hist. nat. Lep. Papillons Fr. 7 (2): 388. Misidentification nec Geometra corticaria Denis & Schiffermüller, 1775 (identity: Parectropis similaria). For synonymy cf. Prout (1915), Fiumi et al. (2013); Flamigni et al. (2016). Duponchel’s figure may also refer to Tephronia lhommaria, but nevertheless, the name is nomenclaturally unavailable. Boarmia cremiaria Freyer, 1838: Neuere Beitr. Schmettkde. 3 (43): 92, pl. 258, figs 1a-e (Germany: Hannover). Syntypes. Validated at subspecies rank (Scoble 1999), but downgraded to synonymy by Redondo & Gaston (1999), Flamigni et al. (2016) and others. The text of the description is dated as ‘1838’, of the plate as ‘1839’ according to Heppner (1982). Mniophila carieraria Herrich-Schäffer, 1848: Syst. Bearb. Schmett. Eur. 3: 93, fig. (‘suppl’) 214 (southern France). Holotype ♀. In Leraut (2009) and Flamigni et al. (2016) validated at subspecific rank with fingalata and teriolica as synonyms. Herewith downgraded to synonymy (syn. n.), see below under “variation”. ‡ Tephrina [sic!] destituta (auct. nec?) Walker, 1870: Entomologist 5: 153 (Egypt: Cairo). Syntypes (NHMUK). Synonym according to Scoble (1999) and Flamigni et al. (2016), synonymy, however, rejected here, requiring further study (see under distribution). The original genus “Tephrina” is a synonym of Isturgia, Macariini). Mniophila fingalata Millière, 1873: Cat. Rais. Lépid. dept. Alpes-Maritimes (2): 159 (France, Alpes Maritimes: Saint-Martin-Lantosque; Berthemont). Syntypes (lost). In Prout (1915) and Lhomme (1930) validated at species rank (misspelled to fingalaria), later recognized as junior synonym. Tephronia sepiaria f. chapmani Prout, 1915: in Seitz, Gross-Schmett. Erde 4: 381 (Spain: Avila, Navalperal, Sierra de Guadarrama). Syntypes 7♂ (NHMUK). Nomenclaturally available, described as ‘form’ with geographical background. Tephronia sepiaria var. teriolica Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 555 (Italy, Alto Adige: Bolzano; Trentino: Torbole). Holotype ♂ (ZFMK, examined). Valid at subspecies rank (Scoble 1999), downgraded to synonymy of carieraria in Flamigni et al. (2016). Tephronia sepiaria var. chapmani Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 Suppl.: 555, raising Prout’s infrasubspecific name to an available rank. Validated at subspecies rank also by Scoble (1999), not accepted here due to transitional patterns between the populations. Unavailable names: (infrasubspecific): nigra Rebel, 1909. – (incorrect subsequent spelling): fingalaria: Millière (1874). – (misidentification): oppositaria: sensu Beshkov & Nahirnic (2017) nec Hufnagel, 1767.

External characters and abdomen: Wingspan ♂♀ 18–22 mm, southern French females sometimes up to 24 mm, specimens of the second generation 15–17 mm only. Forewing dark grey-brown, hindwing somewhat paler. Postmedial line of forewing double s-shaped, slightly curved towards wing base at costa and inner margin, not deeply curved at centre. Antemedial line comparatively straight in central Europe, in the south often strongly bent. Female larger, forewings broader and less pointed, female pattern with more contrast. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ with one pair of hindtibial spurs.

481

Variation: Pale specimens with dotted or slightly dentate postmedial line have often been attributed to cremiaria Freyer, 1838 which has repeatedly been considered as a distinct species, but is not more than an individual (infrasubspecific) form, without any difference in male and female genitalia and sometimes collected together with the nominotypical form. The same conclusion was drawn for Italy by Flamigni et al. (2016) and for Spain by Redondo & Gaston (1999) while Leraut (1997) and Robineau (2007) list cremiaria as valid subspecies in northern and central France (see Remarks). Prout (1915) described cremiaria from northern Italy, France and Spain as “paler than sepiaria, veins partially darkened; ante­ medial line usually more rounded at costa; medial shade usually well developed, often also on hindwing; postmedial line straighter, broken into dots on all wings; terminal line usually absent; palpi and proboscis somewhat better developed”. None of these characters allows a satisfying and clear separation from nominotypical forms which may fly in the same localities. The cremiaria forms prevail in some regions of the Iberian peninsula, southern France and are similar to the taxonomically unclear populations from Turkey (see below). Populations from southern France to western and southern Alps (including Liguria, Pie­ monte, Valesia, Alto Adige) (in literature often named as carieraria, fingalata or teriolica) with darker, often blackish coloration of wings (f. nigra; f. obscura), as a consequence the pattern (transverse lines) is more diffuse. Hindtibial terminal spurs reputedly ‘absent in fingalata’ (Prout 1915), which has to be regarded as an artifact, as no such specimen could be traced in the extensive examined material. Since wing colouration and pattern underlies variation, sharp distributional borders are lacking and no correlated features in genitalia nor in DNA barcodes could be found, subspecies status is not accepted here for these populations. Male genitalia: Uncus (sub-)truncate, rounded, or slightly concave posteriorly (the last often occurring in the Apennine peninsula). Saccus a digitate projection. Juxta comparatively broad, with long posterior process, sometimes bifid at tip. Central process of valva distinct, often digitate. Length of aedeagus 1.4–1.65 mm. Cornutus short (0.37–0.45 mm), stout, usually bent at about 1/2, with large and broad basal sclerite, the latter usually 0.7–1.0 times length of the narrow and straight distal part. Female genitalia: Antrum large, triangular (funnel-shaped). Ductus bursae very long (2.1–2.2 mm), often dilated posteriorly (regularly in the eastern distribution from Austria to Greece), towards corpus bursae usually narrow. Corpus bursae globular, with ring-shaped, spinose signum. Appendix bursae lacking. Distribution and abundance: Sub-Mediterranean (European). North of the Alps very local from France to Germany, Czech Republic and Slovakia (Las˘tu˚ vka 1998). Extinct in Belgium (de Prins 1998), Poland (Malkiewicz 2012) and Lithuania (Müller 1996). In southern ­Europe widely distributed from Iberian peninsula to Balkan countries and Ukraine (­Kostjuk 2004; Crimea: Savchuk pers. comm.). In Sardinia and Corsica replaced by the sibling species T. nuragica (Fiumi et al. 2013; Flamigni et al. 2016). Despite repeated records (Mariani 1939; Raineri & Zangheri 1995; Müller 1996) not present in the Italian provinces of Sicily and Sardinia (Fiumi et al. 2013; Flamigni et al. 2016). In one dissected male with two hind­tibial spurs from ‘Sicily’ in coll. ZSM (’25.V.1917’; gen.prp. ZSM G 6537; cf. Hausmann 1993a) mislabelling of the collecting site cannot be completely excluded. – Abundance strongly declining in the north: extinct in Lithuania since 1901 (Sulcs & Viidalepp 1972), Poland since 19th century (Malkiewicz 2012) as well as in Belgium (de Prins 1998; ­Lambillion 1909), many regions of Germany (Gelbrecht 1999; Trusch et al. 2012) and Austria (­Huemer et al. 2009; Kerschbaum & Pöll 2010). In southern Europe common, often 482

found in ­numbers. – Outside Europe without verified records clearly suggesting conspecifity. In Literature repeatedly recorded from Turkey (e.g. Prout, 1915; Flamigni et al. 2016), but these populations probably refer to another species, as suggested by the genetic data (see below) and morphology. The same probably applies to the records from Georgia (­Didmanidze 2002) and Armenia (Wardikjan 1985, with genitalia figured). Records from Palestine (Prout 1915) erro­neous, referring to Eumannia fumosata Hausmann, 1995. Reputedly also in North ­Africa, e.g. mentioned by Rungs (1981) for the Middle Atlas in Morocco, awaiting confirmation (see Remarks). The synonymy of the Egyptian type specimen of “Tephrina destituta Walker, 1870” – as suggested by Scoble (1999) – is certainly erroneous and may refer to a separate species (see above under nomenclatural remarks and the remarks to the Egyptian ‘T. minutaria’ under Eumannia oranaria). Records for Canary Islands, erroneous, referring to T. bytinskii (Wehrli, 1939).

Phenology: Uni- or bivoltine. In the north univoltine from early June to late August. South of the Alps sometimes emerging from mid-April (in southern Portugal even from late March) and with single specimens from late August to mid-October, belonging to a partial second generation, which is regular in much of the Iberian peninsula, southern Italy and parts of the Balkan peninsula. Overwintering as larva. Adults active at night, readily attracted to light. Biology: Larva oligophagous, feeding on lichens on trees, dead-wood, fences or shrubs but also on walls and rocks. Recorded on Parmelia, ‘Bryopogon’, Sticta, Chrysothrix (= Byssus) candelaris (Flamigni et al. 2016), lichens growing on shrubs and trees (often Juniperus, also Populus, Pyrus, Crataegus recorded: Bergmann 1955; Flamigni et al. 2016). According to an old, doubtful record also feeding on mosses (Millière 1876). Thiele (2001) describes and illustrates the rearing of larvae from the type locality of “cremiaria”. Pupation usually in a cocoon between or under the lichens (Forster & Wohlfahrt 1981; Thiele 2001). 483

Habitat: Xerothermophilous. Forest fringes, hedges, orchards, parks, gardens (near wooden buildings and fences), often in warm, dump sites with much dead wood. In southern Europe in various open habitats with scattered trees and bushes (e.g. Mediterranean macchia), on dry and rocky sites and in scattered oak forest. In central Europe restricted to the lowlands, in southern Europe from sea-level up to 800 m, rarely up to 1600 m in the southern Alps, and up to 1800 m in southernmost Europe. Similar species (see Text-figs 201–203, 204–206 and 210–212): T. sepiaria is, in the area of its distribution, the only species of this genus with two male hindtibial spurs. The same character is only found in the allopatric sister species T. nuragica from Corsica and Sardinia, and in North African T. nigrolineata Zerny, 1934 and T. praerecta Wehrli, 1933. Correct identification is usually possible based on the wing pattern, which in T. sepiaria shows the forewing postmedial line curved towards the base at the inner margin. T. codetaria sicula differing by darker basal area and with strong differences in male and female genitalia. T. theophilaria differing by four male hindtibial spurs, forewings more brown without grey irroration, ante­ medial line at inner termen projecting towards termen, postmedial line of forewing more inclined towards apex, hindwing pale with conspicuous postmedial line. Identification of females sometimes requires dissection. In male genitalia the shape of cornutus allows discrimination from the allopatric sister species T. nuragica, in the latter showing a shorter and smaller basal sclerite. Female genitalia with ring-shaped signum, which is also present in (allopatric) T. nuragica, T. codetaria sicula and T. psyloritaria. T. c. sicula differs from T. sepi­ aria in the divided corpus bursae with two parts in rectangular position. T. nuragica differs in the shorter ductus bursae, which is more dilated towards corpus bursae. T. lepraria with four male hindtibial spurs, postmedial line more strongly concave, male antennal branches shorter. Genetic data: BINs: BOLD:AAD2603 (n=25 from Germany, Austria, France, Italy, Spain, Greece), BOLD:ABV4483 (n=6 from Turkey), both BINs diverging by 2.4%, meriting further taxonomic attention. Genetically homogeneous within the BINs. Nearest species: T. nuragica (2.5%). Remarks: In Leraut (2009) the taxon praerecta Wehrli, 1933 was downgraded to subspecies rank under T. sepiaria, without checking the type specimen. After examination of the type specimen and additional materials we restore the status of T. praerecta Wehrli, 1933 (stat. n.) to species rank, because of striking differential features: male genitalia with longer aedeagus (2 mm), and longer, narrower cornutus (0.6 mm), female with very long corpus bursae without signum, genetic distance 6.5%. Similarly, Leraut’s (2009) synonymization of nigrolineata Zerny, 1934 with praerecta is wrong, as suggested by DNA barcodes (diverging by 6.1% from the latter) and wing pattern: T.  nigrolineata with forewing antemedial line more convex, postmedial line towards costa straight and oblique towards apex (not bent), forewing terminal area without dark shadow, postmedial line on hindwing more diffuse. The populations from Morocco to Libya were still reported as “T. sepiaria subsp. praerecta Wehrli, 1933” in Flamigni et al. (2016), but the North African T. praerecta is clearly a separate species genetically and genitalically. The taxon melloulica Leraut, 2009 (described from Morocco) shows certain similarities in genitalia with T. sepiaria, e.g. female corpus bursae with a ring-shaped spinose signum but ductus bursae is shorter than in T. sepiaria and even the male hindtibia with four spurs does not suggest a closer relationship. In male genitalia the cornutus is long and slender. However, morphometrical analysis of T. bytinskii (Wehrli, 1939) from the Canary islands reveals conspecifity with melloulica Leraut, 2009, the latter is herewith downgraded to synonymy (syn. n.). Genitalia of both Canarian and 484

Moroccan populations are identical and DNA barcodes are overlapping between Moroccan and Canarian populations at a common maximum variation of only 1.2%. The taxon cremiaria Freyer, 1838 had been listed at species rank by many authors (e.g. Prout 1915; Raineri & Zangheri 1995; Müller 1996; Parenzan 1994; 2002; Thiele 2001) at subspecies rank by others (e.g. Scoble 1999; Leraut 1997), but the type locality is very near to that of T. sepiaria. For the classification as infrasubspecific form, see above under variation.

169. Tephronia nuragica Fiumi et al., 2013 Tephronia nuragica Fiumi, Flamigni, Zilli & Hausmann, 2013: Quad. Stud. Not. Storia Nat. Romagna 38: 201, figs 3–9 (Italia, Sardinia: Usellus (Oristano), Santa Lucia). Holotype ♂ (ZSM, examined). ‡ Tephronia sepiaria: sensu auct. nec Hufnagel, 1767 (misidentification).

External characters and abdomen: Wingspan ♂ 16–23, ♀ 19–23 mm. Wing pattern often vague and not well expressed. Postmedial line of forewing double s-shaped, slightly curved towards wing base at costa and inner margin, not deeply curved at centre. Antemedial line curved towards base at forewing costa. Postmedial line absent from hindwing or indistinct. Female larger, forewings broader and less pointed. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ with one pair of hindtibial spurs.

Variation: Ground colour ranging from pale grey to brown or dark grey. Transverse lines (antemedial, medial and postmedial lines) of forewing of variable intensity.

Male genitalia: Uncus (sub-)truncate, edges rounded. Saccus narrow, shortly digitate. ­Juxta comparatively narrow, with long posterior process. Central process of valva triangular, comparatively broad. Aedeagus bent, length 1.5–1.55 mm. Cornutus 1/4–1/3 length of aedeagus (0.37–0.45 mm), on average narrower than equivalent in T. sepiaria, and more curved. Basal sclerite of cornutus small and narrow, approx. 0.25–0.4 times length of narrow, distal part. Female genitalia: Antrum large, triangular (funnel-shaped). Ductus bursae shorter (1.8 mm) than in T. sepiaria, towards corpus bursae dilated. Corpus bursae globular, with ring-shaped, spinose signum. Female genitalia from Corsica correspond well with those of Sardinian specimens (Tautel 2015; Flamigni et al. 2016). Distribution and abundance: Tyrrhenian, endemic to Sardinia and Corsica. – Widespread but not very common in Sardinia, with local occurrence in Corsica. Phenology: Bivoltine. From late June to mid-August (exceptionally earlier), second generation late August to late October. Probably overwintering as larva like the congeners. Adults active at night, readily attracted to light. Biology: Larval stages unknown. Larvae probably feeding on lichens like the congeners. Habitat: Xerophilous. Forest fringes, Mediterranean shrublands, pine forest with small patches of open terrain. From sea-level up to 1100 m. Similar species (see Text-figs 207–209): Allopatric T. sepiaria (see no. 168). In some localities in Sardinia occurring sympatrically at the same localities as T. cyrnea, males of the latter can easily be distinguished by the presence of four male hindtibial spurs. In wing pattern, 485

T. cyrnea shows a much deeper concave postmedial line of the forewing, the antemedial line is bent towards the wing base at the forewing costa rather than to the apex. On the hindwing the postmedial line is more contrasted. In female genitalia T. cyrnea lacks a signum and is characterized by a large, wrinkled, membranous appendix bursae. Genetic data: BINs: BOLD:ABX0094 (n=3 from Sardinia), BOLD:ADE3117 (n=2 from Corsica), both BINs diverging by 1.2%. Nearest species: T. sepiaria (2.5%).

Tephronia codetaria (Oberthür, 1881) Tephrosia codetaria Oberthür, 1881: Études ent. 6: 80, pl. 11, fig. 3 (Algeria: Sebdou). Holotype ♀ (ZFMK, examined, DNA barcode BC ZFMK LepTyp 0001_NGS). Tephronia gerini Leraut, 2009: Moths of Europe, Vol. 2, Geometrid Moths: 200, pl. 53, figs 13–14 (Morocco, Middle Atlas: Ifrane). Holotype ♀, MNHN (gen.prp. Leraut 10769). Herewith downgraded from species to subspecies rank, see Remarks. Unavailable names (incorrect subsequent spelling): cotedaria: Wehrli (1953: 561).

Remarks: Nominotypical subspecies non-European. The external morphology of the examined holotype specimen (ZFMK), its DNA barcode, and dissection of topotypic specimens clearly prove conspecifity with the European subsp. sicula. The nominotypical subspecies is restricted to North Africa, Algeria and Tunisia. Replaced by subsp. gerini Leraut, 2009, stat. n. in the Middle and High Atlas (Morocco), flying from April to August depending on altitude (Leraut 2009). Female genitalia as figured in the original description (fig. 104a) showing just slight differences from T. codetaria in the position of the signum ring in the corpus bursae (supposedly due to different position on the slide or different mating status). The other differential features mentioned by Leraut (l.c.: ‘larger and forewing transverse lines more sinuous’) not sufficient to claim species rank in allopatric populations of this genus (with much site-specific variation in size and wing pattern) and neither confirmed as being constant in 486

our examined material from the Middle and High Atlas. Thus, the taxon gerini is herewith downgraded to subsp. of T. codetaria (T. codetaria gerini stat. n.). For further arguments on this change of taxonomic status see also Genetic data under the following subspecies.

170. Tephronia codetaria sicula (Wehrli, 1933), stat. rev. Mannia codetaria sicula Wehrli, 1933: Amat. Papillons 6: 212 (Italia, Sicily: Madonie). Syntypes 1♂7♀ (ZFMK; 1♂ syntype ‘Typus’ and its genitalia slide no. 5473 examined). Validated at subspecies rank of T. codetaria (Scoble 1999), raised to species rank by Flamigni et al. (2016), herewith downgraded again to subspecies rank.

External characters and abdomen: Wingspan in Sicily ♂ 17–21 mm, ♀ 22–25, in central and southern Italy smaller, ♂♀ 16–19 mm. Forewing with dark basal area, strongly contrasting with medial area. Antemedial line of forewing usually strongly bent twice. Postmedial line deeply concave at centre, almost semicircular. Terminal area usually darker than medial area. Hindwing usually whitish with postmedial line bent near apex. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs. Variation: Variable in the expression of the various elements of the wing pattern. Nominotypical populations from North Africa and those from Middle Atlas (subsp. gerini) in habitus and external structure without constant differences from subsp. sicula. Male genitalia (analysis including slide of type specimen): Uncus usually bilobed and distinctly concave posteriorly. Saccus triangular, strongly projecting. Juxta comparatively broad. Central process of valva developed as short lobe. Aedeagus long and curved (length 1.45–1.8 mm, in nominotypical subspecies 1.25–1.45 mm). Cornutus long, curved, with narrow tip, length of cornutus 0.43–0.5 mm (in nominotypical subspecies 0.35–0.47 mm). Female genitalia: Antrum large, trapezoid (broadly funnel-shaped), posteriorly and anteriorly sclerotized. Ductus bursae short (1.2–1.5 mm), laterally dilated, very broad, of the same width as corpus bursae, joining the latter at 45–90° angle. Lateral extension of ductus bursae membranous, often triangular. Corpus bursae globular but linked with ductus bursae in its whole width. Signum ring-shaped, large, spinose. Distribution and abundance: Central Mediterranean. Endemic to Sicily and the Apennine mountains northwards to Abruzzo, Lazio, Umbria and Tuscany (cf. Parenzan 1994; Flamigni et al. 2001; 2016; Sciarretta & Raineri 2001). An isolated population also on the island of Pantelleria (Flamigni et al. 2016; one male dissected from coll. H. Fischer). Records for Sardinia (e.g. Parenzan 1994; Parenzan & Porcelli 2006) misidentified, referring to T. cyrnea. – Local, but sometimes abundant at the sites of occurrence. – Outside Europe from Algeria to Tunisia (nominotypical subspecies) and in the Middle Atlas in Morocco (subsp. gerini). Phenology: Uni- or bivoltine. In Europe usually univoltine from mid-June to late August, in the mountains somewhat later. Under favourable conditions in the southern lowlands exceptionally bivoltine with two overlapping generations from late May to early October. In North Africa bi- or even trivoltine from early April to early October. The occurrence of a third generation suggested by Wehrli (1953). Probably overwintering as larva like the congeners. Adults active at night, readily attracted to light.

487

Biology: Larval stages unknown for the European populations. Larvae of the nominotypical subspecies reputedly on lichens on Juniperus (Leraut 2009), requiring confirmation of the correct identification and taxonomic interpretation of the taxon in question (‘T. codetaria’). It is possible that this record refers to the old usage of the name codetaria for the French populations, now T. lhommaria, see under ‘biology’ of the following species. Habitat: Xerothermophilous, often in montane habitats. Mediterranean macchia, shrublands, forest fringes, often in canyons. Up to 1300 m in the Apennine peninsula and in Sicily, in the north preferably at lower elevations but rarely below 300 m, in Sicily not below 650 m (Flamigni et al. 2016; data in ZSM). On Pantelleria island collected at 350 m (coll. Fischer). In North Africa from sea-level up to 2000 m. Similar species (see Text-figs 201–203): T. sepiaria (see no. 168). In the European area of distribution, (allopatric) T. lepraria differs in the less concave centre of forewing postmedial line, in the paler basal area of forewing and in the double bent postmedial line of hindwing. In male genitalia the cornutus is much longer in T. lepraria, whilst in female genitalia of T. lepraria the ductus and corpus bursae are very long and narrow and signum is absent. Genetic data: BIN: BOLD:ABY3361 (n=5 from Sicily, southern and central Italy), with six other identical DNA barcodes from Algeria and Tunisia, including that of the holotype of T. codetaria. Nearest taxa: T. codetaria gerini Leraut, 2009 from Morocco (1.4%; BOLD: ACL9884) and an undescribed species from Israel, Jordan and Sinai (2.2%). Nearest species in Europe: T. nuragica (3.8%).

488

Text-figs 201–203. Diagnostic characters (indicated) for wing-pattern-based identification of Tephronia species on the Apennine peninsula and in Istria: Text-fig. 201. T. sepiaria (Hufnagel, 1767) (Italy, Lago di Garda, 27.vii.1990). Text-fig. 202. T. codetaria sicula (Wehrli, 1933) (Italy, Sicily, Madonie, syntype). Text-fig. 203. T. theophilaria Hausmann sp. n. (Italy, Abruzzo, Arischia near Pizzola/Aquila, 4.viii.1980).

171. Tephronia lhommaria (Cleu, 1928) Mannia codetaria var. lhommaria Cleu, 1928: Amat. Papillons 4: 91, pl. 1, figs 9, 10 (France, Hautes Alpes: Saint-Crepin). Syntypes (1♀ ‘cotype’ examined in ZFMK ‘Hautes Alpes: SaintCrepin; 6 VIII 25 e.l. lichens de Jun. thurifera’; one further ♂ with identical labels but without wings, gen.prp. ZFMK 5480). Validated at subspecies rank under T. codetaria (Scoble 1999), raised to species rank by Leraut (2009) and Flamigni et al. (2016).

489

Mannia codetaria var. melaleucaria Schwingenschuss, 1932: in Schawerda, Z. öst. Ent.Ver. 17 (2/3): 13, pl. 3, figs 11, 12 (Spain, Aragon: Albarracin). Syntypes ♂♀ (NHMW?), nomenclaturally available. Not listed in Leraut (2009), mentioned as potentially be valid at subspecies rank in Flamigni et al 2016. Here tentatively validated at subspecies rank. Tephronia codetaria cleui Wehrli, 1953: in Seitz, Gross-Schmett. Erde 4 (Suppl.): 558 (France, Ardèche: Bas-Vivarais). Syntypes (ZFMK, coll. Cleu/MNHN). Validated at subspecies rank (Leraut 1980; 1997; Scoble 1999), downgraded to synonymy by Leraut (2009), here re-­ established at subspecies rank following Tautel (2015). Tephronia lhommaria diniensis Leraut, 2009: Moths of Europe, Vol. 2, Geometrid Moths: 195, pl. 53, figs 1–3 (France, Provence: Digne). Downgraded to synonymy of cleui by Tautel (2015). ‡ Tephronia codetaria: sensu auct. nec Oberthür, 1881 (misidentification). Unavailable names: (incorrect subsequent spellings): lhommearia: Lhomme (1930), followed by Cleu (1933), Wehrli (1933; 1953), Herbulot (1963), Leraut (1980; 1997) and others; meloleucaria: Scoble (1999).

External characters and abdomen: Wingspan of nominotypical subspecies ♂♀ 18–24 mm, females on average somewhat larger and with broader wings. Nominotypical subspecies with grey suffusion over ground colour. Forewing with basal and terminal area usually darker than medial area. Postmedial line deeply concave at centre, almost semicircular. Hindwing usually whitish with postmedial line bent near apex. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs.

Subsp. cleui (from Ardèche and southernmost Alps reputedly smaller (Leraut 2009) but this statement refers to the smaller specimens of the second generations in bivoltine populations (♂♀ 16–20 mm). Typical populations from Dept. Ardèche with very pale, sometimes creamy white ground colour. In other southern French departments with basal and terminal area usually darker than medial area, perhaps without sharp borders from similar populations in the Iberian peninsula (subsp. melaleucaria). Apparently, sharp delimitation is also lacking from the nominotypical subspecies and in some localities different forms are mixed up from very pale specimens to specimens with dark irroration and dark wing pattern. Nevertheless the subspecies rank for cleui and melaleucaria is tentatively maintained here because of the differences from the nominotypical subspecies from the High Alps in phenology and habitat. Variation: Extent of dark suffusion somewhat variable within the same populations. Basal and terminal area sometimes of the same or similar colour as the medial area, e.g. more often in southernmost Iberian peninsula and in the French Dept. Ardèche. Male genitalia: Uncus (sub-)truncate, sometimes slightly concave posteriorly. Saccus digitate, projecting. Central process of valva broad, comparatively short. Aedeagus short (0.9– 1.1 mm), straight. Cornutus short (0.3–0.35 mm). Female genitalia: Antrum small, funnel-shaped. Ductus bursae short (0.75–1.0 mm), in the Iberian peninsula (subsp. melaleucaria) 1/2 – 2/3 length of corpus bursae. Ductus bursae broad, sclerotized, with very short, membranous constriction towards antrum, towards corpus bursae slightly conically dilating. Corpus bursae oval, membranous, without signum. Populations of southern France (Ardèche: subsp. cleui) often with corpus bursae small, globular and length of ductus bursae approaching that of corpus bursae. One examined interesting female from central Italy (gen.fig. 171b) in genitalia similar to those of T. lhommaria requiring further study (see below under Distribution). Examined females from Morocco (conspecific?) with shorter sclerotization of ductus bursae, approx. 1/3 length of corpus bursae. Leraut (2009) figures similar female genitalia from Morocco (Fig 101b) as “T. lhommaria ssp.”. 490

Distribution and abundance: West Mediterranean. Nominotypical subspecies in the French High Alps. Subsp. cleui from north-westernmost Italy (western Liguria: Flamigni et al. 2016) to southern France, subsp. melaleucaria in the Iberian peninsula (Spain and Portugal). For a potential occurrence in central Italy (Abruzzo, San Donato) see remarks under Female genitalia, more material is required to confirm the occurrence in central Italy and to verify the taxonomic status of a potential population in that region. – In Europe rather common, often occurring in numbers. – Outside Europe locally distributed in Morocco (DNA barcoded), reputedly also in Tunisia (Leraut 2009), but conspecifity still requires confirmation, genitalia are slightly different and no DNA barcodes are available for Tunisia, so far.

Phenology: Uni- or bivoltine. Nominotypical subspecies univoltine, from late June to late August, similar to populations at higher elevation in subsp. cleui and subsp. m ­ elaleucaria. At lower elevation under good conditions regularly bivoltine, in southern France from mid-June to mid-September, in the lowlands of southern Iberian peninsula bivoltine, from mid-April to late June; late July to early October. Larval development in southern France observed from October, overwintering, to June (Cleu 1933). Adults active at night, readily attracted to light. Biology: Larva found on and reared on lichens from Juniperus oxycedrus, J. phoenicea, J. communis, J. thurifera, Buxus, Quercus pubescens, Q. pedunculata, and Crataegus (Cleu 1933; Redondo & Gaston 1999; Leraut 2009; Flamigni et al. 2016). Rearing was successful under humid conditions, dry lichens were not accepted (Wehrli 1953). Immature stages described and figured by Gómez de Aizpúrua (1989) under the name of “T. sepiaria” (mis­ identified; cited by Redondo et al. 2001) after rearing from larva, found on lichens on Juniperus oxycedrus. Pupation described as occurring in a loose silken cocoon on the ground, possibly under artificial conditions (Cleu 1933). 491

Text-figs 204–206. Diagnostic characters (indicated) for wing-pattern-based identification of Tephronia species on the Iberian peninsula and in southern France: Text-fig. 204. T. sepiaria (Hufnagel, 1767) (France, Vaucluse, Puyvert, 10.ix.1988). Text-fig. 205. T. lhommaria (Cleu, 1928) (Portugal, Raposeira, 15.ii.2006). Text-fig. 206. T. espaniola (Schawerda, 1931) (Spain, Teruel, Moscardon, 28.vii.2014).

492

Habitat: Xerothermophilous, nominotypical subspecies xeromontane. In a large variety of habitat types from sand and marsh localities at sea-level to semi-open sites including Mediterranean macchia, scattered juniper or oak forest or scrubland, forest fringes, often on calcareous soils. Subsp. cleui usually from sea-level up to 1000 m, nominotypical subspecies up to 1500 m, subsp. melaleucaria up to 2150 m. Similar species (see Text-figs 204–206): Sympatric T. sepiaria (see no. 168), allopatric T. c. sicula (170). T. theophilaria (allopatric but with potential small overlap of distribution in Liguria, cf. Flamigni et al. 2016) with similar wing pattern, but forewings darker and more uniform brown coloured, pattern less contrasted. Genetic data: BIN: BOLD:AAE4162 (n=13 from north-westernmost Italy, France, Spain, Portugal, Morocco). Genetically heterogeneous, maximum variation 1.1%. BIN-sharing but not exactly barcode-sharing with T. cyrnea (0.5%), T. psyloritaria (0.6%), and T. lepraria (0.6%). In previous European literature, this species was misidentified as “T. codetaria”, for the DNA sequence of the holotype see under T. codetaria sicula. Remarks: Subsp. cleui was described from the French department Ardèche as did also ­Leraut (2009) in his description of diniensis. It is debatable if the populations from French High Alps, southern France and Iberian peninsula are really sharply separated from each other, and thus if subsp. cleui and subsp. melaleucaria can be validated at subspecies rank (cf. Flamigni et al. 2016). The taxon melaleucaria Schwingenschuss, 1932 was erroneously listed as synonym of T. espaniola (and misspelled to meloleucaria) by Scoble (1999). The figures in the original description clearly show forms of T. lhommaria.

172. Tephronia cyrnea (Schawerda, 1932) Mannia codetaria var. cyrnea Schawerda, 1932: Z. öst. Ent.Ver. 17 (2/3): 13, pl. 3, figs 9, 10 (France, Corsica: Evisa). Syntypes: 4♂ (not traced in NHMW and SMNK). Treated as synonym/subspecies of codetaria by Leraut (1980; 1997) and Scoble (1999), re-established at species rank by Hausmann et al. (2004; 2011a). ‡ Tephronia codetaria: sensu auct. (e.g. Raineri & Zangheri 1995), nec Oberthür, 1881 (mis­ identification). Tephronia tonnara Tautel, 2015: Oreina 30: 21, figs 2, 6c, 7c, 8 (France, Corsica: Bonifacio, Pont de Ventilegne). Holotypus ♂ (MNHN), two paratypes examined and DNA barcoded. Herewith downgraded from species rank to synonymy (syn.n.), see Remarks.

External characters and abdomen: Wingspan ♂♀ 18–22 mm. Ground colour whitish, often with grey suffusion, basal and/or terminal area usually darker. Pattern variable, reminiscent of that of T. lhommaria. Postmedial line of forewing strongly concave at centre. Postmedial line of hindwing usually well developed. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. Length of ♂ antennal branches 0.6–0.65 mm, 6–7 times width of flagellum. ♂ and ♀ with two pairs of hindtibial spurs.

Variation: Ground colour varies from very pale to darker. Basal and terminal area sometimes not darker than medial area. Male genitalia: Uncus rounded. Saccus large, shape variable in length and width. Central process of valva developed as a distinct lobe. Aedeagus large (length 1.65–1.75 mm). Cornutus long (0.6–0.7 mm), needle-shaped, straight or slightly curved, with narrow basal stalk shorter than the distal part. On Sardinia saccus narrower, somewhat shorter, digitate.

493

Female genitalia: Antrum small, with v- or u-shaped sclerotization. Ductus bursae narrow, short (0.6–0.9 mm), membranous, continuously dilating towards corpus bursae. Corpus bursae large (2.0–2.5 mm), membranous, pyriform or oval, at junction with ductus bursae with strongly wrinkled cuticle, usually dilating to corrugate appendix bursae. Signum absent. No constant differences observed between populations from Sardinia and Corsica. Distribution and abundance: Tyrrhenian. Endemic to Corsica and Sardinia. Mentioned for Sardinia and Sicily under the name ‘codetaria‘ by Raineri & Zangheri (1995), in the case of Sicily probably referring to T. c. sicula. – Local, with wider distribution only in central Sardinia, usually found in limited numbers.

Phenology: Usually univoltine, from late June to mid-August, at sea-level sometimes bivoltine, from late May to late June and from late August to mid-September. Type specimen of cyrnea from late July/early August. Probably overwintering as larva like the congeners. Adults active at night, readily attracted to light. Biology: Larva reported as feeding on lichens of Juniperus (Leraut 2009), possibly based on assumption. Habitat: Xerothermophilous. Mediterranean macchia and scattered Mediterranean forest and forest fringe at lower altitudes, associated with Quercus ilex, Q. suber, Pistacia lentiscus, Phillyrea, Juniperus, and Arbutus unedo (Flamigni et al. 2016). Also in old costal sand dunes with Juniperus and Pinus (P. Skou own data). Most data from sea-level up to 500 m. Type locality of cyrnea at 850 m, examined and verified specimens from Corsica up to 1100 m (Ciamanacce, ZSM, coll. P. Skou), in Sardinia up to 1000 m (Fonni, ZSM). Records from higher altitudes in Corsica (Tautel 2015) may be misidentified, see remarks.

494

Text-figs 207–209. Diagnostic characters (indicated) for wing-pattern-based identification of Tephronia species on Sardinia and Corsica: Text-fig. 207. T. nuragica Fiumi et al., 2013 (France, Corsica, Camping Lozzi, 5.–8.viii.2012). Text-fig. 208. T. cyrnea (Schawerda, 1932) (Italy, Sardinia, Stagno de Platamona, 23.v.2004). Text-fig. 209. T. theophilaria Hausmann sp. n. (Italy, Abruzzo, Arischia near Pizzola/Aquila, 4.viii.1980).

495

Similar species (see Text-figs 207–209): Sympatric T. nuragica (see no. 169). In male genitalia, shape of cornutus similar to that of sympatric (even in the same collecting nights!) T.  lepra­ria, but slightly curved and with shorter basal stalk in T. cyrnea whilst straight in T. lepraria, female genitalia with huge differences between both species. Allopatric T. lhommaria not clearly distinguishable in wing pattern. Genetic data: BIN: BOLD:AAE4162 (n=6 from Corsica and Sardinia). BIN-sharing but with defined, constant differences in the COI barcode from T. lhommaria (0.5%), and T. psyloritaria (0.8%). Populations from Corsica and Sardinia diverging by 1.2% (one Corsican paratype of “tonnara”, however, clustering with the Sardinian haplotype), both without genetic variation within the cluster. The Corsican cluster genetically is identical with the COI barcode of E. lepraria (attributed to ‘oppositaria’ in Fiumi et al. 2013; Flamigni et al. 2016; based on an earlier misidentification by the author A. Hausmann), suggesting introgression or regular gene flow in the recent evolutionary history of both species. Remarks: Until recently, subordinated as subspecies under T. codetaria by many authors, e.g. Leraut (1980; 1997) and Scoble (1999), raised to species rank by Hausmann et al. (2004, ‘Fauna Europaea’) and Leraut (2009). Further information in Fiumi et al. (2013). The taxon tonnara Tautel, 2015 was described from Corsica, based on a holotype male from Corsica (Bonifacio, Pont de Ventilegne), which in the description is stated to have one single pair of hindtibial spurs whilst the holotype figure (Tautel 2015: Fig. 2) clearly shows two pairs. DNA barcoding of two paratypes resulted in belonging to two different species, T. theophilaria (new for the fauna of Corsica) and T. cyrnea. The wing coloration and pattern of the holotype clearly suggests synonymy with T. cyrnea, though the figures of male genitalia of the holotype presented in Tautel (2015: Fig. 6c) show some unusual characters (e.g. smaller saccus, but neither matching those of the sympatric T. theophilaria). The genitalia of the female paratype (Fig. 7d) perfectly match those of T. cyrnea, in accordance with the cyrnea-like wing coloration and pattern. Unfortunately, the figures (showing two of the four syntypes) in the original publication of T. cyrnea do not allow an unambiguous attribution to one of the two species. Since we were not able to trace any of the four syntype specimens of T. cyrnea (failed in NHMW, SMNK, ZSM), it is not excluded that they may (all or partly) refer to the new taxon from Italy, Istria and Corsica (T. theophilaria sp. n.). In that case, cyrnea would have priority over theo­philaria, and tonnara should be validated for what is named as ‘cyrnea’ here (maintaining the taxonomic practice of the last 85 years). It is not recommendable to change the taxonomic understanding of the identity of cyrnea as it was conceived in all publications and collections, just based on an assumption that ‘the type photographs in the original description may also be similar to the wing pattern in T. theophilaria’. The question cannot be solved definitely without examination of the types (DNA, dissections), however, a neotype designation cannot currently be made because of the possible existence of (some of) the type specimens. Here, we prefer to maintain the conservative view and practice in classification and nomenclature. If the syntype series turns out to be mixed, the lectotype should be chosen in favour of stabilizing the traditional taxonomic concept as applied in this book.

496

173. Tephronia lepraria Rebel, 1909, comb. n. Tephronia sepiaria var. lepraria Rebel, 1909: Mitt. Ent. Ver. Polyxena 4 (1): 2 (Hungary: Buda­ pest, ‚Ofen‘). Syntypes ♂. Validated at species rank (Müller 1996; Scoble 1999, Leraut 2009), in all citations combined with Eumannia. Formally transferred to genus Tephronia by Flamigni et al. (2016), but based on misidentification (see under T. theophilaria). Herewith definitely transferred to genus Tephronia based on the integrative analysis including molecular and morphological characters (e.g. venation, female genitalia).

External characters and abdomen: Large, wingspan ♂♀ 20–24 mm, exceptionally only 18 mm. Forewing elongate and narrow. Ground colour of forewing pale grey with dark grey suffusion, darker than hindwing. Pattern contrasting. Postmedial line of forewing strongly concave and dentate. Hindwing pale with slight brown suffusion, postmedial line distinct, often dentate. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate with long and narrow antennal branches (0.4–0.6 mm = 5–7 times width of flagellum), in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs. Venation as described for Tephronia.

Variation: Variation mainly concerns suffusion over the ground colour and distinctness of pattern. Male genitalia: Uncus (sub-)truncate and shallowly concave posteriorly. Saccus strongly projecting, narrow. Valva narrow, bent subapically. Spinulose process in rather basal position, comparatively small. Aedeagus very large, straight or slightly curved (length 1.9–2.0 mm). Cornutus very long and narrow, length 0.6–0.8 mm, basal extension sometimes longer than the rest of the needle- or sword-shaped distal part. Female genitalia: Apophyses very long, apophyses anteriores 0.5–0.55 mm, apophyses posteriores 1.2–1.3 mm. Antrum funnel-shaped. Ductus bursae narrow, membranous, very long (3–4 mm), towards corpus bursae corrugated. Corpus bursae membranous, sometimes globular but not always sharply separated from ductus bursae. Signum absent. Distribution and abundance: (South-east) European-West Asiatic. Described from H ­ ungary, found also in south-eastern Austria (vouchers in ZSM; cf. Forster & Wohlfahrt 1981 though not confirmed by Huemer & Tarmann 1993 nor by Huemer et al. 2009) and in the Balkan peninsula from Croatia to Greece, Bulgaria and European Turkey. Earlier records from Romania (Popescu-Gorj 1980) doubted by Székely (1999). Recorded for ‘eastern Europe’ (Müller 1996 ‘EE’), the only confirmed records being from Crimea (Efetov & Budashkin 1990; Viidalepp 1996; Kostjuk 2004; dissected). – Usually local and rare, last verified record in Hungary from 1980 (L. Ronkay pers. comm.). – Outside Europe in Turkey (material in ZSM). Phenology: Univoltine, flying in late summer and early autumn. In Europe from early August to late September, rarely from late July. In Turkey and Crimea at higher elevation regularly emerging earlier, from mid-July. Adults active at night, readily attracted to light. Biology: Immature stages unknown. Larval food-plants probably lichens as in the congeners. Habitat: Xerothermophilous. Open or semi-open lowland and coastal habitats, forest fringes, Mediterranean macchia, riverside habitats. In the north usually restricted to lowlands from sea-level up to 1000 m, in southern Balkan countries and Turkey more common on hills and mountains (compare Eumannia oppositaria), up to 1300 m.

497

Similar species (see Text-figs 210–212): Sympatric T. sepiaria with two male hindtibial spurs (further differential features see no. 168). In Balkan countries sometimes syntopic with ­Eumannia oppositaria. The latter differs by an earlier flight period (in Europe usually late May to mid-July), a smaller wingspan (usually 16–19 mm), forewing broader and rounded, ground colour of both wings usually paler, glossy brownish, pattern less contrasting, ante­ medial line of forewing more zigzagging, male antennal branches very short (3–4 times width of flagellum). Allopatric T. theophilaria (no. 175) smaller, forewing colour fawn, postmedial line of forewing less concave, pattern less contrasting, male antennal branches longer and narrower (0.6–0.7 mm = 6–8 times width of flagellum), flying earlier in the year, in the south bivoltine. In male genitalia the uncus posteriorly rounded. Female genitalia of T. theo­ philaria completely different, bursa copulatix much longer. Allopatric T. cyrnea in habitus very similar, but on average larger, forewing somewhat less irrorated by dark scales. In male genitalia aedeagus longer, basal stalk of cornutus comparatively short. Female genitalia of T. cyrnea with much shorter ductus bursae, cuticle between ductus and corpus bursae more strongly wrinkled. Genetic data: BIN: BOLD:AAE4162 (n=12 from Croatia, Serbia, Bulgaria, Greece, Turkey). Genetically homogeneous. BIN-sharing and exactly DNA barcode-sharing with one of the two clusters of T. cyrnea, suggesting introgression or regular gene flow in the recent evolutionary history of both species. BIN-sharing but not exactly barcode-sharing with T. psyloritaria (0.6%), and T. lhommaria (0.6%).

498

Text-figs 210–212. Diagnostic characters (indicated) for wing-pattern-based identification of Tephronia and Eumannia species in the Balkan peninsula, eastern Austria and Hungary: Textfig. 210. Tephronia sepiaria (Hufnagel, 1767) (Italy, Lago di Garda, 27.vii.1990). Text-fig. 211. T. lepraria Rebel, 1909 (Austria (inf.), Marchfeld, Oberweiden, 25.vii.1936). Text-fig. 212. Eumannia oppositaria (Mann, 1864) (Macedonia, Sweti Wratschi, 10.vii.1933).

499

174. Tephronia psyloritaria (Reisser, 1958), comb. n. Mannia psyloritaria Reisser, 1958: Z. wien. ent. Ges. 43 (7): 123 (Greece, Crete: Silva Rouva). Holotype ♂ (SMNK?), photos of holotype and paratypes examined. In earlier publications combined with Eumannia, herewith transferred to genus Tephronia (comb. n.) based on clear relationship with the species group around T. lhommaria in COI barcodes, venation and other differential features.

External characters and abdomen: Wingspan ♂♀ 19–21 mm. Coloration uniform in ♂, basal and terminal area much darker and medial area whitish in ♀. Medial area sometimes very broad. Ante- and postmedial lines of forewing comparatively straight. Hindwing paler, here postmedial line straight, bent near apex. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs.

Male genitalia: Uncus truncate, very slightly concave posteriorly. Saccus very large. Central process of valva developed as broad lobe. Aedeagus broad and large (length 1.6 mm). Cornutus large (length 0.65 mm).

Female genitalia: Antrum large, funnel-shaped. Ductus bursae short (0.6–0.7 mm), membranous, narrow. Corpus bursae large (length 1.8–2.3 mm), oval, anteriorly membranous, dilated laterally. Posteriorly more strongly sclerotized, wrinkled (as in T. cyrnea and T. lepraria) with double spinulose ring (signum). Distribution and abundance: Endemic to the mountains of central Crete. – Rare and local.

Phenology: Probably univoltine at higher elevation, (partially?) bivoltine at lower elevations. Most data from late June to mid-August, two specimens (at low elevation) from early to mid-October, and one (e.l.!) from late May. Adults active at night, readily attracted to light. 500

Biology: Larval stages unknown. Larvae probably feeding on lichens like the congeners. Habitat: Xeromontane. Warm and dry slopes, canyons. Vertical distribution from 300 m up to 1600 m a.s.l. Similar species: On the island of Crete without sympatric, congeneric species, thus unmistakeable. Genetic data: BIN: BOLD:AAE4162 (n=2 from Greece, Crete island). Genetically heterogeneous, both specimens diverging by 0.5% from each other. BIN-sharing but with defined, constant differences in the COI barcode from T. lhommaria (0.6%), T. lepraria (0.6%), and T. cyrnea (0.8%). Remarks: Until recently, this species was combined with genus Eumannia (Scoble 1999; Leraut 2009). Genetic similarity clearly supports a position in Tephronia and even suggests a potential downgrading to subsp. of T. lepraria or T. lhommaria but the huge differences in male and female genitalia do not allow any change of taxonomic status.

175. Tephronia theophilaria Hausmann, sp. n. ‡? Tephronia cremiaria: sensu Verity (1906) nec Freyer, 1838. Misidentification. Based on 1♂ (late July) from Vallombrosa (Tuscany), presumably belonging here. ‡ Eumannia oppositaria: sensu Viidalepp (1988a) nec Mann, 1864. Misidentification. Viidalepp (1988a: pl. 37 fig 78) illustrates the male genitalia of an Italian specimen. ‡ Tephronia oranaria: sensu Parenzan (1994) nec Staudinger, 1892. Misidentification. ‡ Tephronia tonnara: sensu Tautel (2015; partim) nec Tautel, 2015 (holotype). Misidentification of at least one paratype (see under ‘genetic data’). ‡ Tephronia lepraria: sensu Flamigni et al. (2016) nec Staudinger, 1892. Misidentification, based on an earlier, erroneous identification by the present author (A. Hausmann). Examined material. Holotype: ♂, Italia centr. [Abruzzo], Mte. Sirente, 10.VII.1914, leg. Dannehl, coll. ZSM, DNA barcode specimenID BC ZSM Lep 103417_NGS. Paratypes: Coll. ZSM (32♂2♀): 1♂, Italy, Toscana, Grosseto, l’Argentario, 2 km N Cala Piccola, 100 m, 5.–11.VII.1997, leg. P. Schwarzbauer; 5♂, Toscana, Anchiano, Vinci (FI), 100 m, 20.VII-21.IX.1995, leg. S. Scalercio (gen.prp. SS212, 213); 1♂, Reppublica di San Marino, Monte Cucco (sud), bosco, 350 m, Castello San Marino, 4.VII.2005, leg. L. Rezbanyai-Reser & E. Schäffer; 1♂, Italia centr., Monti Albani, leg. Dannehl, [19]14; 1♂ Italia, Apennin, Lago di Ortiglieto, 300 m, 9.–23.VIII.1977, leg. G. Behounek; 15♂, Italien, Abruzzen, oberh. Arischia, 1000 m, bei Pizzola/Aquila, 4.VIII.1970, leg. Gross, coll. EMEM in ZSM (gen.prp. ZSM G 13279); 1♂, Italia mer. Calabria centr., Donnici inf. (Cosenza), 600 m, 5.IX.1997, leg. A. Hausmann; 1♂, Calabria, loc. Donnici (CS), Fosso Cucolo, 550 m, 15.VII.1995, leg. S. Scalercio; 2♂, id., 20.VII.1996 (DNA barcode BC ZSM Lep 14333); 1♀, id., 13.VII.1995 (gen.prp. ZSM G 13297); 1♂, Lago Angitola, Scuotrapiti (VV), 44 m, 23.V.2001, leg. S. Scalercio; 1♂, id., 21.IV.1999, leg. S. Scalercio, M. Infusino & Bonacci; 1♂, Italia mer., Basilicata mer., Maratea, statua del Redentore, 650 m, 19.VIII.2004, leg. A. Hausmann (DNA barcode BC ZSM Lep 14301); 1♀, Italia mer., Basilicata mer., upper Noce valley, by Lago­ negro, 700 m, 2.IX.1998, leg. A. Hausmann (DNA barcode BC ZSM Lep 10677 – gen.prp. ZSM G 13296); 1♂, Basilicata, Oasi San Giuliano (MT), loc. Ponte Cagnolino, 250 m, 26.– 31.VIII.1992, leg. P. Parenzan (DNA barcode BC ZSM Lep 14309 – gen.prp. ZSM G 6280).

Coll. P. Skou (19♂1♀): 18♂1♀, [Italia, Tuscany], Albiano, Arezzo, 18.VII.[19]79, leg. A. Moberg; 1♂, [Italy], Umbria, Orvieto, 25.VII.[19]58, leg. C. Prola.

Coll. Centro di Ricerca Foreste e Legno (CREA-FL), all leg. S. Scalercio & M. Infusino (15♂2♀): 1♂: Italia Calabria, SR_A2, Il Palmento, S.S. Bruno (VV), 840 m, 3.VIII.2016, 38.5625°, 16.3140°; 1♂: Italia Calabria, SR_D2, Rosarella, S.S. Bruno (VV), 917 m,

501

3.VIII.2016, 38.5560°, 16.3350°; 1♂: Italia Calabria, SR_A1, Santa Maria, S.S. Bruno (VV), 860 m, 3.VIII.2016, 38.5550°, 16.3109°; 2♂: Italia Calabria, SR_D1, Rosarella, S.S. Bruno (VV), 884 m, 3.VIII.2016, 38.5552°, 16.3329°; 1♂: Italia Calabria, CC_A2, Fiego di San Fili (CS), 740 m, 19.VIII.2015, 39.3302°, 16.1293°; 1♂1♀: Italia Calabria, CC_A2, Fiego di San Fili (CS), 740 m, 19.VIII.2015, 39.3372°, 16.1290°; 1♂: Italia Calabria, PO_A2, Serrapaolo, Saracena (CS), 1010 m, 24.VIII.2015, 39.8225°, 16.0883°; 2♂1♀: Italia Calabria, PO_A2, Serrapaolo, Saracena (CS), 1010 m, 20.VII.2015, 39.8225°, 16.0883°; 5♂: Italia Calabria, PO_A1, Serrapaolo, Saracena (CS), 990 m, 20.VII.2015, 39.8225°, 16.0911°. Coll. G. Govi: 1♂, Italy, Romagna, sopra Corniolo (FC), San Paolo in Alpe, 1000 m, 1.VIII.2014, leg. G.Govi. Coll. G. Fiumi (14♂1♀): 2♂, Romagna Castrocaro loc. Cozzi (FC), 100 m, 8.VIII.1980; 1♂, id., 17.VII.1991; 1♂, id., 2.VII.1980; 1♂, Romagna sopra Meldola (FC), 150 m, 13.VII.1979; 1♂, Romagna Predappio loc. Tontola (FC), 200 m, 19.VII.1988; 1♀, Romagna Premilcuore loc. Fiumicello (FC), 700 m, 15.VIII.1987; 1♂, id., 26.VII.1995; 1♂, Romagna Alfero loc. Riofreddo (FC), 800 m, 15.VII.1982; 1♂, Romagna Villagrande loc. Calvillano (RN), 1000 m, 22.VIII.2012; 4♂, Romagna (crinale appenninico) Passo del Muraglione (FI), 850 m, 3.VIII.2011; 1♂, Romagna Campigna loc. San Paolo in Alpe (FC), 1000 m; 2.VIII.2014. Coll. C. Flamigni (9♂): 2♂, Emilia-Romagna: Bologna, Monte Adone località Raieda, 500 m, 29.VIII.1987, leg. C. Flamigni; 2♂, Emilia-Romagna: Bologna, Monzuno località la Bettola, 440 m, 27.VII.1988, leg. C. Flamigni; 3♂, Emilia-Romagna: Bologna, torrente Idice presso Savazza, 250 m, 26.VII.1993, leg. C. Flamigni; 1♂, Toscana: Siena, Sarteano, 600 m, 14–16.VIII.2002, leg. L. Dapporto & C. Pansolli, 1♂, Puglia: Bari, Pulo di Altamura, 385 m, 8.VIII.1999, leg. L. Dapporto. [the type series is restricted to material from central and southern Italy to warrant geographical homogeneity]. Further material examined: 43♂1♀ from northern Italy and Istria (colls T. Grünewald, ZSM); 1♂ from Liguria (coll. G. Fiumi), 5♂ from Sicily (colls G. Fiumi, C. Flamigni), 3♂ from Corsica (colls P. Skou, ZSM). The type series is restricted to material from central and southern Italy to guarantee geographical homogeneity. Etymology: The species is named in honour of Dr. Theo Grünewald (Landshut) who donated his huge collection to the ZSM, who supported our studies in many ways and who had correctly separated his samples of Tephronia and Eumannia to the different morphospecies, including T. theophilaria sp.n.

Description of external characters and abdomen: Wingspan ♂♀ 17–21 mm. Forewing homo­ geneously fawn brown, much darker than hindwing. Antemedial line of forewing comparatively straight, at inner termen projecting towards termen, postmedial line conspicuous, concave, usually slightly dentate. Hindwing pale with slight brown suffusion, post­medial line distinct. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. Length of ♂ antennal branches 0.6–0.7 mm = 6–8 times width of flagellum. ♂ and ♀ with two pairs of hindtibial spurs. Variation: One of the least variable Tephronia species, only slight variation of ground colour and distinctness of pattern. Male genitalia: Uncus posteriorly rounded. Saccus strongly projecting, narrow. Valva narrow, bent sub-apically. Spinulose process in rather basal position, comparatively small. Aede­agus large, straight or slightly curved (length 1.55–1.65 mm). Cornutus very long and narrow, length 0.65–0.8 mm, basal extension sometimes longer than rest of needle- or sword-shaped distal part.

502

Female genitalia: Antrum funnel-shaped. Ductus bursae narrow, membranous, very long (3–4 mm). Corpus bursae membranous, sometimes globular but not always sharply separated from ductus bursae. Signum absent. Distribution and abundance: (South-) European. Distributed in Slovenia, Croatia (Istria), and in north-easternmost Italy. Widely distributed on the Apennine peninsula from southern Piemonte (Baldizzone et al. 2013), Liguria, Emilia-Romagna and San Marino (Flamigni et al. 2008) across central and southern Italy to northern Sicily (Parenzan 1994; Grillo & ­Parenzan 1995; Flamigni et al. 2001; 2002; Zilli et al. 2001; Sciarretta & Zahm 2002), records including many citations under the names ‘cremiaria’ (Tuscany: Verity 1906), ‘ora­naria’ (e.g. sensu Parenzan 1994) and ‘lepraria’ (in recent publications). Isolated records from Alto Adige (‘Bolzano’: Flamigni et al. 2016; ‘Schnalstal’ (side valley of Val Venosta): own data from ZSM) and Lake Garda (ZSM). Present also on Corsica (see remarks to T. cyrnea). – Not rare and sometimes common at the sites of occurrence. – Not distributed outside Europe.

Phenology: Univoltine, flying in late summer, from early July to early September, most data from late July to late August. In Tuscany one record as late as late September, in Sicily as early as late June. Two very early records from southern Calabria at sea-level from late April and late May indicating regional, occasional and/or partial bivoltinism. Adults active at night, males readily attracted to light but ♀-rate at light very low (5%).

Biology: Larva on lichens on wood or rocks (Deutsch 2006), probably based on assumption. Habitat: Meso- to xerothermophilous. Open or semi-open lowland and coastal habitats, forest fringes, Mediterranean macchia, scattered oak forests, riverside habitats and valleys with Alnus, Populus and Salix (cf. Flamigni et al. 2016). Most data from 500 m up to 1400 m, but sometimes (e.g. Istria, Romagna) down to sea-level and lower coastal hills. 503

Similar species (see Text-figs 201–203 and 207–209): Sympatric T. sepiaria with two male hindtibial spurs (further differential features see no. 168), T. c. sicula (170) usually with more strongly concave postmedial line and darker basal area of forewing. The fawn ground colour of the forewing of T. theophilaria and the much paler hindwing with distinct postmedial line usually allows discrimination from sympatric species. Genetic data: BIN: BOLD:AAF8352 (n=6 from Italy, France: Corsica, including one paratype of “T. tonnara”). Genetically homogeneous (maximum variation 0.15%). Nearest species: An undescribed species from the Levant (4.1%), T. sepiaria (4.6%), T. nuragica (5.3%). COI data suggesting a quite isolated evolutionary history within the genus. Remarks: Female genitalia are similar to those of the North African T. praerecta Wehrli, 1933 but genetically it is far from the latter.

176. Tephronia espaniola (Schawerda, 1931) Mannia oranaria var. espaniola Schawerda, 1931: Mitt. münch. ent. Ges. 20 (3/4): 137, pl. 7, fig. 10 (Spain, Aragon: Sierra Alta near Albarracin). Syntypes 2♂ (NHMW). Unavailable names: (incorrect subsequent spelling): espagnolia: Tautel (2015).

External characters and abdomen: Large, wingspan ♂♀ 20.5–24 mm. Forewings broad, apex rounded. Ground colour of forewing grey, usually with strong brown suffusion, rendering pattern diffuse. Hindwing slightly paler than forewing. Postmedial line of forewing rather straight, slightly concave, usually broken into dots. Medial line of forewing sometimes distinct, zigzagging. Postmedial line of hindwing indistinct or absent. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate, in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs.

Variation: Ground colour variable, pale grey to dark grey. Several examined specimens (♂♀) with pale, creamy ground colour and with very sharp pattern. Male genitalia: Uncus broad at tip, sometimes slightly concave posteriorly. Saccus strongly projecting, narrow. Central process of valva lobed. Aedeagus very large, straight, length 1.6–1.7 mm, without cornutus.

Female genitalia: Antrum with two long, lateral sclerites. Sterigma oval, sclerotized, with two minute posterior projections. Ductus bursae and corpus bursae narrow, membranous, very long (4.5 mm), posteriorly with longitudinal furrows. Signum absent. Distribution and abundance: West Mediterranean. Endemic to a small area from ­central to north-eastern Spain, with isolated occurrences in France/Hérault (Leraut 2009), south-­ eastern Spain (Guerrero et al. 2010; Aistleitner et al. 2017). Recently recorded for north-­ eastern and central Portugal (Corley et al. 2018). – Local, usually in limited numbers, but now and then up to 20 specimens found in one night. Phenology: Univoltine. From early July to mid-August, sometimes emerging two to three weeks earlier. Adults active at night, readily attracted to light. Biology: Larva reported as feeding on lichens as the congeners (cf. Leraut 2009, probably on assumption). Immature stages are described and figured in Gómez de Aizpúrua (1989), after rearing from egg on lichens. Habitat: Xeromontane. Apparently related to habitats with scattered Quercus and fringe of denser oak forest. Often in slightly cool places within hot landscapes, such as north-facing 504

slopes or canyons with small rivers/streams, covered by smaller oak trees and bushes. From 700 m up to 1400 m a.s.l., absent from coastal regions. Similar species (see Text-figs 204–206): Sympatric T. lhommaria with less dark suffusion, postmedial line of forewing stronger concave, postmedial line of hindwing more strongly marked. T. fatimaria Bang-Haas, 1906 from Morocco and Algeria (allopatric) with narrower forewings, transverse lines more strongly contrasted on all wings, in male genitalia aedeagus narrower and with cornutus, female genitalia (corpus bursae) with a double spinose ring. Genetic data: BIN: BOLD:ADE1551 (n=4 from Spain). Genetically homogeneous. Nearest species: T. lhommaria, T. lepraria and T. cyrnea (all 5.2%). COI data suggesting a quite isolated evolutionary history within the genus. Remarks: Female genitalia suggest certain relationships between T. espaniola and T. theo­ philaria, whilst DNA barcodes indicate rather isolated positions of both taxa. The taxon proutiana Turati, 1936, described from Libya, is erroneously listed as a synonym of T.  espan­iola in Leraut (2009). The taxon melaleucaria Schwingenschuss, 1932 was erroneously listed as a synonym of T. espaniola (and misspelled to meloleucaria) by Scoble (1999), see remarks under T. lhommaria.

Eumannia Fletcher, 1979 ‡ Mannia Prout, 1915, in Seitz, Gross-Schmett. Erde 4: 382. Type species: Tephronia oppositaria Mann, 1864, by original designation. Junior homonym of Mannia Davidson, 1874 [Brachio­ poda]. Eumannia Fletcher, 1979, The Generic Names of the Moths of the World Vol. 3: 82. Type species: Tephronia oppositaria Mann, 1864, by original designation. Replacement name for Mannia Prout, 1915. Validity as a separate genus from Tephronia confirmed here, as stated in all earlier publications.

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Genus with six species in the western Palaearctic region, two in Europe, three in the ­Middle East (Eumannia arenbergeri Hausmann, 1995 from Cyprus, E. fumosata Hausmann, 1995 from the Levant and E. neoppositaria Beljaev & Vasilenko, 1998 from Turkmenistan) and one in North Africa (Libya): E. syriaca (Turati & Krüger, 1936), for nomenclature see Hausmann (1995a). The taxon oxygonaria Püngeler, 1900 was transferred to genus Haber­ mania by Viidalepp (1988a), lepraria Rebel, 1909 was (re-)transferred to genus Tephronia by F ­ lamigni et al. (2016), psyloritaria Reisser, 1958 is transferred here to genus Tephronia. Differential characters: ♂ hindtibia with 4 spurs. Proboscis short and weak. ♂ antennae with short branches (0.2–0.4 mm). Venation similar to that of Tephronia but on hindwing veins Sc and Rs+M1 fused, common stalk of Rs and M1 comparatively short. Vein M2 absent from hindwing in E. oppositaria, tubular but weak in E. oranaria. ♂ genitalia with two weak, membranous latero-posterior processes of uncus, aedeagus short, valva with spinulose process in apical third; ♀ genitalia with signum developed as bipartite, flat sclerite, usually with two spinules. Biology lichens.

and immature stages:

Larva lichenophagous, with well camouflaged coloration on

Genetic data: DNA barcodes (COI) support combination of the above mentioned species with genus Eumannia. DNA barcodes of Eumannia significantly diverging from those of the genera Tephronia and Phyllometra (cf. Text-fig. 200), independent of the chosen algorithm and on the tree construction settings (at maximum likelihood bootstraps >60). Remarks: Genus Eumannia validated as separate genus from Tephronia in Fletcher (1979), Müller (1996), Scoble (1999) and Leraut (2009).

177. Eumannia oppositaria (Mann, 1864) Tephronia oppositaria Mann, 1864: Wien. ent. Monatschr. 8 (6): 178, pl. 4, fig. 4 30: 21 (north-western Turkey: Bursa). Syntype(s) (NHMUK), examined. Tephronia obscuraria Staudinger, 1892: Dt. ent. Z. Iris 5; 178 (northern Iran: Astrabad). Syntypes (MNHU: 5♂♀ syntypes, examined). Currently validated at subspecific rank (Hausmann 1995b; Scoble 1999), see remarks. Unavailable names: (infrasubspecific): altmannii: Koutsaftikis (1973).

External characters and abdomen: Wingspan ♂♀ usually 16–19 mm, females on average larger than males, sometimes up to 21 mm. Forewing broad, rounded at apex. Ground colour pale grey-brown, on forewing not much darker than on hindwing. Hindwing colour slightly glossy. Postmedial line of forewing slightly concave, slightly dentate. Antemedial line zigzagging, sharply bent twice. Postmedial line of hindwing usually vague. Palpi very short. Proboscis vestigial. ♂ antennae bipectinate with short and broad branches (0.2–0.35 mm = 3–4 times width of flagellum), in ♀ filiform. ♂ and ♀ with two pairs of hindtibial spurs. Hindwing veins Sc and Rs+M1 fused. Variation: Sometimes with darker suffusion of ground colour (Peloponnese, north-western Turkey), reminiscent of Near Eastern E. fumosata Hausmann, 1995. Male genitalia: Uncus with tapering lateral processes, deeply (rounded) concave between. Saccus short, digitate. ‘Central’ process of valva long, appressed to membranous tip of ­valva, spinulose only at tip. Aedeagus very short and broad (length 0.95–1.05 mm). Cornutus long (0.5 mm), stout, bent, basal part approx. 1/4 length of distal, longitudinally fur506

rowed part. Genitalia of north-western Turkish populations well corresponding with those from European populations. In the (doubtfully conspecific) populations from northern Iran the aedeagus is somewhat longer. In the sister species E. fumosata Hausmann, 1995 from Israel and Jordan the valva process is less appressed to the tip of valva, length of aedeagus 0.8–0.9 mm, cornutus fine, needle-shaped, very short (0.2–0.25 mm). Female genitalia: Antrum sclerotized, laterally rounded. Sterigma posteriorly sclerotized as a waved band. Ductus bursae short (0.5–0.6 mm), broad. Corpus bursae oval or globular, membranous, with a small bipartite and bicornute signum, both parts almost separate. In the populations from northern Iran female genitalia are similar, but the antrum is more swollen. Sister species E. fumosata Hausmann, 1995 from the Levant (Israel, Jordan) with edges of antrum straight, signum fused to a plate with paired spinules, but without the longitudinal crests as an extension of these spinules. Sterigma band narrower. Distribution and abundance: (South-east) European-West Asiatic. In Europe restricted to the southern Balkan peninsula (Macedonia, Bulgaria and Greece). Recorded for southern Serbia by Beshkov & Nahirnic (2017) but erroneously attributed to E. oppositaria (after misidentification by the author AH) and partly referring to T. lepraria and T. sepiaria. Reputedly in the Ukraine (Savchuk 2017), the photos however suggesting misidentification and confusion with T. sepiaria and also the other records (Efetov & Budashkin 1990; Kostjuk 2004) could not yet be confirmed by dissection or DNA barcodes. Records for the European part of Russia (Leraut 2009) doubtful, data awaiting confirmation. – In Europe sometimes abundant. – Outside Europe in Turkey (type locality: north-western Turkey). In the Caucasus (Pyatigorsk), Transcaucasus (e.g. Georgia: Didmanidze 2002) and northern Iran replaced by subsp. obscuraria Staudinger, 1892 (for taxonomic status see genetic data and remarks), in the Levant by sister species E. fumosata Hausmann, 1995.

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Phenology: On hills and mountains usually univoltine, usually from early June to late July, exceptionally up to ten days earlier or later, single females until mid-August (protandry). Under good conditions in the lowlands regularly bivoltine, from mid-May to mid-June and from late July to late September, sometimes until mid-October. Larvae found on Samos from September, pupation after hibernation late May (Fritsch et al. 2014). Adults active at night, females readily attracted to light, males apparently less, thus with high ♀-rate at light (70–80%). Biology: Larva feeding on lichens. Frequently found on beard lichens (Usnea) (Samos: Fritsch et al. 2014).

Habitat: Xerothermophilous. Dry and warm habitats with Mediterranean vegetation. From sea-level up to approx. 1400 m, more common in the lowlands, in Samos abundant above 700 m (Fritsch et al. 2014). Similar species (see Text-figs 210–212): Sympatric T. sepiaria (see no. 168), in males easily separated by number of hindtibial spurs and length of antennal branches. Sympatric T. lepraria (no. 173) larger, forewings elongate and narrower, forewing colour pale grey with dark grey suffusion, postmedial line of forewing more strongly concave, pattern more contrasting, hindwing much paler than forewing, male antennal branches longer and narrower (0.4–0.6 mm = 5–7 times width of flagellum), flying later in the year. Genetic data: BIN: BOLD:AAZ7045 (n=6 from Greece, Turkey, northern Iran), Euro­pean and Turkish populations diverging by 1.4% from Iranian subsp. obscuraria Staudinger, 1901. Nearest species: E. fumosata Hausmann, 1995 from the Levant (2.2%), and E. arenbergeri from Cyprus (6.0%). Remarks: The taxa syriaca (Turati & Krüger, 1936) (locus typicus Libya, Cyrenaica) and obscuraria Staudinger, 1901 (locus typicus northern Iran) are listed as synonyms in Leraut (2009), apparently without knowledge of Hausmann (1995b). Nevertheless, both have to be regarded as separate valid taxa (species or subspecies): E. obscuraria obscuraria shows remarkable COI divergence from both E. oppositaria and E. fumosata Hausmann, 1995, see also diagnosis in Hausmann (1995b). Recently some specimens of E. syriaca from Cyrenaica have been investigated and dissected by F. Weisert (pers. comm.), clearly showing them to be different from E. fumosata and E. oppositaria. E. neoppositaria Beljaev & Vasilenko, 1998 from Turkmenistan was described with (correct) differential analysis from E. oppositaria but without deeper analysis of E. fumosata from the Levant. The single mentioned character (‘dilation of the apex of the saccus’) is surely not a valid differential character, on the contrary, the figured male genitalia suggest potential downgrading of the Turkmenistan taxon to subspecies of E. fumosata. However, for a formal taxonomic change it would be good to base it also on data from DNA barcoding, which are not yet available for populations from Turkmenistan.

178. Eumannia oranaria (Staudinger, 1892), comb. n. Tephronia oranaria Staudinger, 1892: Dt. ent. Z. Iris 5; 179 (Algeria, Oran: Sebdou). Syntypes (MNHU: 3♂1♀ syntypes, photos examined, two further reputedly on loan; 1♂ ‘paratype’ in coll Püngeler/MNHU; 1 further ♂ from Sebdou examined in NHMUK, genitalia slide NHMUK Geom. 4870 examined). Herewith transferred to genus Eumannia based on a number of synapomorphies shared with E. oppositaria, e.g. the short male antennal branches, the fusion of hindwing veins Sc and Rs+M1 (stated even for the type specimens in the original description!), the distal position of the spinulose process of valva, the shape of the signum, the similarity in the COI barcode etc.

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Tephronia castiliaria Staudinger, 1892: Dt. ent. Z. Iris 5; 180, pl. 2, fig. 14 (not ‘11’ as erroneously stated in the caption and in the text of the original description) (Spain, Castilia: St. Ildefonso; Andalusia: Chiclana). Syntypes ♂♀ (MNHU: 4♀ syntypes examined, two further reputedly on loan). In Prout (1915) validated as separate species, placed in ‘Tephronia’ intended as the group with two male hindtibial spurs. Validated at subspecies rank under E. oranaria (Wehrli 1953; Scoble 1999; Leraut 2009). Tephronia cebennaria Chrétien, 1898: Bull. Soc. ent. Fr. 1898: 20 (France, Ardèche: La Voulte). Lectotype ♀ (ZSM/Herbulot, examined and barcoded: BC ZSM Lep 85886_NGS; 1♂1♀ paralectotypes – labelled as ‘paratypes’ – in MNHU, examined). Validated at species rank in Prout (1915), later downgraded to subspecies rank under E. oranaria (Scoble 1999; Flamigni et al. 2016). Mannia cebennaria var. stephaniaria Lhomme, 1930: Amateur Papill. 5: 92 (France, Massif Central: St. Étienne V.F.). Syntype(s). Though one time called ‘aberration’ in the text, introduced as ‘var.’ and classified as ‘form of St. Étienne’ thus with geographical background, not infrasubspecific, nomenclaturally available. Spelling and nomenclatural availability accepted by Wehrli (1953). Listed as synonym of “T. oranaria subsp. castiliaria” in Leraut (1980, 1997; misspelled to ‘stephenaria’), not listed in Scoble (1999). In Leraut (2009) correctly spelled but with the authorship of Cleu, 1933. Junior synonym. ‡ Mannia cebennaria var. stephanaria Lhomme, 1930b [Dec 31]: Catal. Lépid. France et Belgique 1 (2): 420: Not infrasubspecific, classified as ‘race’. Incorrect subsequent spelling (i.e. without explicit indication for an intended emendation). Misspelling subsequently followed by Cleu (1933) and others. ‡ Mannia cebennaria ab. luminosaria Cleu, 1933: Amateur Papill. 6: (14): 231 (locus typicus not stated, syntypes ♂♀, one ‘cotype’ ♂ ex coll. Cleu: France ‘Ardèche, Vallon, 13.VII.32 e.l. Jun. oxycedrus’, examined in ZFMK). Infrasubspecific, non available (thus not listed in Scoble 1999). Tephronia oranaria alpinaria Leraut, 2009: Moths of Europe, Vol. 2, Geometrid Moths: 202, pl. 54, figs 2, 3 (France, Hautes Alpes: Château Queiras). Holotype ♂ (MNHN). Here validated at subspecies rank. Tephronia oranaria hubertcleui Leraut, 2009: Moths of Europe, Vol. 2, Geometrid Moths: 202, pl. 54, figs 4, 5 (France, Ardèche: Lagorce). Holotype ♂ (MNHN). Herewith downgraded from subspecies rank (Leraut 2009) to synonymy of subsp. cebennaria because of the lack of constant and sharp delimitation from the latter taxon (see text below). Unavailable names: (incorrect subsequent spellings): orangaria: Wehrli (1953); oronaria: Wehrli (1953); onanaria: Nestorova (1998); stephenaria: Leraut (1980; 1997); stephanaria: Lhomme (1930b).

External characters and abdomen: Wingspan ♂♀ 17–20 mm, females on average larger than males. Nominotypical subspecies rather variable in ground colour, contrast and extent of wing pattern. Forewing much darker than hindwing. Postmedial line of forewing comparatively straight (slightly waved), dentate, whitish bordered distally in North African populations. Antemedial line strongly zigzagging. Palpi very short. Proboscis short or vestigial. ♂ antennae bipectinate, antennal branches comparatively short (0.35–0.4 mm), ♀ antennae filiform. ♂ and ♀ with two pairs of hindtibial spurs. Hindwing veins Sc and Rs+M1 fused.

Subsp. castiliaria from Iberian peninsula smaller than nominotypical subspecies, wingspan ♂♀ 14–18 mm. Females with broader forewings. Ground colour very variable, from whitish or greyish to brown. Postmedial line of forewing usually broken into rows of dots. Hindwing darker than in nominotypical subspecies. Forewing terminal area with dark fascia. Subsp. cebennaria occurring in three infrasubspecific forms (Flamigni et al. 2016), (a) the nominotypical form paler and better contrasted than in E. o. oranaria; (b) f. stephani­ aria (described from Massif Central) usually being very dark; (c) f. luminosaria with whitish

509

ground colour and reduced or absent dark pattern in terminal area (dominant in some localities of the southern calcareous edge of the French Massif Central). The last was renamed by Leraut (2009) as ‘subsp.’ hubertcleui, but we agree with Flamigni et al. (2016) that such subspecific status cannot be accepted because of the lack of constant differential features from subsp. cebennaria described from the same French department (Ardèche) and belonging to a population with various individual forms in changing proportions. Wingspan of examined material of all three forms similar, ♂♀ 14–18 mm.

Subsp. alpinaria from French High Alps is described as [having forewings] ‘far more slender and even paler, with transverse lines reduced (Leraut 2009), or ‘paler and larger, but with considerable variability in size, coloration and pattern’ (Flamigni et al. 2016). The material examined by us supports the latter description, showing transitions to the darker populations from Basses Alpes and Alpes Maritimes and causing doubts on the taxonomic status. Male genitalia (European populations): Genitalia small. Uncus triangular, posteriorly narrow, truncate but edges rounded, rarely slightly concave posteriorly. Saccus broad, usually shortly projecting and truncate. Valva broad at base, narrow at tip, ventrally edged, shape of valva unique within the genus. Spinulose process of valva comparatively large, lobed, in rather distal position. Aedeagus very short (0.8–0.9 mm), slightly bent and narrow. Cornutus narrow, needle-shaped, long (0.35–0.4 mm), i.e. 0.3–0.45 times length of aedeagus. Genitalia of the nominotypical subspecies from North Africa similar, aedeagus slightly longer (0.9–1.1 mm) and basal sclerite of cornutus slightly broader.

Female genitalia: Antrum small, with two lateral sclerites. Sterigma with posterior edge narrowly sclerotized to transverse ridge. Ductus bursae membranous, in European populations comparatively short (0.7–0.9 mm) and narrow (in the Iberian peninsula somewhat broader), towards corpus bursae somewhat dilated, often longitudinally furrowed. Corpus bursae membranous, oval. Signum developed as flat sclerite with two small posterior spinules, as it is characteristic for the genus Eumannia. In subsp. castiliaria from Spain signum with transverse ridge, in France (subsp. cebennaria) without transverse ridge but with two more distinct spines. In the nominotypical subspecies from North Africa ductus bursae sigmoid, comparatively long (0.8–1.0 mm), corpus bursae more elongate, signum with transverse ridge and two minute spinules. Distribution and abundance: West Mediterranean. Nominotypical subspecies non-European. Subsp. castiliaria from the Iberian peninsula (Portugal, Spain, Andorra) to southernmost France. From French Central Massif to north-westernmost Italy (Liguria: Raineri 1986b; Raineri & Zangheri 1995; Arnscheid 2000; Flamigni et al. 2016) replaced by subsp. ceben­ naria. In the French High Alps replaced by subsp. alpinaria. Erroneous records (misidentifications) from Bulgaria (Ganev 1983b; Nestorova 1998; Müller 1996; Leraut 2009; Flamigni et al. 2016) and Macedonia (Ganev & Beshkov 1987). – In south-western Europe widespread, sometimes in numbers. – Outside Europe subsp. castiliaria in northernmost Morocco. From central and southern Morocco to Tunisia replaced by the nominotypical subspecies T. oranaria oranaria. Phenology: Usually univoltine, from mid-June to early August, exceptionally up to ten days later, at high elevation in the Alps emerging from early July. Larvae emerging from September, overwintering, until June. Reputedly from March to September (Leraut 2009), possibly partly based on misidentifications: such extended phenology data not confirmed by the comprehensive material examined by us. Adults active at night, females readily attracted to light, males apparently less, thus with very high ♀-rate at light (75–85%). 510

Biology: Larva found on and reared on lichens of Buxus sempervirens and Juniperus oxycedrus (France: Cleu 1933; Flamigni et al. 2016). Larva found on (beaten from) low plants, e.g. Thymus vulgaris (Prout 1915). Pupation on the ground between leaves or small stones in a loose silken cocoon (Chrétien 1898; Cleu 1933). Habitat: Xerothermophilous, often xeromontane. Dry slopes, scattered oak forest, scrubland with steppe vegetation, often in escarpment habitats on calcareous soils, with occurrence of juniper (Cleu 1933; 1957; Flamigni et al. 2016). In France and Spain from 300 m (subsp. cebennaria) up to 1500 m (subsp. alpinaria), exceptionally lower. In the Iberian peninsula up to 2200 m. Nominotypical populations in North Africa from 1500 m up to 2500 m. Similar species: In Europe, discrimination from congeners based on wing pattern is usually possible without problems. In North Africa confusion is possible with certain forms of the sympatric Tephronia fatimaria Bang-Haas, 1906 (with synonym ismailaria Oberthür, 1913), but the latter with postmedial line less dissolved and distally not bordered by whitish scales. Genetic data: Nominotypical E. oranaria Staudinger, 1892 (described from western ­Algeria; three specimens barcoded from Morocco) at a distance of 1.9% from European E. o. castili­ aria, with separate BIN (BOLD:ACM0415). European populations with BIN: BOLD:AAI0761 (n=9 from Spain, north-westernmost Italy, France, including holotype of cebennaria and the distribution areas of alpinaria and hubertcleui). Genetically homogeneous, infraspecific maximum variation very low (0.17%), not supporting the validity of the taxa cebennaria Chrétien, 1898, alpinaria Leraut, 2009 and hubertcleui Leraut, 2009 at subspecies rank. Nearest species: Tephronia fatimaria (Bang-Haas, 1906) from Morocco (5.9%), Eumannia oppositaria (6.7%) and E. arenbergeri Hausmann, 1995 from Cyprus (7.1%). In various tree constructing algorithms most closely clustering with E. arenbergeri. Remarks: The distribution borders of the three subspecies as tentatively (but questionably) validated here for Europe, are probably not sharply defined. 511

Combination with genus Eumannia is necessary because of a number of synapomorphies shared with E. oppositaria, e.g. short male antennal branches, fusion of hindwing veins Sc and Rs+M1, distal position of spinulose process of valva, shape of signum, similarity in the COI barcode etc. One male from Egypt (Natron valley) with a (very) short COI sequence of 270bp revealing sequence identity with E. oranaria in this DNA fragment. Although the moth corresponds to the description of Tephronia minutaria Turati, 1934: Atti Soc. ital. Sci. nat. 73 (1–2): 177, fig. (Libya, Cyrenaica: Barce) the taxon is not synonymized here with E. oranaria because the male genitalia strongly differ in the shape of uncus and valva.

Phyllometra Boisduval, 1840 Phyllometra Boisduval, 1840, Genera Index meth. eur. Lepid.: 193. Type species: Phyllometra gracilaria Boisduval, 1840, by monotypy. Synonymized with Tephronia, for the type species gracilaria, by Müller (1996), and then again by Leraut (2009) as ‘syn. n.’. Herewith reverted to validated genus rank (stat. rev.) because of separate COI-clustering and several significant synapomorphies (see below). Egea Duponchel, 1845, Cat. meth. Lepid. Eur.: 245. Type species: Idaea culminaria Eversmann, 1843, by subsequent designation (Prout 1910). Still retained as valid genus by Fletcher (1979), synonymized with Phyllometra in Scoble (1999). Eremia Herrich-Schäffer, 1847, Syst. Bearb. Schmett. Eur. 3: 35. Type species: Idaea culminaria Eversmann, 1843, by monotypy. Unavailable names (incorrect subsequent spellings): Eugea: Lederer (1853); Aegea: Rambur (1866); Ereuxa: Hedemann (1881).

Genus including five Palaearctic species, two in Europe and three others in North Africa and central Asia: P. teneraria (Staudinger, 1892) from central Asia; P. argentaria (Bang-Haas, 1910) with synonym planaria (Chrétien, 1911) from Tunisia; P. proutiana Turati, 1936 from Libya. Genus Phyllometra downgraded to synonymy of genus Tephronia in Expósito (1978) and then again by Leraut (2009), herewith re-established based on significant differential features (see below). Differential characters: ♂ antennae bipectinate to tip, branches longer (0.7–0.8 mm) than those of the genera Tephronia (0.4–0.6 mm) and Eumannia (0.2–0.4 mm). Palpi very short in both sexes (less than diameter of eye), proboscis vestigial. ♂ hindtibia with 4 spurs. Hindwing dark with pale postmedial fascia, fringe chequered. Female with reduced length of wings (transition to genus Ithysia?). Tympanum very small (specially in ♀), ansa minute, functionality for acoustic perception doubtful. Venation similar to that of Tephronia but on hindwing discal cell shorter, common stalk of Rs and M1 comparatively short. Hindwing without M2, discocellular vein weak. ♂ genitalia small, uncus round and broad, juxta and valva similar to equivalents in Tephronia, aedeagus short and broad, cornutus with broad base. ♀ genitalia without or with minute signum; if present, reduced to a short transverse ridge and/or a hardly visible minute sclerite. Biology lichens.

and immature stages:

Larva lichenophagous, with well camouflaged coloration on

Genetic data: DNA barcodes of the representatives of Phyllometra clustering together and significantly diverging from those of the genera Tephronia and Eumannia (cf. Text-fig. 200), independent of the chosen algorithm and tree construction settings (maximum likelihood bootstraps >95). 512

Remarks: Vasilenko (2005) figures the forewing venation for a group of central Asian species, i.e. Pleotrichia kurenica Vasilenko, 2005 and Autotrichia lysimeles Prout, 1934 as well as the male genitalia for A. lysimeles, A. karanguica Vasilenko, 2002 and P. kurenica. Venation and male genitalia of both ‘genera’ appear to match their equivalents in Phyllometra very well, suggesting a potential synonymisation of the two genus names Autotrichia and Pleotrichia with Phyllometra. All further features (except wing pattern) are similarly well matching, e.g. male hindtibia with four spurs, head hairy, proboscis vestigial etc. Close relationship was earlier recognized by Prout (1934) and Wehrli (1953). It is desirable to study female genitalia and DNA barcodes as well before taking a formal decision on this question.

179. Phyllometra culminaria (Eversmann, 1843) Idaea culminaria Eversmann, 1843: Bull. Soc. imp. Nat. Moscou 16: 552 (Russia, Urals: Majo). Until recently combined with Phyllometra (e.g. Müller 1996; Scoble 1999; Hausmann et al. 2004), transferred to Tephronia in Leraut (2009), herewith transferred back to Phyllometra based on the below mentioned differential features (comb. rev.). Fidonia desertaria Freyer, 1844: Neuere Beitr. Schmettkde. 5 (73): 95–96, pl. 438, fig. 2 (Russia). Syntypes.

External characters and abdomen: Wingspan ♂ 15–18 mm, forewing comparatively small, with apex tapered, ♀ with reduced wings, wingspan 12 mm only. Ground colour greybrown. Forewing with pale streak along the costa. Ante- and postmedial lines broken into black-brown dots, the latter distally bordered by white fascia on both wings. Terminal line proximally bordered by row of white spots on both wings. Fringe chequered brown and white. Hindwing dark with pale postmedial fascia. Palpi very short in both sexes. Proboscis vestigial. ♂ antennae bipectinate to tip, branches longer (0.7–0.8 mm) than those of genera Tephronia (0.4–0.6 mm) and Eumannia (0.2–0.4 mm). ♀ antennae filiform. ♂ and ♀ with two pairs of hindtibial spurs. Variation: Little variation all over the distribution area. Male genitalia: Genitalia similar to those of P. gracilaria, uncus more rounded, saccus shortly projecting, triangular. Valva broader in P. culminaria, central spinulose projection of valva broad, shallow, rounded. Aedeagus short (0.9–1.0 mm) and broad, cornutus large (0.55 mm), strongly sclerotized, with latero-basal sclerite. Female genitalia: Apophyses posteriores three times length of apophyses anteriores. Antrum trapezoid. Ductus bursae sclerotized and strongly swollen laterally, longitudinally wrinkled. Corpus bursae oval, membranous, signum absent. Distribution and abundance: Eurasiatic. In Europe restricted to a discontinuous area in the southern Urals, and in Hungary (Pilis, Veszprém). – In Europe very local and rare. – Outside Europe from Turkey and Turkmenistan to Kazakhstan and central Siberian Russia (Tuva, Altai, Sajan). Phenology: Univoltine, from late April to late May, exceptionally until June. Males active during daytime, occasionally attracted to light. ♀-rate in collections low.

Biology: Larval stages unknown. Larvae probably feeding on lichens like species of Tephro­ nia and Eumannia. Females with reduced potential for dispersal because of reduced wing length. K-strategy.

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Habitat: Xerothermophilous. Inhabiting open habitats on low-nutrient soils and on rocky, often calcareous ground with sparse, petrophilous vegetation, also on dry steppes and low, sparse grassland. Usually preferring lowlands, in Turkmenistan up to 1000 m above sea-level. Similar species: Allopatric P. gracilaria darker, larger, wings broader, forewing without pale streak along costa, postmedial line of forewing more strongly marked, pale distal border of postmedial line usually narrower. Genetic data: BIN: BOLD:AAE6536 (n=8 from Hungary, Turkey, Kazakhstan). Infraspecific variation low, despite the geographic distances (0.8%). Nearest species: P. gracilaria (2.6%). Remarks: Transferred to Tephronia in Leraut (2009), herewith reverted to combination with Phyllometra based on differences in wing pattern, external structure (e.g. length of antennal branches), genitalia, and dissimilarity of COI barcodes.

180. Phyllometra gracilaria Boisduval, 1840, comb. rev. Phyllometra gracilaria Boisduval, 1840: Genera et index methodicus europaeorum lepidopterorum: 193 (Spain: Andalusia). Syntype(s). Until recently combined with Phyllometra (e.g. ­Scoble 1999; Hausmann et al. 2004), transferred to Tephronia in Vives Moreno (1994) and Müller (1996), followed by Leraut (2009), herewith combined again with Phyllometra (stat. rev.). ‡ cacuminaria Rambur, 1866: Cat. syst. Lépid. Andal. 2: pl. 19, fig. 2 (Spain, Andalusia). Non binominal. Egea cacuminaria Prout, 1915 (“Rmb.”): in Seitz, Gross-Schmett. Erde 4: 4 fig. 1b. Validated at species rank by Prout (1915), thus made available nomenclaturally. Junior synonym.

External characters and abdomen: Wingspan ♂ 17–24 mm, ♀ 15–17 mm. Ground colour dark grey-brown. Ante- and postmedial lines broken into black brown dots, the latter distally 514

bordered by narrow white fascia. Terminal line proximally bordered by row of white spots on both wings. Fringe chequered brown and white. Hindwing dark with pale postmedial fascia. Palpi very short in both sexes. Proboscis vestigial. ♂ antennae bipectinate to tip, branches longer (0.7–0.8 mm) than those of genera Tephronia (0.4–0.6 mm) and Eumannia (0.2–0.4 mm). ♀ antennae filiform. ♂ and ♀ with two pairs of hindtibial spurs. Variation: Extent of white elements of wing pattern slightly variable.

Male genitalia: Genitalia similar to those of P. culminaria. Uncus small, often bilobed, rounded at tip. Saccus short, digitate or sub-truncate. Valva narrow. Central process of valva broad, shallowly rounded. Aedeagus short (0.75–0.8 mm) and broad, tapering to tip. Cornutus slightly curved, needle-shaped, weakly sclerotized, length 0.3–0.35 mm. Female genitalia: Apophyses posteriores twice length of apophyses anteriores. Antrum developed as rectangular sclerite. Ductus bursae very short (0.2 mm), longitudinally wrinkled. Corpus bursae globular or pyriform, membranous. Signum small, very weakly sclerotized, with transverse ridge. Distribution and abundance: (West-) European. Spain, with isolated occurrence in southern France. One record for Portugal (Cruz & Gonçalves 1974), treated as doubtful by Corley (2015). – Local, rarely abundant at the sites of occurrence.

Phenology: Univoltine. In the lowlands from late April to early June (exceptionally emerging from early April), in the mountains from early May to late June, single females until late July. Males active during daytime, occasionally attracted to light. ♀-rate in collections low.

Biology: Larval stages unknown. Larvae probably feeding on lichens like species of Tephronia and Eumannia (cf. Leraut 2009). Females with reduced potential for dispersal because of reduced wing length. K-strategy. 515

Habitat: Xeromontane. Preferring open habitats, steppes, dry grassland, scattered scrubland on hills and mountains. Usually from 800 up to 1600 m a.s.l., in the steppes of the Ebro plain (prov. Huesca, e.g. Candasnos) down almost to sea-level. Similar species: Allopatric P. culminaria (see no. 179). Easily distinguishable from sympatric species of Tephronia by the chequered fringe, the pale postmedial line of hindwing etc. Genetic data: BIN: BOLD:AAH8468 (n=3 from Spain). Genetically homogeneous. Nearest species: P. culminaria (2.6%). Distance from central Asian P. teneraria 8.2%. Remarks: Genus Phyllometra (including the taxon gracilaria) subordinated under Tephronia in several works (e.g. Expósito 1978; Leraut 2009), here not accepted considering the dissimilarities in wing pattern, external structure and separate clustering of DNA barcodes.

Ithysia Hübner, 1825 Ithysia Hübner, [1825] 1816, Verz. Bekannter Schmett.: 319. Type species: Geometra pravata Hübner, 1813, by subsequent designation by Fletcher (1979).

Monotypic, one species in the western Palaearctic region. The classification as monotypic genus and the isolated position in phylogeny supported by DNA data (see below). The position beside Tephronia (specially ‘Phyllometra’) supported by similar wing pattern, wing shape, reduction of wing size in females and ecological niche as ‘winter moths’ (incl. late autumn and spring) of steppes. Differential external characters and abdomen: ♀ brachypterous (almost apterous). Forewing narrow, apex pointed. Forewing postmedial line strongly curved. Thorax and palpi hairy, the latter longer than in Tephronia. Abdomen white at tip. ♂ antennal branches shorter than in Tephronia. Venation: Discal cells shorter than in Tephronia and Eumannia. On forewing Sc and R1–3 reaching costa, Sc free or shortly touching R1. R1 and R2 arising separately from cell. R3–R5 shortly stalked, connate with M1 or shortly separate. On hindwing Sc+R1 not fused with cell. Rs and M1 shortly separate. M2 absent. M3 and CuA1 connate. Abdomen: ♂ abdomen very long, ♂ sternites sclerotized, tergum A1 and A2 broad, strongly sclerotized, tergum A2 with long and narrow lateral appendages. Tergites and sternites of ♀ membranous. ♂ sternum A8 trilobed, tergum A8 triangular, elongate. Tympanum in both sexes vestigial, almost absent. Genitalia: See description of species. Biology and immature stages: Larva apparently polyphagous. Pattern of larval cuticle with numerous longitudinal lines (cf. Budashkin & Savchuk 2013). Adults diurnal. Genetic data: Analysis of DNA barcodes (COI 5’) not revealing any closely related genera, supporting the hypothesis of an old phylogenetic lineage (see below, under Remarks). Remarks: The small tympanal organs of Tephronia, Eumannia and Phyllometra may suggest relationship to Ithysia where the tympanal organs are almost absent, which is a very rare character state in Geometridae. However, external morphology, venation, genitalia and COI barcodes do not support a closer relationship and rather suggest an independent origin of tympanum reduction (analogy). However, small tympanal organs and morphology of male and female genitalia also reminiscent of genus Lignyoptera (P. Sihvonen own observation), probably pointing to true relationships. In the multi-gene analysis of Murillo-Ramos et al. (2019) clustering with the “Apeirini + Hypochrosini + Epionini” complex of tribes, as a separate lineage. 516

181. Ithysia pravata (Hübner, 1813) Geometra pravata Hübner, 1813: Samml. Eur. Schmett. 5 (1): pl. 84, fig. 432. Syntype(s) lost. Geometra pravaria Hübner, 1825: Verz. Bekannter Schmett.: 319. Emendation.

External characters and abdomen: Wingspan ♂ 19–22 mm, ♀ brachypterous (almost ­apterous). Male: Forewing narrow, apex pointed. Ground colour of forewing brown, along veins white, with curved brown postmedial line from inner margin near wing base to costa near apex. Cell spot large and distinct on forewing. Hindwing with extended whitish ground colour, darker towards termen, postmedial line brown, discal spot present. Thorax with hairy scales. Thorax and abdomen dark, whitish towards posterior tip. ♂ antennae shortly bipectinate, branches approx. three times width of flagellum. Palpi comparatively long, hairy. Proboscis absent. Hindtibia of both sexes with four spurs. Venation and abdomen: see description of genus. Variation: Little variation, one examined specimen from Sarepta with reduced dark pattern, just the postmedial line, the fine terminal line and the cell spot remaining. Male genitalia: Uncus long, narrow, strongly sclerotized. Socii minute, setose. Gnathos present, weakly sclerotized. Juxta with two broad digitate postero-ventral projections, terminally rounded. Saccus shortly projecting. Valva long spatulate, anterior and posterior margins parallel, rounded at tip, without appendages, costa more strongly sclerotized. Aedeagus narrow, length 1.1–1.2 mm, tapering at tip. Terminal half with row of 5–7 small cornuti. Female genitalia: Ovipositor short. Apophyses narrow, apophyses posteriores twice length of apophyses anteriores. Lamella antevaginalis sclerotized in narrow band. Antrum funnel-shaped, membranous. Ductus bursae narrow, short (0.5 mm). Corpus bursae broader, conical, divided into a slightly sclerotized posterior half and a delicately membranous anterior half. Signum absent. Distribution and abundance: East European. In Europe restricted to Ukraine (Crimea: ­Budashkin & Savchuk 2013), Volga plain and southern Ural region. – Usually rare and ­local. – Not yet recorded from outside Europe (Viidalepp 1996), but occurrence in northern steppes of Kazakhstan not excluded. Phenology: In the Crimea from late September to late November (Budashkin & Savchuk 2013). One adult in ZSM labelled as “May”, perhaps by error. Larvae under rearing conditions hatching from mid-April and moths emerging from September. Adults active during daytime, occasionally attracted to light. Biology: Larva probably polyphagous. Reared on Jurinea stoechadifolia, J. sordida, Artemisia santonica, A. taurica, and Galatella villosa (Budashkin & Savchuk 2013). Habitat: Xerophilous, lowlands. Inhabiting steppes, dry grassland. From sea-level up to approx. 200 m. Similar species: No similar species. Genetic data: BIN: BOLD:ACD5536 (n=4 from Ukraine). Genetically slightly heterogeneous, maximum variation 1.1%. Analysis of COI not revealing any closely related genera, supporting the hypothesis of an old phylogenetic lineage. Nearest species (long branch attraction): Australian Crypsiphona ocultaria (Donovan, 1805) (6.7%), belonging to subfamily Geometrinae. Genetic distance from the closest ally in the Tephronia genus group

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(Tephronia lepraria) 8.4%, distance from Lignyoptera fumidaria 10.4%. Hence, the systematic relationship between Ithysia and Lignyoptera (as suspected from morphology) not supported by similarity in the COI gene. In the multi-gene analysis of Murillo-Ramos et al. (2019) clustering with the “Apeirini + Hypochrosini + Epionini” complex of tribes, as a separate lineage.

Text-fig. 213: Wingless female of Ithysia pravata (Hübner, 1813) (Ukraine; photo by the courtesy of V. Savchuk).

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New species for the fauna of Europe after publication of the previous volumes In this section we present 21 new species for the fauna of Europe, discovered after the publication of the related GME volume (Hausmann 2001; Mironov 2003; Hausmann 2004; Hausmann & Viidalepp 2012; Skou & Sihvonen 2015). A few of them have been described as new to science (improved taxonomy), others are new to the fauna of Europe (described earlier and later found in Europe). In eleven additional chapters important taxonomic changes are briefly mentioned and explained. Updated species lists for the six GME Volumes: GME1 (Archiearinae to Geometrinae), Hausmann 2001: Three new species recorded here, three species (Alsophila, Epirranthis) moved to Ennominae (actual total: 42). GME2 (Sterrhinae), Hausmann 2004: Seven new species recorded here, four species (Lythria) added in Vol. 3 – one species now synonymized (actual total: 206). GME3 (Larentiinae I), Hausmann & Viidalepp 2012: One new species recorded here, four species (Lythria) belonging to Sterrhinae (actual total 265). GME4 (Larentiinae II), Mironov 2003: Eight new species recorded here (actual total 159). GME5 (Ennominae I), Skou & Sihvonen 2015: Two new species recorded here, three species (Alsophila, Epirranthis) added from Vol. 1 (actual total 146). GME6 (Ennominae II): present volume: 181 species. With these new faunistic records and improved taxonomy, the number of European geometrid species is raised to 999 (cf. summary to the complete checklist in this volume).

Archiearinae Leucobrephos Grote, 1874 Leucobrephos Grote, 1874: Bull. Buffalo Soc. Nat. Sci. 2: 53. Type species: Anarta brephoides Walker, 1857 by original designation (Canada).

External characters and abdomen: Ground colour of hindwing whitish or pale grey. Anten­ nae shortly bipectinate in male, filiform in female. Palpi long, palpi and head densely covered with hairy scales. Abdomen densely covered by long, bifurcate scales dorsally and laterally. Tympanum with ansa broad at base, conically narrowing towards tip. Male genitalia: Gnathos present, weak. Socii absent. Valva long, narrow, curved. Aedeagus without cornuti. Female genitalia: Papillae anales with fine setae and apophyses anteriores long as in Boudinotiana (thus unlike Archiearis). Ductus bursae and posterior part of corpus bursae narrow, long. Signum round. Habitat: Associated with cold habitats in polar tundra and mountainous valleys with riverside vegetation. Remarks: Two species in the northern parts of the Holarctic region. In multi-gene analysis Leucobrephos is the most basal Archiearinae taxon, sister to Boudinotiana + Archiearis lineage (Murillo-Ramos et al. 2019).

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182. Leucobrephos middendorfii (Ménétriés, 1858) Amphidasis middendorfii Ménétriés, 1858: Bull. classe phys.-mat. Acad. impériale Sci. St.-Pétersbourg 17 (12–14): 221 (eastern Russia, eastern Siberia: Udskoe; Vilui). Lectotype ♂ (ZISP, designated by V. Mironov, examined). Brephos ussuriensis Moltrecht, 1914: Ezheg. gosud. Muz. N.M. Mart’yanova 2 (1): 73 (eastern Russia, Ussuri Region: 300 km north of Vladivostock). Syntypes 5♂. Validated at subspecies rank (Scoble 1999), but the type locality relatively close to that of the nominotypical subspecies suggesting potential synonymy. Leucobrephos middendorfii nivea Kozhantschikov, 1924: Ent. Rd.sch. 31 (6): 34 (central Russia, Sajan region: River Tansybei). Syntype(s). Valid at subspecies rank (Scoble 1999). Leucobrephos mongolicum Vojnits, 1977: Ann. hist.nat. Mus. natn. Hung. 69: 166 (Mongolia: Chentej Aimak Aduein). Validated at species rank (Scoble 1999), recently downgraded to synonymy (Beljaev 2016). Unavailable names: (incorrect subsequent spelling): middendorffii: Ménétriés (1859), followed by Leraut (2009).

External characters and abdomen: Wingspan ♂ 25–28 mm, ♀ 24–25 mm. Female forewing shorter than that of male, at apex more rounded. Ground colour of forewing dark grey, of hindwing whitish. Forewing with blackish ante- and postmedial lines. Postmedial line zigzagging and projecting towards termen in centre, distally bordered by whitish or ochreous fascia. Terminal area dark. Hindwing with broad, black band at termen and at inner margin. Underside of forewing dark grey, with much broader postmedial fascia than on upperside, with small rectangular spot subapically on costa. Hindwing as described for upperside. Fringe weakly chequered white and grey. Female with ground colour of forewing grey, paler than in male, ante- and postmedial lines well contrasted. Hindwing pattern as in male but ground colour with stronger dark suffusion. Eyes small, vertically elongate, tapering ventrally. Antennae of ♂ shortly bipectinate, in ♀ filiform, flagellum black, each flagellomere with white scales, appearing slightly lamellate. Palpi long, palpi and head densely covered with hairy scales. Variation: Varying in the width of the blackish terminal area of hindwing, and in the density of the dark suffusion along the inner margin of hindwing. Male genitalia: Uncus short and broad, triangular. Gnathos present, weakly sclerotized. ­ ocii absent. Saccus broadly rounded. Juxta large, trapezoid, posteriorly with short projecS tion. Valva long, narrow, strongly curved in distal third. Aedeagus narrow, curved, without cornuti. Everted vesica showing minute spinules at base. Female genitalia: Papillae anales with fine setae. Apophyses anteriores comparatively stout, long (1.0 mm), 2/3 length of apophyses posteriores. Tergum A8 trapezoid. Sternum A7 (lamella antevaginalis) a comparatively broad band. Ductus bursae and posterior part of corpus bursae narrow, long, longitudinally wrinkled, antrum dilated funnel-shaped, more strongly sclerotized. Anterior part of corpus bursae oval. Signum at dorsal side of corpus bursae, round, with shallow, sclerotized teeth at anterior margin. Distribution and abundance: Siberian. In Europe single specimens recorded in the Polar Urals (K. Nupponen pers. comm.). – Very rare and local in Europe. – Outside Europe in northernmost Asia (Siberia) to Far East Russia. In North America replaced by the allopatric sister species L. brephoides (Walker, 1857). Phenology: Univoltine, scarce data from late April to mid-May. Adults active during daytime, starting to fly when ground is still covered by snow (K. Nupponen pers. comm.). 520

Biology: Larval stages unknown. Habitat: Riverside vegetation in partly forested polar tundra, flying around in the first thawed patches in the Betula zone (K. Nupponen pers. comm.). Similar species: No similar species in Europe. North American sister species L. brephoides (Walker, 1857), described from north-eastern Canada similar, but with darker, blackish grey forewings and pale ante- and postmedial lines. Genetic data: Not yet DNA barcoded. North American sister species L. brephoides with BIN BOLD:AAH0638. Nearest species: Phallaria ophiusaria Guenée, 1858 (Oenochrominae) from Australia (6.1%), which is a small distance for species currently classified in two different subfamilies. Nevertheless, the observed genetic similarity should be due to long branch attraction, not to true phylogenetic relationship.

Desmobathrinae 183. Myinodes constantina (Hausmann, 1994) Myinodes constantina Hausmann, 1994: Nota lepid. 17 (1/2): 36, figs 1, 2, 9, 11a-d, 15 (eastern Algeria: Lambèse). Holotype ♂ (ZFMK, examined and dissected Hausm. 7910).

External characters and abdomen: Wingspan ♂ 26–29 mm, ♀ 20–26 mm. Postmedial line not dentate, at inner margin strongly inclined towards wing base. White intervenal line in forewing apex very short, length about 1/3 of subterminal area. Postmedial and intervenal lines indistinctly bordered. Antemedial line absent. Small terminal dots black, punctiform. Intervenal lines near termen neither forked nor surrounding terminal dots. Hindwing pale.

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♂ antennae ciliate, length of cilia (0.18 mm) 1.5 times width of flagellum. Frons with one central projection. Palpi long in both sexes (1.4–1.6 mm).

Variation: Little variation observed so far. Male genitalia: Uncus long, with very small subapical projection. Juxta with very deep posterior notch, base convex, latero-posterior lobes tapering. Valva with sinus-shaped costa. Harpe prominent, less pointed than in M. interpunctaria, strongly curved, without spinules. Basal lobe of harpe absent. Aedeagus narrow, very long (1.8–2.0 mm), with smooth, digiti­ form, sclerotized process (without teeth). Cornutus broader and more strongly sclerotized than in M. interpunctaria. Female genitalia: Ductus bursae long, straight. Corpus bursae small, irregularly shaped, not longitudinally ribbed. Distribution and abundance: West Mediterranean. Recorded as new for the fauna of ­Europe in central Spain by Expósito Hermosa (2006), confirmed by King (2008). – In Europe rare and very local. – Outside Europe in north-west Africa from Morocco (Middle Atlas) to north-western Tunisia (Hausmann 1994b).

Phenology: In Europe late February to mid-March. In North Africa March and April, exceptionally earlier or later. Biology: Larval stages unknown. Habitat: Xerothermophilous. In Europe, so far, only collected in a nature reserve near ­Madrid with halophilous and gypsicolous vegetation and hot microclimate. In Europe at 550 m a.s.l., in North Africa from sea-level up to 1100 m. 522

Similar species: Palpi in both sexes considerably longer than in the two congeneric species M. interpunctaria (Herrich-Schäffer, 1839) and M. shohami Hausmann, 1994. Forewing of the two last-mentioned species with dentate postmedial line, at inner margin not strongly inclined towards wing base, white intervenal line of forewing apex longer. M. interpunctaria furthermore differing in a double projection of frons, and in triangular terminal dots, encircled by the forked, white intervenal line, hindwing not considerably paler. Differential analysis of genitalia and text-figs for external features see Hausmann (1994b). Genetic data (cf. Hausmann et al. 2013): BIN: BOLD:AAG5649 (n=2 from central Spain), infraspecific variation low (0.16%). So far, no barcodes available from North Africa. Nearest species: M. shohami from Israel (8.1%) and M. interpunctaria from Spain (8.2%).

Text-figs 214–215. Diagnostic characters (indicated) for wing-pattern-based identification of Myinodes species: Text-fig. 214. M. interpunctaria (Herrich-Schäffer, 1839) (Morocco, Fez, 11.iii.1929). Text-fig. 215. M. constantina Hausmann, 1994 (Tunisia, Thala, vi.1911).

Subfamily Geometrinae Tribus Hemitheini The tribes Hemistolini, Comostolini, Jodini, Thalerini, and Microloxiini were validated in GME1 (Hausmann 2001), but later on these were downgraded, together with several non-European tribes, to subtribe rank or synonymy (Beljaev 2016; Ban et al. 2018).

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Kuchleria insignata Hausmann, 1995 Kuchleria insignata Hausmann, 1995: Atalanta 25 (3/4): 584, figs 15, 41, 47 (south-eastern Spain, Alicante: Cabezon de Oro-Busot). Holotype ♂ (ZSM, examined, dissected and DNA barcoded). Kuchleria garciapitai Expósito, 2005: Shilap Revta. lepid. 33 (132): 437, figs 1, 4, 6, 8. (Spain, Madrid: El Regajal, Aranjuez). Holotype ♂ (coll. Expósito, Madrid; one topotypic specimen DNA barcoded). Accepted at species rank in Redondo et al. (2009), downgraded to synonymy in Leraut (2009), no more listed as separate species in Redondo et al. (2015), because of the mentioned differential features revealing as not significant. Synonymy herewith confirmed by the shared COI sequence (cf. Hausmann et al. 2013).

For full treatment of K. insignata with figures of adults and genitalia see Hausmann (2001). Leraut (2009) subordinated the taxon insignata under K. menadiara without justification (‘in my view synonym’). Herewith reversed to species rank based on considerable differential features in morphology and genetics (see under K. menadiara). The taxon garciapitai was described as separate species from central Spain (Expósito Hermosa 2005b), based on slight differences in male genitalia (bifid saccus). Molecular data (COI barcoding, including one topotypic specimen, provided by the courtesy of A. Expósito Hermosa) show these minor differences to fall under the variation of K. insignata (Hausmann et al. 2013).

184. Kuchleria menadiara (Thierry-Mieg, 1893) Phorodesma menadiara Thierry-Mieg, 1893: Naturaliste 15 (2, 142): 40 (north-eastern Algeria: Bône). Holotype ♂ (MNHN). Microloxia menadiara atlagenes Prout, 1935: in Seitz, Gross-Schmett. Erde 4, Suppl.: 16 (­Morocco, High Atlas: Tinmel). Holotype ♂ (NHMUK, examined). Valid at subspecies rank (Hausmann 1995c; Scoble 1999). Microloxia menadiara ephedrae Prout, 1935: in Seitz, Gross-Schmett. Erde 4, Suppl.:16 (­Morocco, Middle Atlas: Bekrit). Holotype ♂ (NHMUK, examined). Valid at subspecies rank (Hausmann 1995c; Scoble 1999). Unavailable names: (incorrect subsequent spellings): menadaria: Staudinger, 1901; menadiaria: Prout, 1913.

External characters and abdomen: Wingspan ♂ 19–24 mm, ♀ 24–31 mm. Ground colour deep green when fresh, often faded to yellowish green in collections. Fringe concolorous. Postmedial line of both wings broken into slightly waved row of usually conspicuous white dots. Antemedial line of forewing inconspicuously marked by 2–4 white dots. Forewing costa white, sometimes with rosy tinge. Frons flat, unicolorous green, vertex white. Palpi narrow, green, tip reddish. Length of ♂ palpi 1.2–1.3 times diameter of eye. Variation: Slight variation in the intensity of the dotted postmedial lines. Male genitalia (Sicily): Length of aedeagus 1.75 mm. Aedeagus with large distal patch of microcornuti (differential feature from the West Mediterranean sister species K. insignata), fused with central patch forming together a very large field. Cornuti comparatively longer than in K. insignata. See also Hausmann (1995c; 2001). Female genitalia (Sicily): Apophyses anteriores shorter (0.35–0.4 mm) than in K. insignata. Sterigma triangular. Corpus bursae pyriform. See also Hausmann (1995c; 2001).

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Distribution and abundance: (Central) Mediterranean. Recently recorded in Sicily (Parenzan et al. 2006; P. Skou own data; J. Bittermann pers. comm.). – Local and usually rare, occasionally occurring in numbers. – Outside Europe in the Mediterranean parts of eastern Algeria and northern Tunisia (nominotypical subspecies), subsp. ephedrae in the Moroccan Middle ­Atlas, subsp. atlagenes in the Moroccan High Atlas, the taxonomic status of both taxa requiring further study with more material.

Phenology: In Sicily bivoltine, recorded from early June to late June and from mid-August to mid-October. In North Africa emerging from mid-April. Biology: Larvae associated with Ephedra, found on Ephedra major in Morocco (Prout 1935a: ‘E. nebrodensis’). In Sicily E. fragilis occurs at the sites of occurrence and is likely to be the larval food-plant there. ♀ ratio at light low, about 10–20%. Habitat: Xerothermophilous. In Europe collected in dry and hot habitats on sandy soils, usually near the coast, from sea-level up to 400 m.

Similar species: All congeneric species very similar in habitus (cf. Hausmann 1995c; 2001), The allopatric sister species Kuchleria insignata on average smaller, the postmedial line usually less distinct, with differential features also in the genitalia of both sexes. K. therapaena (Prout, 1924) from eastern Algeria and Tunisia differs in the white forewing costa, and in the more pointed forewing apex. Large females of Microloxia herbaria may be similar too, but their antennae are shortly bipectinate, frons laterally with red scales. Genetic data (cf. Hausmann et al. 2013): BIN BOLD:ABZ0774 (n=6 from Sicily and Algeria). Infraspecific variation low for the Sicilian and Algerian specimens (0%), distance from three barcoded northern Tunisian specimens 0.7%. Nearest species: K. menadiara atlagenes from Morocco (1.7%), K. insignata Hausmann, 1995 from Spain (2.2%, n=6), and K. gisisi 525

Hausmann, 1995 from Israel (2.6%; n=2) the last two having own BINs (BOLD:AAJ3108 / BOLD:AAX0736). One specimen from central Tunisia, Sfax (M. therapaena) diverging by 6.2%. Remarks: The differential diagnosis between K. menadiara and K. therapaena still needs further investigation with more ample material (cf. Hausmann et al. 2013). The locus typicus of the former in north-easternmost Algeria, with its Mediterranean vegetation and habitat types, corresponds much better with the Sicilian landscape than the locus typicus of the latter which is an oasis at the border of the Sahara desert (central Tunisia: Gafsa).

Chlorissa pretiosaria (Staudinger, 1877) Nemoria pretiosaria Staudinger, 1877: Stettin. ent. Ztg. 38 (4–6): 202 (southern Caucasus). Holotype ♂ (MNHU), examined.

Remarks: Reputedly recorded as new for the fauna of Europe from Astrakhan desert, southern European Russia (Anikin pers. comm.), subsequently posted on the website for European Geometridae (www.zsm.mwn.de/lep/gme) and mentioned in Hausmann et al. (2013). Later on corrected as misidentification (V. Zolotuhin pers. comm.; Anikin et al. 2017). Hence to be excluded from the European fauna, based on misidentification. Locus typicus in southern Caucasus, possible occurrence in Europe (southernmost European Russia), however, not excluded.

Subfamily Sterrhinae Tribus Sterrhini 185. Idaea barbuti Tautel & Lévêque, 2013 Idaea barbuti Tautel & Lévêque, 2013: Alexanor, 25 (6), 2012 (2013), 363, figs 1, 2, 47, 53 (Portugal, Région de Minho, District de Braga, Tebosa). Holotype ♂ (MNHN, DNA barcoded BC AL Geo 00488).

External characters and abdomen: Wingspan ♂ 17 mm ♀ 19–20 mm. Ground colour beige, ante- and postmedial lines of forewing dark brown, distinct, zigzagging, medial line indistinct. Terminal area of forewing with two conspicuous central spots and another blotch at apex. On forewing costa three dark spots at endings of transverse lines. Hindwing with similar pattern. Discal spots blackish, usually distinct. Terminal dots black, distinct, elongate. Fringe concolorous with ground colour. Frons grey, vertex creamy white. ♂ hindtibia without spurs. Variation: Little variation. Females with forewing apex slightly more rounded. Male genitalia: Very similar to those of I. alyssumata (cf. Hausmann 2004), cornuti in I. barbuti somewhat stouter, but with great variation in number. Length of aedeagus approx. 1.7 mm in I. barbuti, but shorter (1.25–1.4 mm) in I. alyssumata (Tautel & Lévêque 2013). Female genitalia: Very similar to those of I. alyssumata (cf. Hausmann 2004), I. barbuti with smaller spinules on the semicircular ring on the corpus bursae. Some of the differential features mentioned in Tautel & Lévêque (2013) are very small and doubtfully constant.

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Distribution and abundance: West Mediterranean. Endemic to western Iberian peninsula, so far only recorded for northern Portugal (type locality: Braga distr.), and reputedly in western Spain (‘La Granja’, prov. Caceres: Tautel & Lévêque 2013; Lepiforum 2017). Possibly ­occurring in the adjacent regions of Spain as well.

Phenology: Scarce data from late July to mid-August, thus apparently univoltine, but bivoltinism not excluded (Tautel & Lévêque 2013). Biology: Larval stages unknown. Habitat: The type locality is at 160 m a.s.l., the putative occurrence at La Granja at about 400 m. Similar species: Idaea alyssumata (Millière, 1871) – so far without confirmed sympatry – with less contrasting wing pattern, discal dot absent or very weak, slightly different in genitalia (see above), clearly diverging in the DNA barcode (see below). The allopatric Idaea typicata with one male hindtibial spur and different genitalia. Genetic data (cf. Hausmann et al. 2013): BIN BOLD:AAL5780 (n=6 from Portugal; all part of the type series), infraspecific variation low (0.47%). Nearest species: Idaea alyssumata at a minimum distance of 3.5%. The latter barcoded from Spain and France (n=13), diverging into five (!) different BINs. Remarks: Despite the unusual zoogeographical distribution (endemic to northern Portugal and westernmost Spain with a sister species widely distributed on the Iberian peninsula) the species rank for Idaea barbuti is supported by COI barcodes. The differences in morphology, i.e. wing pattern, size, minor differences in male and female genitalia as stressed in the original description are prone to variation and require re-examination based on more 527

ample material (M. Corley pers. comm.). If species rank for I. barbuti can be confirmed, the taxonomy of the (so far) five major genetic lineages (BINs) recorded for Idaea alyssumata is in question as well and needs to be investigated in more depth.

Idaea incisaria albarracina (Reisser, 1934), stat. rev. Ptychopoda incisaria albarracina Reisser, 1933: Eos 9: 250 (Spain, Aragon: Albarracin). Syntypes (SMNK, one syntype examined by R. Trusch; other syntypes at NHMW).

Remarks: In Hausmann (2004) the taxon albarracina was proposed to be validated at species rank, based on slight morphological differences. New data from DNA barcoding (Hausmann et al. 2013: divergence 0.16% = 1bp only), however, suggest the taxon should be downgraded from species rank to subspecies rank under Idaea incisaria, as it was earlier classified in Scoble (1999).

186. Idaea textaria (Lederer, 1861) Acidalia textaria Lederer, 1861: Wien. ent. Monatschr. 5 (5): 153, pl. 2, fig. 3 (southern Turkey, Hatay: Antiochia). Holotype ♀.

Genetic data (cf. Hausmann et al. 2013): BIN BOLD:AAD4546 (n=8 from Samos, southern Turkey and western Syria). Genetically homogeneous, intraspecific variation low (0.3%). Nearest species: Sister species Idaea laszloi Hausmann, 1999 from Turkmenistan (1.2%), Idaea striolata (Staudinger, 1892) from Lebanon and Israel (3.0%). Nearest European species: Idaea obliquaria (4.0%) and Idaea incisaria (4.5%).

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Remarks: For full treatment of this species with figures of adults and genitalia and for the suggested potential occurrence in Europe see Hausmann (2004). The species was recorded for the first time on European territory on the island of Samos by Fritsch et al. (2014) as well as on the island of Chios (Taylor 2018; D. Fritsch pers. comm.). On Samos polyvoltine, with records in late March, mid-May to late June, late August to early October, mid-­November, overwintering as larva, vertical distribution up to 300 m above sea-level, exceptionally up to 500 m (D. Fritsch pers. comm.). Reared on Thymus and Origanum (D. Fritsch pers. comm.).

Idaea humiliata davidi Gastón & Redondo, 2005, stat. n. Idaea davidi Gastón & Redondo, 2005: Bol. Soc. Ent. Aragonesa 36: 51 (Spain, Andalusia: Sierra Nevada, Ruta de Veleta). Holotype ♀ (MNCN Madrid). Herewith downgraded from species to subspecies rank, quoting the minor divergence in the DNA barcodes (see below).

Genetic data (cf. Hausmann et al. 2013): BIN BOLD:ACF2945. DNA barcodes from southern Spain (subsp. davidi) diverging from the main cluster of I. humiliata (n=13 from six countries from western, central and northern Europe) by only 0.7% whilst Greek and southern Italian populations of I. humiliata diverge by 1.7%. Remarks: Justification of species rank for the Andalusian (Sierra Nevada) populations based on weak evidence and recently questioned following the results of DNA barcoding (Hausmann et al. 2013). In wing pattern and coloration without significant and constant differ­ ences from its allopatric sibling species, Idaea humiliata (Hufnagel, 1767) (cf. Hausmann 2004). However, the question merits further attention and analysis. In allopatric distributions the decisions on classification at species or subspecies rank are inevitably subjective (Mutanen et al. 2012). For full treatment of the Andalusian populations with figures of adults and genitalia see Hausmann (2004) and Gastón & Redondo (2005).

187. Idaea lobaria (Chrétien, 1909) Acidalia lobaria Chrétien, 1909: Le Naturaliste 31: 30 (south-eastern Algeria: Biskra Bi Mora). Lectotype ♂ (ZSM, examined; DNA barcoded BC ZSM Lep 84175: failure). Lectotype designated in Herbulot (1982). Acidalia balestraria Lucas, 1909: Bull. Soc. ent. Fr. 1909 (4): 72 (Tunisia: Tozeur). Lectotype ♀ (ZSM, examined; barcoded BC ZSM Lep 84175: 94bp). Lectotype designated and synonymy recognized in Herbulot (1984).

External characters and abdomen: Very small, wingspan ♂♀ 10–12 mm, exceptionally somewhat larger, but not reaching ‘15 mm’ as stated in the original description of lobaria (type specimens examined and found to measure 12 mm). Forewing narrow, hindwing margin strongly concave between M1–M3. Ground colour creamy white, sometimes with pale brown suffusion. Transverse lines brown, ante- and postmedial line sometimes sharp. Terminal area pale brown with paler wavy line. Cell spots black, sharp, usually conspicuous. Antennae of ♂ ciliate-fasciculate, cilia curved, 2.5 times width of flagellum. Hindtibia slender, tarsus slender, not shortened, 0.8–1.1 times length of tibia. Variation: Ground colour sometimes with pale brown suffusion. Medial line sometimes conspicuously dark. Populations of the Levant often with darker brown terminal area, well constrasting on all wings.

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Male genitalia: Uncus short, triangular, at apex narrowly truncate. Saccus shortly projecting. Valva shortly spatulate, length 0.6–0.7 mm, without spinules at tip. Length of aedeagus 0.75–0.85 mm, narrow, straight. Aedeagus without cornuti in North Africa and the Levant, with narrow, straight, weakly sclerotized cornutus in Spain. Female genitalia: Apophyses posteriores (0.3 mm) twice length of apophyses anteriores (0.15 mm). Antrum laterally slightly swollen, sclerotized, in the Spanish subspecies laterally much more strongly swollen. Ductus bursae and posterior part of corpus bursae longitudinally furrowed. Corpus bursae pyriform. Distribution and abundance: South Mediterranean. In Europe restricted to ­south-easternmost Spain. Mentioned for the European fauna in Hausmann et al. (2013), based on one single female (dissected and DNA barcoded) recorded in south-eastern Spain, near Murcia. More data are presented in Guerrero et al. (2018) and Lepiforum (2018), including a record from prov. Almeria. – In Europe rare and local. – Outside Europe from south-western Morocco (ZSM, barcoded) to Tunisia (ZSM) and Libya (coll. P. Skou), and in the southern Levant (Sinai, southern Israel, Jordan; ZSM).

Phenology: Bivoltine: In Europe recorded from late June to mid-July and in early October (Guerrero et al. 2018). In North Africa scarce data from early May to early June, in Libya in mid-August (P. Skou own data). In the Levant scarce data indicating bivoltinism: from early April to late June; mid-October to mid-November. Biology: Larval stages unknown. Habitat: Xerothermophilous. Inhabiting deserts, semi-deserts and oases. In Spain recorded from 80 m up to 150 m above sea-level (Guerrero et al. 2018). In North Africa from sea-­ level up to 1500 m in Morocco (Anti-Atlas). 530

Similar species: Sympatric Idaea fractilineata sometimes very similar, in the spring generation larger (wingspan ♂♀ 12–16 mm), discal dots small, usually less conspicuous, male hindtarsus shortened, 0.4–0.5 times length of tibia. In ♂ genitalia aedeagus with cornutus. In ♀ genitalia antrum not laterally swollen, ductus bursae without longitudinal furrows. Genetic data (cf. Hausmann et al. 2013): BIN BOLD:ACD0706 (n=1 from Spain; A. Ortiz pers. comm.). DNA barcodes from North Africa and Israel (n=8) clustering to two different BINs (BOLD:ABZ0361 and BOLD:AAE5597) at distances of 1.9% and 2.8% from the Spanish specimen. Nearest species in Europe: I. consanguiberica (5.5%), I. belemiata (5.8%). The species was tentatively listed in the infirmaria species-group in Hausmann (2004), which is not well supported by the genetic data (see above). It is probably the product of a phylogenetically isolated lineage. Genetic distance from I. infirmaria 6.4%. Remarks: Potential divergence at species rank from North African populations requiring further study and more material.

188. Idaea gelbrechti Hausmann, 2003 Idaea gelbrechti Hausmann, 2003: Entomologische Zeitschrift Stuttgart 113 (11): 322, figs 6, 18, 19 (Morocco, Middle Atlas: Ifrane prov. 33.32 N, 5.07 W). Holotype ♂ (ZSM, examined, DNA barcode BC ZSM Lep 84135; paratypes dissected).

External characters and abdomen: Wingspan ♂♀ 22–24 mm. Forewing with apex pointed, slightly concave at termen subapically. Ground colour white, slightly suffused with light grey scales. Transverse lines fine, grey, rather straight. Antemedial line indistict. Medial line thin on forewing passing distal to discal spot, on hindwing proximally. Postmedial line sub-dentate, running straight to costa. Terminal area concolorous with ground colour. Terminal line and fringe dots dark grey, distinct. Discal spots present. Proboscis well developed. Length of palpi about 0.8 times diameter of eye. Frons black, vertex white, collar with slight sand-coloured tinge. Antennae of ♂ with intersegmental indentations, ciliate-setose, length of cilia about 1.2–1.4 times width of flagellum, measured at intersegmental indentations. Hindtibia of ♂ strongly dilated, with very large pencil from base of tibia, tibial scales covering tarsus. Tarsus 0.3–0.4 times length of tibia. Variation: Ground colour of Spanish populations with ochreous tinge, less whitish than the populations in Morocco. Female wing pattern and coloration not different from that of male. No polychroistic wing patterns observed in contrast to the case in I. aversata. Male genitalia: Uncus long and slender, digitiform. Gnathos modified into a pair very large, spinose lobes. Saccus projecting, triangular. Valva slender, distal third bent ventrad, at tip sub-truncate, not spinulose. Aedeagus with five cornuti, length 2.4–2.5 mm. Female genitalia: Ductus bursae very long, narrow and straight, at antrum strongly dilated, trumpet-shaped. Ductus bursae looped at transition towards corpus bursae. Corpus bursae ovoid, spinulose all over. Distribution and abundance: West Mediterranean. Recently recorded in southernmost Spain (Müller 2010; Guerrero et al. 2012). – In Europe rare and local. – Outside Europe in ­Morocco (Middle Atlas).

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Phenology: Univoltine in nature. In Spain and Morocco recorded from mid-July to late ­August (Müller 2010; Guerrero et al. 2012). In captivity with a subsequent generation emerging in September. Biology: Probably polyphagous on dry and withered leaves. Easily reared on withered leaves of Taraxacum, Polygonum and on various flowers, e.g. Campanula (B.  Müller, M. Leipnitz pers. comm.). Habitat: Mountain shrubland with Juniperus and Genista, mountainous mixed forest with Pinus and Quercus. In Spain from 1100–1700 m (Müller 2010; Guerrero et al. 2012). In Morocco, the species has been collected in a small area, in mountain habitats at 1700–1750 m above sea-level. Similar species: The (allopatric) sister species Idaea aversata (Linnaeus, 1758) differs in the pale ochreous ground colour, sometimes with orange tinge, in the postmedial line of forewing bent at costa and in the filiform flagellum of ♀ antennae. Difference in coloration is striking in the nominotypical populations from Morocco whilst Spanish populations are more reminiscent of I. aversata and require dissection. Male genitalia of I.  aversata with shorter, broader uncus, much smaller gnathos lobes, saccus less projecting, valva slightly curved rather than bent, spinulose and tapered at tip; female genitalia of I. aversata with ductus bursae broader, antrum less dilated. Genetic data (cf. Hausmann et al. 2013): BIN BOLD:AAA8256 (n=5 from Spain and ­Morocco). BIN-sharing with Idaea aversata (n=46; from eight European countries), but at a constant distance of 0.16% (1bp). Remarks: Despite the small genetic distance from I. aversata and the wing coloration of the Spanish populations approaching that of I. aversata, the large differences in genitalia clearly justify species rank for I. gelbrechti. The taxon indeviata (Prout, 1929), examined in specimens from Algeria and Tunisia, clearly belongs to I. aversata, subspecific value may be discussed. 532

Brachyglossina exilaria (Guenée, 1858), comb. n. Acidalia exilaria Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 9: 449 (southern France). Syntypes 1♂2♀ (ZFMK, examined).

Remarks: Herewith transferred from genus Idaea to Brachyglossina: Brachyglossina ­exilaria (Guenée, 1858), comb. n., because of morphological characters, larval behaviour and genetic similarity: Larvae often with transverse black and white rings (J. Gelbrecht pers. comm.; not confirmed by M. Leipnitz), which are characteristic for many species of Brachyglossina. With typical feeding behaviour of genus Brachyglossina, i.e. photophobous, on the ground, in loosely spun ‘caves’, preferring ‘dirty conditions’ like mouldy, wet food (J. ­Gelbrecht, M.  Leipnitz pers. comm.). Adult morphology: ♂ genitalia with more than one cornutus (three) and subapical, ventral process of valva, ♀ genitalia with very short and broad ductus bursae reminiscent of equivalents in representatives of genus Brachyglossina, e.g. in B. culoti. Similar structures, however, also occur in a few Idaea species. Genetically nearest species (COI barcodes): North African Brachyglossina culoti (Wehrli, 1930) (5.0%), B. tantalidis (Turati, 1924) (5.1%), Spanish B. hispanaria (Püngeler, 1913) (5.4%), and Moroccan B. truschi (6.6%) (see below; cf. Hausmann et al. 2013). For full treatment of this species with figures of adults and genitalia see Hausmann (2004).

The two North African sister species of B. exilaria have to be transferred, too: ­Brachyglossina erubescens (Hausmann, 2003), comb.n. from Tunisia and Brachyglossina truschi (Hausmann, 2003), comb.n. from Morocco. Genetically nearest species of B. truschi: B. culoti (5.2%), B. erubescens (5.3%), B. tantalidis (5.5%), Turkish B. consociata (Staudinger, 1900) (5.7%), B. hispanaria (5.8%), B. exilaria (6.6%) and B. manicaria (6.9%). Genetically nearest species of B. erubescens: B. truschi (5.3%), B. consociata (6.1%), B. culoti (6.6%), and B. tantalidis (6.7%). Further study should examine whether the gracilipennis species-group from the Near East (Idaea purpureomarginata (Bohatsch, 1880), I. gracilipennis (Warren, 1901), I. subpurpurata (Staudinger, 1900) also needs to be transferred to the genus Brachyglossina. So far, the species have not been reared, DNA barcodes suggest relationship to Brachyglossina, wing pattern and ♂ genitalia with a large number of cornuti are reminiscent, too.

Brachyglossina manicaria (Herrich-Schäffer, 1852), comb. n. Acidalia manicaria Herrich-Schäffer, 1852: Syst. Bearb. Schmett. Eur. 6 (55): 67. (Spain, Andalusia: Ronda). Holotype ♂ (MNHU, examined).

Remarks: Herewith transferred from genus Idaea to Brachyglossina for similar reasons as in the case of B. exilaria (see above): Larvae with blackish head capsule (M. Leipnitz pers. comm.; photo examined) and weak transverse black and white rings (T. Drechsel, J. ­Gelbrecht pers. comm.; not observed by M. Leipnitz), with typical feeding behaviour of genus Brachyglossina, i.e. photophobous, on the ground, in loosely spun ‘caves’, preferring ‘dirty conditions’ like mouldy, wet food (M. Leipnitz, T. Drechsel, J. Gelbrecht, R. ­Blaesius pers. comm.). Adult morphology: ♂ genitalia with a large bundle of parallel cornuti, ♀ genitalia with very short and broad ductus bursae reminiscent of equivalents in some species of Brachyglossina. Genetically nearest species (COI barcodes): Turkish Idaea proclivata (Fuchs, 1902) (6.5%), European Idaea ostrinaria (Hübner, 1813) (6.7%) and North African Brachyglossina truschi (6.9%; see above). The generic combinations of Idaea proclivata (Fuchs, 1902) and Idaea ostrinaria await further analysis, the ostrinaria species-group was already 533

considered to be closely related to Brachyglossina in Hausmann (2004). Further study should test if also Idaea nexata (Hübner, 1813) needs to be transferred to the genus Brachyglossina. So far, I. nexata has not been DNA barcoded, and no behavioural and morphological data from rearings are available currently.

Tribus Scopulini 189. Oar pratana (Fabricius, 1794) Phalaena pratana Fabricius, 1794: Ent. Syst. 3 (2): 161 (North Africa, ‘Barbaria’; neotype from Algeria: Bône). Original syntype(s) not traced in ZMUC, MNHN. Neotype, ♂, [eastern Algeria], ‘Bône, 22.V., L. Lhomme’, ZSM/Herbulot, examined.

For full treatment of this species with synonymy, description, figures of adults and genitalia see Hausmann (2004). Distribution and abundance: Eremic. Recently recorded on Lampedusa island (Fiumi & Guidi 2011). – Occurrence in Europe very local and rare. – Outside Europe widely distributed from central Algeria to Egypt (nominotypical subspecies), from southern Levant (‘subsp. mortuaria’) across Arabian peninsula to Yemen and Sudan (‘subsp. oppressa’) (Hausmann 2006a; Hausmann et al. 2016). On Canary islands and southern Morocco (Agadir region) replaced by subsp. baezi Hausmann, 2004, in Sokotra island by sister species Oar tertia Hausmann, 2009. Phenology: In Europe only one single record from late September. For extralimital records see Hausmann 2004. Habitat: Xerothermophilous, halophilous. In Europe collected at light (!) at 150 m above sea-level in an open habitat on a sun-exposed slope (Fiumi & Guidi 2011). For extralimital records see Hausmann (2004). Similar species: For differences from the West Mediterranean O. reaumuraria (Millière, 1864) see Hausmann (2004). Genetic data: Three BINs BOLD:ADM2637 (n=4 from Algeria, Tunisia including the neotype of O. pratana and short sequences of the holotypes of ab. pallens and ab. lactea, cf. Hausmann 2004); BOLD:AAI3418 (subsp. mortuaria (Staudinger, 1898); n=3 from ­Israel, northern Egypt); BOLD:AAF2861 (subsp. oppressa (Walker, 1870); n=3 from Yemen), subspecies status of the Middle East taxa requiring further integrative taxonomic study, at large genetic distances from nominotypical subspecies (up to 6%). Genetic distance between North African nominotypical subspecies and the European sibling species O. reaumuraria 9.2%. Not yet DNA barcoded from Lampedusa island, but supposedly BIN- or sequence-sharing with the Algerian and Tunisian populations.

190. Scopula drenowskii Sterneck, 1941 Scopula decorata f. drenowskii Sterneck, 1941: Z. Wien. Ent. Ver. 26 (2): 53 (Bulgaria: Alibotusch-Bereng). Though formally introduced as a ‘form’, Sterneck (1941: 54) calls it ‘a separate species, hence not understood at infrasubspecific rank. Replacement name of preoccupied Acidalia decorata var. rebeli Drenowski, 1930, being a junior primary homonym nec Prout, 1913 (identity: Scopula beckeraria). For nomenclatural availability see Hausmann

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(2004). Male genitalia slide of the type specimen from Rebel collection NHMW (Museum of Natural History of Vienna) examined. Validated at subspecific rank (Scoble 1999), downgraded to synonymy (Hausmann 2004), raised to species rank (Can 2009). ‡ Acidalia ornata var.? aequata Staudinger, 1879: Horae Soc. Ent. Ross. 14 (4): 440 (northern Turkey, Pontus: Amasia). Holotype ♂ MNHU (Museum für Naturkunde der Humboldt-­Universität, Berlin). “Preliminary considered as aberratio” by Staudinger, hence infrasubspecific and unavailable.

External characters and abdomen: Wingspan ♂♀ 25–29 mm. Hindwing slightly concave between M1 and M3. Ground colour creamy white. Antemedial and medial lines usually present but weak, the latter sometimes dentate, with costal spots distinct. Postmedial line dentate, distally with grey spots reaching costa, brownish double spots only on forewing. Near termen with small, strongly confluent spots, not reaching forewing apex. Discal spot usually grey on forewing, black on hindwing. Fringe chequered dark grey and white. Frons blackish brown, vertex whitish, collar brown. Antennae of ♂ almost filiform, ciliate-­ fasciculate, cilia 1.0–1.3 times width of flagellum. Male hindtibia stout, but not dilated, without spurs but with pencil, tarsus 0.7–0.9 times length of tibia. Male genitalia: Similar to those of S. decorata (Denis & Schiffermüller, 1775) but left arm (ceras) of sternum A8 much longer in S. drenowskii, right arm more strongly curved at tip, base of sternum more strongly projecting (cf. Can 2009). Female genitalia: Not investigated in Can (2009). Two examined (DNA barcoded) females without significant and constant differences from genitalia of the sister species S. decorata. Distribution and abundance: (South-east-) European-West Asiatic. In Europe restricted to Bulgaria (locus typicus!) and north-easternmost Greece. – Occurrence in Europe very rare and local. – Outside Europe from western to eastern Turkey.

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Phenology: Scarce data from early June to late June. Biology: Larval stages unknown. Habitat: So far, verified specimens (dissected males and/or DNA barcoded vouchers) were collected at altitudes from 1200 up to 1800 m, in montane habitats. Similar species: Coloration and wing pattern very similar to that of the sympatric sibling species S. decorata, for secure species identification dissection (males) and/or DNA barcoding is necessary. On average, slightly larger than S. decorata, in male ‘genitalia’ (terminalia) left arm (ceras) of sternum A8 much longer, right arm more strongly curved at tip, base of sternum A8 more projecting. Currently, females cannot be identified reliably without DNA barcoding. Genetic data: BIN: BOLD:AAC1708 (n=15 from Turkey). Genetically homogeneous, maximum variation 0.4%. Nearest species: S. decorata (1.6%), S. submutata (2.7%). Remarks: Upgraded from synonymy to species rank by Can (2009), based on correlated differences in male genitalia and DNA barcodes. This species deserves further research to confirm the taxonomy and to provide bionomic data. In detail it should be investigated whether further differential features can be found (e.g. different larval food-plants, different habitats) and whether the correlation between differential characters in male genitalia and DNA barcodes can be confirmed also for the distribution area in Europe.

191. Scopula donovani (Distant, 1892) Lycauges donovani Distant, 1892 (April): Annals and Magazine of Natural History (6) 10: 240. pl. 2, fig. 4 (South Africa, Gauteng (‘Transvaal’): Pretoria). Syntype(s) (NHMUK). Longula extraordinaria Staudinger, 1892 (June): Dt. ent. Z. Iris 5: 157, pl. 2, fig. 7 (Lebanon: Beirut). Syntypes 8♂4♀ (MNHU: examined). Lycauges extremata Warren, 1897: Novit. Zool. 4: 57 (Nigeria, River Niger: Akassa). Syntypes 2♀ (NHMUK: examined).

External characters and abdomen: Wingspan ♂♀ 19–26 mm, autumn generation(s) smaller, 16–21 mm. Ground colour ochreous with warm pale brown tinge. Antemedial and ­medial lines absent. Postmedial line dentate but broken into single dots, proximally bordered by vague, straight grey-brown fascia. Terminal dots sharp, proximally bordered by usually indistinct subterminal line, the latter sometimes broken into elongate streaks. Discal spots blackish, conspicuous on all wings. Abdomen ochreous, often with a central line of black dots. Frons flat, pale brown, vertex ochreous. Proboscis weak, somewhat shortened. Male palpi very short, length of palpi half diameter of eye, in female slightly longer. Male antennae with intersegmental incisions, ciliate-fasciculate, length of cilia twice width of flagellum. Female antennae filiform. Male hindtibia slender, without pencil, with two terminal spurs of unequal length, tarsus not shortened. Variation: Considerably varying in size, wing pattern, however, characteristic and constant. Male genitalia: Socii narrow, of medium length (0.3 mm). Fibula (ventral arm of valva) sclerotized, tapering. Saccus projecting, v-shaped. Aedeagus long (1.75 mm), constricted at centre, tapering at tip, with small tapered cornutus. Sternum A8 with round base and round mappa. Left ceras half length of mappa, right ceras longer than mappa (0.8 mm), both cerata with spinules at tip.

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Female genitalia: Apophyses long and narrow, apophyses anteriores as long as apophyses posteriores (1.1–1.2 mm). Lamella antevaginalis sclerotized, broad. Ductus bursae short (0.7 mm), dilated towards corpus bursae. Signum elongate, spines in fusiform position. Distribution and abundance: Afrotropical with extension to the eremic parts of North Africa and Arabian peninsula. In Europe recently recorded from southern Spain (Granada, ­Motril: Gastón et al. 2013) and then repeatedly confirmed from southernmost Spain (P. Hale, J. Garzon pers. comm.). – Occurrence in Europe rare and local. – Outside Europe widely distributed from South Africa to North Africa (Morocco: Rungs 1981; Egypt: Wiltshire 1949), and from Yemen to Israel and Lebanon (Hausmann 2006a; Hausmann et al. 2016).

Phenology: Probably bi- or polyvoltine. In Europe scarce data from mid-March to mid-April. In North Africa recorded in winter, from November to March (Rungs 1981) and in spring, from mid-April to late April (ZSM). In Israel bi- or polyvoltine with data from early June to late July, mid-September to early November, with one record as late as early December (ZSM). In Egypt flying in successive broods almost throughout the year (Wiltshire 1949). Biology: Larva recorded on rice (Oryza, Graminaceae) from Nigeria, which surely is not a natural food-plant (Scoble 1999; Robinson et al. 2010; Hausmann et al. 2016a). Habitat: Xerothermophilous. In Europe and North Africa restricted to hot and dry open habitats or scattered scrubland in lowlands from sea-level up to 200 m. In Africa inhabiting steppes and savannahs, in the eremic zone in oases. In Israel collected in hot habitats near lake-side vegetation with Phragmites and Cyperus papyrus. Similar species: No similar species in Europe.

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Genetic data: BIN BOLD:AAC9910 (n=16 from South Africa, Mauretania, Mali, Gambia, Yemen, Israel). Infraspecific variation elevated (maximum 2.5%), requiring further attention. Nearest species: Scopula yamanei Inoue, 1978 from Taiwan (5.9%). Remarks: The species was listed in the ternata species-group (subgenus Calothysanis) in Hausmann (2004).

Subfamily Larentiinae Tribus Larentiini Entephria byssata (Aurivillius, 1891), stat. n. Cidaria byssata Aurivillius, 1891: Nordens Fjärilar: 245 (central Lapponia). Lectotype ♂ (NHRS, examined, designated in Troubridge 1997). Earlier considered as junior synonym of E. punctipes (Troubridge 1997; Scoble 1999), recently raised again to species rank (Aarvik et al. 2017).

Remarks: In GME3 (Hausmann & Viidalepp 2012) this taxon was synonymized with the North American E. punctipes (Curtis, 1835), referring to the taxonomic concepts of Troubridge (1997) and Scoble (1999). The taxon was raised to species rank by Aarvik et al. (2017) which is justified by the considerable divergence of COI barcodes (5.1%) between Euro­pean and North American populations. The amplitude of the divergence exceeds that of the divergence between the two European species E. byssata and E. polata (Duponchel, 1830) (4.4%).

Tribus Trichopterygini 192. Nothocasis rosariae Scalercio, Infusino & Hausmann, 2016 Nothocasis rosariae Scalercio, Infusino & Hausmann, 2016: Zootaxa 4161 (2): 177, figs 1–4, 6, 10–12, 14, 16 (Italy, Calabria, Sila mountains: Vivaio Sbanditi (CS)). Holotype ♂ (coll. CreaSam, Italy; examined and dissected; several paratypes DNA barcoded).

External characters and abdomen: Wingspan ♂ 22–28 mm, ♀ 23–29 mm. Ground colour light grey, especially in ♂ irrorated by darker scales, wing pattern not being well contrasted. Transverse lines of forewings parallel to termen, black or dark grey, bordered by brownish scales especially along basal, medial and postmedial lines. Terminal line broken into a row of paired black dots. Discal spots black, surrounded by a pale ring. Fringe concolorous with wings, sometimes darker at vein terminations. Hindwing without transverse lines, terminal area slightly darker, discal spots black. Frons, thorax and abdomen concolorous with wings. Antennae ciliate-setose in males and scarcely ciliate in females, flagellum dorsally chequered black and white. Hindtibia of both sexes with one pair of distal spurs, male hindtibia with a pencil longer than the tibia. Variation: In Sila mountains less variable than on Mte. Pollino. Forewing ground colour varying from dark grey to light grey. Forewing pattern more variable than pattern of hindwing. Forewing sometimes with two anastomoses of medial and postmedial lines on veins CuA1 and CuA2. Sometimes (♂♀) with dark grey medial fascia, bordered by brownish antemedial and postmedial fasciae. 538

Male genitalia: Uncus long, tubular, distal portion sparsely setose, width constant from base to tip, inversely T-shaped. Valva setose, bilobed at tip. Apices of cleft valva densely setose. Costa strongly sclerotized. Harpe v-shaped distally, with two pointed tips. Juxta large, trapezoidal. Saccus short and flat. Aedeagus short, narrow at base and at tip, slightly dilated at 4/5 of its length. Vesica without spinose cornuti (Scalercio et al. 2016). Female genitalia: Antrum well sclerotized, cup-shaped, showing in mounted genitalia a width/length ratio 3.3–3.6 (n=3) and bidimensional shape with apices pointed and widely projecting laterally. Ductus bursae short, membranous. Ductus seminalis projecting from short membranous appendix bursae between ductus bursae and corpus bursae. Posterior part of corpus bursae narrowly cylindrical, weakly sclerotized, slightly granulated or with sparse microspines. Anterior part of corpus bursae elliptic, membranous, without signum (Scalercio et al. 2016). Distribution and abundance: Central Mediterranean (Trans-Ionian), montane. Mountains of southern Italy (Calabria, Basilicata), with some isolated populations from Greece (Epirus). In southern Italy sometimes abundant at the sites of occurrence. – Replaced by the allo­patric sister species N. sertata (Hübner, 1817) in the mountains of central Europe, northern Balkan peninsula (confirmed from Croatia by DNA barcoding; I. Mihoci pers. comm.), and in northern Europe, one further record confirmed as N. sertata from Tuscany through dissection. Populations from Umbria (Prola & Racheli 1979), Marche (Teobaldelli 1976; 1978; 2009), Abruzzo (Sciarretta & Zahm 2002) need further study.

Phenology: Univoltine. From late August to mid-November, single specimens emerging from mid-August. Proterandrous, ♂♂ prevailing at the beginning of the flight season, in November exclusively ♀♀ having been collected. 539

Biology: Larval stages unknown. Adults most abundant in pure beech forests, suggesting Fagus sylvatica as food-plant (Scalercio et al. 2016), while larvae of the sibling species N. sertata are monophagous on Acer pseudoplatanus (Hausmann & Viidalepp 2012). Recently collected in extremely large numbers in a forest of Acer, questioning the hypothesis of Fagus-feeding (S. Scalercio pers. comm. 2017). Habitat: Silvicolous, montane. From 500 m up to 1500 m. Inhabiting forested mountain habitats, especially forests dominated by beech (Fagus sylvatica) and various species of Acer, but found also in Calabrian black pine forests (Pinus nigra laricio). Preferring young forests with closed canopy to old, more scattered forests (Scalercio et al. 2016). Similar species: Allopatric sister species N. sertata differing in less diffuse wing pattern, m ­ edial and postmedial lines arranged in well-defined fasciae; in male genitalia uncus wider at base and inversely y-shaped, harpe bilobed; in female genitalia antrum laterally less diverging and less pointed, posterior part of corpus bursae more strongly sclerotized. Genetic data: BIN BOLD:AAM2755. Genetically homogeneous in southern Italy (maximum variation (0.6%). Distance from sister species N. sertata 2.7%. The single specimen from Epirus, Greece, BIN-sharing with N. rosariae, but diverging by 1.77% (minimum pairwise distance) from the Italian populations, by 2.85% from N. sertata, preliminarily placed with N. rosariae in Scalercio et al. (2016) but requiring further study.

Tribus Perizomini 193. Perizoma juracolaria (Wehrli, 1919) Lar[entia] alpicolaria f. juracolaria Wehrli, 1919: Mitt. Ent. Ver. l 11: 2, pl. I, II (Switzerland, Solothurner Jura: Weissenstein, inferred from labels of type specimen). Lectotype ♀ (ZFMK), designated and figured in Bryner & Ziegler (2014). Larentia reisseri Schawerda, 1932: Int. ent. Z. 26 (26): 283, figured (Corsica: Monte Rotondo). Syntype(s) (NHMW). Synonymized with P. obsoletata in Mironov (2003), raised to subspecies rank in Leraut (2009), tentatively raised to species rank in Taurand & Tautel (2019). Here accepted at subspecies rank under P. juracolaria, requiring further integrated taxonomic study, including DNA barcoding analysis. ‡ Perizoma alpicolaria: sensu Leraut, 2009 (genitalia figs) nec Herrich-Schäffer, 1848 (identity: P. obsoletata; cf. Bryner & Ziegler 2014). Misidentification.

External characters and abdomen: Wingspan ♂ 22–26 ♀ 24–28 mm. Pattern well and sharply contrasted against background colour. Forewing antemedial line at inner margin oblique. Medial area at inner margin with trapezoid or paralellogram-shaped spot. Forewing discal dot usually in area of whitish ground colour. All other characters without constant and significant differences from P. obsoletata (Herrich-Schäffer, 1838), cf. Mironov (2003). Male genitalia: Uncus short and stout, broad at base. Valvae at tip regularly rounded, costa and ventral margin parallel. All other characters without constant and significant differences from P. obsoletata (Herrich-Schäffer, 1838), cf. Mironov (2003). Female genitalia: Sclerotized part of ductus bursae comparatively long, cylindrical. Signum more or less rounded. All other characters without constant and significant differences from P. obsoletata (Herrich-Schäffer, 1838), cf. Mironov (2003).

540

Distribution and abundance: European, montane. French and Swiss Jura, southern Germany (Baden-Württemberg: Rennwald 2015, southern Bavaria: Wolf 2014; Huemer et al. 2015), Swiss and south-eastern French Alps, Liechtenstein, western Austria (Vorarlberg, North Tyrol: Huemer et al. 2015), north-easternmost Italy (isolated record from Friuli province: Huemer et al. 2015), Apennines (Lazio; Abruzzo), further data from southern Italy (Infusino et al. 2016; own, unpublished data). Studied material from Macedonia (ZSM) suggesting that all populations from the Balkan peninsula belong to P. juracolaria. For populations from Corsica see Taurand & Tautel (2019), requiring further integrated taxonomic study, for the moment accepted at subspecies rank of P. juracolaria. – Local, sometimes common. Partially sympatric with the sister species P. obsoletata (Herrich-Schäffer, 1838), e.g. in some localities in the French Alps from 1700 up to 2300 m (Berard et al. 2005), or in Liechtenstein, Vorarlberg and in Bavaria (data from ZSM; Wolf 2014), repeatedly being collected at the same sites on the same days (Huemer et al. 2015). P. obsoletata (cf. distribution indicated in Mironov 2003) seems to be absent from the Apennine peninsula while P. juracolaria seems to be absent from most parts of the eastern Alps, in the Pyrenees and the Iberian peninsula.

Phenology: Univoltine, from late June to mid-August. Larval development from August to October, overwintering as pupa (Bryner & Ziegler 2014). Biology: Larva monophagous on capsules of Gentiana, mainly G. lutea. Found in capsules of Gentiana lutea (Dietze 1874; Wehrli 1919; Berard et al. 2005). Recorded also on G. asclepiadea (Bryner & Ziegler 2014), assumed to be the larval food-plant also in Germany (Ebert 2003; Wolf 2014), however requiring confirmation according to Huemer et al. (2015). Sister species P. obsoletata found on Gentiana purpurea and G. punctata (Bryner & Ziegler 2014; Huemer et al. 2015). More exact data on food-plant preferences are required. ♀ rate high in collections, perhaps due to lower affinity of males to light sources. 541

Text-figs 216–217. Diagnostic characters (indicated) for wing-pattern-based identification of Perizoma obsoletata (Herrich-Schäffer, 1838) and P. juracolaria (Wehrli, 1919): Text-fig. 216. P. obsoletata (Austria sup., Kl. Pyhrgas, Gowilalm, e.l., 10.v.1935). Text-fig. 217. P. juracolaria (France, Hautes Alpes, Le Lautaret, 30.vi.1952).

Habitat: Open habitats, low-nutrient grasslands on slopes, with occurrence of yellow gentian. From 1000 m up to 2300 m, exceptionally down to 600 m, on average at lower elevation than the sister species P. obsoletata, the latter from 850 m up to 2550 m (Huemer et al. 2015), but usually above 2000 m, sometimes lower (Berard et al. 2005). Similar species: Sister species P. obsoletata with forewing discal spot less conspicuous because on darker background, antemedial fascia at inner margin less oblique. Less reliable differential characters: P. obsoletata on average slightly larger, medial area at inner margin with rectangular spot, hindwing discal dot and postmedial line often less distinctly marked. ♂ genitalia of P. obsoletata with longer, narrower uncus, valva broader, tip of valva asymmetrical, slightly lobed at tip towards costa or at centre. In ♀ genitalia the sclerotized part of ductus bursae shorter, usually triangular, suitability as differential feature, however, questioned by Bryner & Ziegler (2014) and Huemer et al. (2015). Signum often elongate. Genetic data (cf. Huemer et al. 2015): BIN BOLD:AAD4226 (n=10 from Germany, Austria, France, central and southern Italy). Infraspecific variation very low (0.38%). Nearest species: P. obsoletata at a distance of 7.7% (n=14 from Germany, Austria and northern Italy; BIN BOLD:AAD4225). Remarks: In Mironov (2003) treated as an infrasubspecific form of P. obsoletata (Herrich-­ Schäffer, 1838), validated and raised to species rank by Berard et al. (2005). In Leraut (2009) misidentified as ‘Perizoma alticolaria’. According to Mironov (2003) the type locality 542

of P. obsoletata is ‘Sicily’. Since the species was not refound in Sicily (cf. map in Mironov 2003) this type locality may be wrong. If the type locality ‘Sicily’ is correct, the distribution area in Italy as revealed by our study would suggest P. obsoletata to be the correct name for what is named ‘Perizoma juracolaria’ in recent literature and in this volume. Taurand & Tautel (2019) validate P. reisseri from Corsica and P. avilaria from Iberian peninsula at species rank. These questions, however, require further attention and accurate, integrated taxonomic revision including DNA barcode analysis.

194. Perizoma barrassoi Zahm, Cieslak & Hausmann, 2006 Perizoma barrassoi Zahm, Cieslak & Hausmann, 2006: Mitt. münch. ent. Ges. 95:31, figs 1a-b, 2a-b (Italy, Abruzzo: Maiella Mts., Valle Cannella, Manzini-Hut). Holotype ♂ (coll. Zahm/ZSM, examined and dissected, DNA barcoding failed). Female paratype erroneously attributed to this species, after DNA barcoding revealing to belong to genus Colostygia.

External characters and abdomen: Wingspan ♂ 19.5 mm, ♀ 23 mm, forewings slender in the holotype, apex strongly pointed. Ground colour whitish pale grey, silky shining. Transverse pattern of forewing grey with slight brown tinge, towards forewing costa darker. Medial area towards inner margin whitish, with only narrow anterior border reaching inner margin. Discal spot distinctly developed in centre of medial area. Dark terminal area dilating towards apex. Fringe chequered on forewing. Hindwing silky pale grey, darker towards termen, with dark terminal line. Underside almost without wing pattern. Palpi short, length 0.8–1.0 times diameter of eye. ♂ antennae ciliate-setose, flagellum dorsally chequered grey and white. ♂ with two pairs of hindtibial spurs. Variation: Little variation observed, but forewing less narrow and apex less pointed in the second known specimen, collected by H. Rietz, as was expected for females. Male genitalia: Valvae comparatively narrow, ventral and dorsal margins approximately parallel. Costa of valva narrow from base to apex. Anal tube strongly developed, strongly spinulose at tip. Head of (fused) labides comparatively large. Aedeagus with nine small cornuti of almost equal length (0.1–0.15 mm). Length of aedeagus 0.9 mm. Female genitalia: Apophyses tapered at apices. Colliculum elongate, with almost constant width over the whole length, slightly wider at centre. Corpus bursae small, globular. Scobinate signum small, round. Diagnosis based on genitalia of the specimen from H. Rietz; female paratype misidentified, see Remarks. Distribution and abundance: Central Mediterranean. Endemic to central Italy, strongly isolated from the closest distribution of the sister species P. incultaria (Herrich-Schäffer, 1848) in the Italian Alps. So far, only the male holotype from Majella mountains, and one female from Mt. Terminillo (prov. Lazio; leg. et coll. H. Rietz) is known. Recently generated genetic data suggesting the potential presence of P. barrassoi in the Alps as a sibling species of P. incultaria (see Genetic data). Phenology: Univoltine, two records from mid-July to late July. Biology: Larval stages unknown. Larvae of the sister species P. incultaria feeding on Primulaceae and Saxifragaceae, both being abundant at the collecting sites of P. barrassoi.

543

Habitat: Montane. Collected on karstic slopes at 2500 m in the Maiella and 1800 m on Mt. Terminillo. Collected in larger valleys at the transition zone from (gapless) low vegetation to alpine polster vegetation. Similar species: The allopatric sister species P. incultaria differing in broader forewings, broader medial area of forewing at inner margin, wing pattern more strongly developed on wing underside. ♂ genitalia of P. incultaria with broader costa of valva, dilating towards apex and ending in a small tapered process, anal tube less sclerotized and less spinulose at tip, head of labides smaller, aedeagus with larger but less cornuti (5–7), the distalmost being the largest. Female genitalia of P. incultaria very similar, signum larger, colliculum broader in the posterior part. Genetic data: The specimen from Mt. Terminillo was DNA barcoded with a Next Generation Sequencing approach, resulting in a 658bp sequence including a 89bp n-gap. BIN: BOLD:ACJ5976 (n=1 from central Italy). Nearest species: P. incultaria (8.5%; n=19 from Germany, Austria and northern Italy; BIN BOLD:AAF5044). A rare ‘haplotype of P. incultaria’ (n=5 from Germany: Bavaria and Austria: Tyrolia) BIN-sharing, at a distance of only 1.1% from P. barrassoi. The distinctness of this second Alpine cluster from P. incultaria at species level was confirmed by RAD-sequencing involving nuclear genes (Mutanen et al. 2017); we therefore suspect the presence of P. barrassoi (or a subspecies of it) in the Alps (Huemer et al. in prep.). Remarks: COI barcoding revealed misidentification of the female paratype (mentioned in the original description Zahm et al. 2006), referring to an unusually small specimen of Colostygia aquaeata. The taxonomic status of the taxon barrassoi merits further attention (e.g. analysis of DNA barcodes; larval food-plants).

544

Tribus Eupitheciini 195. Eupithecia carpophilata Staudinger, 1897 Eupithecia carpophilata Staudinger, 1897: Dt. ent. Z. Iris 10: 108, pl. 3, fig. 69 ([Russia]: Amur; Ussuri – other syntypes erroneously also mentioned from: Spain, Aragon, Sierra de Albarracin: misidentified and not conspecific). Syntypes (MNHU), lectotype designation necessary, preferably to be chosen from eastern Russia. Eupithecia carpophilata collega Dietze, 1908: Dt. ent. Z. Iris 21: 199 ([Uzbekistan]: Aksu). Syntypes 9♂2♀. Junior synonym according to Scoble (1999). Eupithecia kaszabi Vojnits, 1974: Acta zool. Hung. 20 (3–4): 445, figs 1a, 2a (Mongolia: Central Aimak, SE Somon Bajanzogt). Holotype ♂ (TMB). Junior synonym according to Mironov (1990), Viidalepp (1996) and Scoble (1999). Unavailable names: (incorrect subsequent spelling): corpophilata: Mironov & Trochimova (2007).

External characters and abdomen: Wingspan 20 mm. Ground colour homogeneously grey with slight brown tinge, transverse lines of forewing sharp, thin, black. Basal line present. Antemedial, medial and postmedial lines double, undulating. Forewing with narrow, elongate discal spot, surrounded by double medial line. Hindwing unicolorous grey brown, discal spot fine, transverse lines absent. All wings with fine, black, uninterrupted terminal line. Male genitalia: Uncus short, biapical. Valva medium length, medially broadened. Saccus rounded. Vesica armed with two cornuti, one straight and narrow, the other stouter and sigmoid-curved in distal half. Sternum A8 elongate, with two long, curved, sclerotized, apically pointed horns; basal hollow deep and basal lateral projections narrow; sternum length (1.7 mm) about 0.85 times valva length. Female genitalia: Anterior part of bursa copulatrix (perhaps homologous to the minute ante­ rior diverticulum in E. alliaria Staudinger, 1870) globular, completely covered with small spines. Posterior part of bursa copulatrix conical, covered with stout spinules, with bulbed, membranous diverticulum at ductus seminalis. Colliculum comparatively large and broad. Distribution and abundance: East Palaearctic. One specimen collected in Bashkiria (Southern Urals: Kraka, 53°46’57” N, 58°44’13” E; Mironov & Trofimova 2007). – Outside Europe across central Asia to the Far East. Putative occurrence in Spain as suggested in the original description (Staudinger 1897) based on misidentification (see above). Phenology: The only European specimen collected in early August (Mironov & Trofimova 2007). Outside Europe with data from mid-July to late August. Biology: Immature stages unknown. Habitat: Xerophilous. In the Urals collected on stony, dry, steep slopes at 600–900 m above sea-level (Mironov & Trofimova 2007). Similar species: Well characterized by the homogeneous grey ground colour (with slight brown tinge) and the thin, sharp, double transverse lines. European E. alliaria Staudinger 1870 (almost allopatric, distribution areas touching in easternmost European Russia) very similar, postmedial line of forewing more strongly (inwards) curved at costa, terminal area with slighly better expressed pale wavy line dilated to a pale spot in the forewing tornus. Genitalia of both sexes diagnostic, allowing reliable identification.

545

Genetic data: BIN: BOLD:AAY2745 (n=3 from Kyrgyzstan, north-eastern China). Genetically homogeneous. Nearest species: Eupithecia satyrata (Hübner, 1813) (6.4%), E. irrigu­ ata (Hübner, 1813) (6.5%), closest species of the venosata species-group at a distance of 8.3% (E. alliaria Staudinger, 1870) and 10.7% (E. schiefereri Bohatsch, 1893), cf. Remarks. Remarks: To be listed in the venosata species-group (numbers 42–47 in GME4: Mironov 2003), questioned by genetic dissimilarity (see above).

196. Eupithecia dubiosa Dietze, 1910 Eupithecia dodoneata f. dubiosa Dietze, 1910: Biol. Eupithecien 1: pl. 78, figs 769, 770; ibidem (1913), 2: 141 (Lebanon: Beirut). Lectotype ♀ (MNHU). Valid at subspecific rank.

External characters and abdomen: Wingspan ♂♀ 15–18 mm. Forewing comparatively short and broad. Ground colour grey with pale brownish tinge. Transverse lines more or less conspicuous, dark brownish grey; basal, postbasal and antemedial lines angled near costa; ­medial area between antemedial and medial lines darker and between medial and post­ medial lines usually lighter than rest of wing; the darker medial fascia touching discal spot (or slightly separate) and obtusely curved from it toward costa. Postmedial line inwards curved near costa, on its inner side with series of elongate, black dashes on veins. Terminal area narrow, pale wavy line present. Discal spot small, usually well developed, sometimes absent. Hindwing with ground colour slightly paler than forewing. Transverse lines represented by series of dark dashes or wedge-shaped marks, well marked along dorsum. Terminal area narrow, usually darker. Pale wavy line invisible. Discal dot minute, indistinct. Fringe chequered light and dark brownish grey. Variation: Little variation in coloration and pattern, but contrast of pattern against the background colour varying. 546

Male genitalia: Valva with evenly curved ventral margin, without medial ventral process. Aedeagus large, length 1.5–1.6 mm. Vesica armed with one very long (0.95–1.1 mm) and narrow cornutus, two short and tapered spines, and one elongate, curved sclerite. Additionally a small, irregularly shaped sclerite at the base of ductus ejaculatorius. Sternum A8 elongate, narrow, at tip asymmetrical, with two tapered spines, one straight, the other pointing downwards, curved. Female genitalia: Bursa copulatrix covered with spinules at base, with larger marginal spines, and with elongate patch of spinules in ductus bursae. Ductus bursae broad, parallel-­ bordered. Tergum A8 posteriorly rounded. Distribution and abundance: Recorded on Samos (D. Fritsch pers. comm.). – In Europe local, at the sites of occurrence usually common. – Outside Europe from southern Turkey and Cyprus to the Levant. In Turkey (prov. Antalya, Malatya) recorded by Mironov (2013), the records for Maras province in Koçak & Kemal (2009) under the name of Eupithecia dodo­ neata very likely belong to this species.

Phenology: From Europe (Samos) only scarce data from collecting of larvae (late March, early April). Adult moths in Israel bivoltine, from mid-March to late April and late September to early November. Overwintering as larva. Biology: Scarce data. Found on Pyrus spinosa (Rosaceae) and Pistacia lentiscus (Anacardia­ ceae) (Samos: Fritsch pers. comm.). Habitat: Xerothermophilous. Mediterranean macchia, scattered forest with oak and pistacia. Usually in the lowlands and lower hills up to 500 m above sea-level, on southern slopes up to 1000 m.

547

Similar species: Allopatric E. dodoneata (Guenée, 1858) hardly distinguishable in wing pattern and coloration, but strongly diverging in genitalia and DNA barcodes. Allopatric E. abbreviata Stephens, 1831 very similar, too, but larger, forewing more elongate (wingspan 20–25 mm, exceptionally 18 mm only), and diverging in genitalia and DNA barcodes. Genetic data: BIN BOLD:AAJ0162 (n=4 from Israel). Infraspecific variation very low (0.38%). Nearest species: E. abbreviata at a distance of 3.2% (n=38 from seven European countries and Tunisia). Remarks: To be listed in the interruptofasciata species-group (numbers 48–61 in GME4: Mironov 2003). The taxon has been taxonomically revised by Mironov (2013).

Eupithecia dodoneata Guenée, 1858 Eupithecia dodoneata Guenée, 1858: in Boisduval & Guenée, Hist. nat. Insectes (Spec. gén. Lépid.) 10 (2): 344, pl. 11: 7 ([England, France and Belgium]). Syntypes ♂♀ not traced. Eupithecia andrea Weisert, 2005: Z. Arb. Gem. Ost. Ent. 57: 47 (southern Spain, Andalusia: Province Cadiz, Tarifa). Holotype ♀ (NHMW). Downgraded from species rank to synonymy in Redondo et al. (2009) because of the lack of constant and significant differential features (previously stated by V. Mironov pers. comm.), here confirmed.

Remarks: The taxon andrea Weisert, 2005, described from southern Spain, needs to be synonymized with E. dodoneata (V. Mironov pers. comm.). The taxon was not mentioned in Leraut (2009), but formally downgraded in Redondo et al. (2009). For description, data and illustrations for E. dodoneata see Mironov (2003).

197. Eupithecia furcata Staudinger, 1879 Eupithecia furcata Staudinger, 1879: Horae Soc ent. ross. 14 (4): 479 (Turkey: ‘Caraman or Jenikeui high plain’, the author apparently not being sure about the locality; perhaps Amasia, see below). Holotype ‘♀’ (MNHU, examined). 1♂ labelled as ‘origin’ (=type) from ‘Amasia’ from the Staudinger collection examined, dissected by de Laever, though only ‘one female’ is mentioned in the original description, probably after erroneous determination of sex by Staudinger. The collection date on the labels is the same as indicated for the holotype in the original description: 7 June.

External characters and abdomen: Wingspan ♂ 12–14 mm, ♀ larger, 14–16 mm. Forewing narrow, forewing apex pointed. Ground colour dirty white, with grey suffusion towards forewing costa and forewing termen. Ante- and postmedial lines of forewing double, straight and oblique, bent shortly before costa. Forewing apex divided by whitish streak. Discal dots small and inconspicuous on all wings. Hindwing dirty whitish grey, with white postmedial fascia. Fringe chequered white and grey. Variation: Without remarkable variation in wing coloration and pattern. Male genitalia: Uncus narrow, tapering, biapical at tip. Valva narrow, with evenly curved ventral margin, in distal half considerably narrowing. Aedeagus short and broad, length 1.2 mm, width 0.5 mm. Vesica armed with two narrow sclerites of 0.5 mm length. Sternum A8 triangular, anteriorly broad, posteriorly narrow, tip rounded, base concave. Female genitalia: Ductus bursae with an elongate patch of minute spinules and some larger lateral spinules. Corpus bursae long oval, with oblique border of spiniferous area, almost reaching posterior border of corpus bursae but not reaching colliculum. 548

Distribution and abundance: Recently recorded in Macedonia: Pletvar (coll. H. Rietz, leg. L. Traxler – H. Rietz pers. comm.; adult photo and genitalia examined by V. Mironov pers. comm. and by the author). – In Europe local and rare. – Outside Europe from northern to southern Turkey, northern Syria, northern and central Iran (V. Mironov, U. Ratzel pers. comm.).

Phenology: Univoltine, in Europe and Turkey from late May to early July; in Iran from midApril to mid-May. Biology: Early stages and larval food-plant unknown. Habitat: Xeromontane. In Europe collected at 950 m above sea-level. Dry slopes with with sparse, dry, steppe-like vegetation. In Turkey and Iran from 800 m up to 1800 m. Similar species: Small size, whitish ground colour, oblique, double transverse lines and white apical forewing streak are unique. No similar species in Europe. Female genitalia are close to those of E. thurnerata Schütze, 1958 (H. Rietz pers. comm.) which is synonymized with E. inconspicuata Bohatsch, 1893 in Mironov (2013), see below. Wing coloration and pattern, however, completely different: E. inconspicuata with darker, more dirty ground colour, apical streak of forewing indistinct, pattern more diffuse, transverse lines less oblique etc.. Genetic data: BIN: BOLD:AAJ0145 (n=3 from Turkey). Genetically homogeneous. Nearest species: E. cooptata Dietze, 1904 (5.1%), E. lanceata (Hübner, 1825) (5.4%), E. graphata (Treitschke, 1828) (5.6%). Remarks: In GME4 (Mironov 2003), the year of the original description is erroneously dated as ‘1848’. To be listed in the graphata species-group (numbers 87–92 in GME4: Mironov 2003). 549

Eupithecia inconspicuata Bohatsch, 1893 Eupithecia inconspicuata Bohatsch, 1893: Dt. ent. Z. Iris 6: 12 (northern Turkey: Amasia). Syntypes ♂♀. Eupithecia thurnerata Schütze, 1958: Ent. Z., Frankf. a.M. 68 (5): 62, pl. 1: 21–30 (Macedonia: Stari-Dojran, Drenovo near Kavadar). Holotype ♂ (ZSM; examined), downgraded to synonymy in Mironov (2013).

Remarks: In Scoble (1999) and Mironov (2003) both taxa thurnerata Schütze, 1958 and inconspicuaria Bohatsch, 1893 were validated at species rank, in different species-groups. The synonymy was recognized by Mironov (2013). Further studies should address the question if the European populations can be validated at subspecies rank (‘thurnerata’).

198. Eupithecia gypsophilata Skou, Mironov & Rietz, 2017 Eupithecia gypsophilata Skou, Mironov & Rietz, 2017: Zootaxa 4272 (2): 291, figs 1–4, 9, 11 (Spain, Huesca: 2 km W Peñalba). Holotype ♂ (coll. P. Skou; examined). ‡ Eupithecia gemellata: sensu auct., e.g. Mironov, 2003 (partim) nec Herrich-Schäffer, 1861 (identity: E. gemellata). Misidentification.

External characters and abdomen: Wingspan ♂♀ 15–17 mm. Forewing costa almost straight. Ground colour ash-grey. Basal, medial, postbasal and antemedial lines oblique, straight, sharply angled at costa. Medial transverse band distinct, whitish, touching discal spot. Forewing apical streak pale. Terminal area with pale wavy line. Discal spot small, blackish, round. Hindwing pale ash-grey, transverse lines fine, curved, usually distinct. Basal area darker, terminal area narrow. Discal spot small, round, less distinct than on forewing. Fringe chequered pale and dark grey. Palpi with mixed dark and pale grey scales, short, equal to diameter of eye. Variation: Wavy line more or less distinct. Transverse lines on hindwing sometimes more vague. Male genitalia: Without significant and constant differences from those of E. gemellata (cf. Skou et al. 2017b and GME4: Mironov 2003). Female genitalia: Without significant and constant differences from those of E. gemellata (cf. Skou et al. 2017b and GME4: Mironov 2003). Distribution and abundance: West Mediterranean, replacing Eupithecia gemellata Herrich-­ Schäffer, 1861 in Spain. For occurrence of the latter see Mironov (2003). – Local, sometimes common at the sites of occurrence. – Outside Europe a sister species in Turkey and Lebanon, the genetic data suggesting these populations to be another undescribed species requiring accurate integrative revision. Phenology: Probably bivoltine. Recorded from late June to mid-August; early September to mid-October. Larvae found from mid-September to early November (King & Viejo Monte­ sinos 2010; Skou et al. 2017b). Biology: Larva oligophagous. Repeatedly found on Gypsophila struthium, sometimes also on Limonium dichotomum (King & Viejo Montesinos 2010; Skou et al. 2017b). Adults nocturnal, readily attracted to light. Habitat: Xerothermophilous. Steppe habitats on gypsiferous soil. From 175 m up to 1000 m (Skou et al. 2017b), exceptionally up to 1700 m (Redondo et al. 2009). 550

Similar species: From allopatric E. gemellata differing in paler ground colour, finer, oblique and straight transverse lines of forewing, a fine pale medial fascia strongly bent at costa, pale streak in the apex, and in less distinct hindwing discal spot. In male and female genitalia no clear-cut differences, but with considerable differences in their DNA barcodes. E. cooptata Dietze, 1904 (occurring in Spain but so far not recorded in the same localities with E. gypso­ philata) similar, but medial area between ante- and postmedial line wider, postmedial line at forewing costa more curved and rounded (see Remarks). Genetic data: BIN: BOLD:AAC9936. Genetically very homogeneous (maximum variation 0.0%; n=4 from southern Spain). Nearest species: E. gemellata Herrich-Schäffer, 1861 from Turkey (3.3%; potentially representing an undescribed species); E. gemellata from Italy, Croatia and Greece (7.5%). Remarks: To be listed in the graphata species-group (numbers 87–92 in GME4: Mironov 2003). Several genitalia dissections made in the ZSM indicate that all ‘E. gemellata’ records from the eastern Aragon region (e.g. Albarracin) as indicated on the distribution map in Mironov (2003) belong to E. cooptata. This applies also for figure 90d on the colour plates.

199. Eupithecia pindosata Weidlich, 2008 Eupithecia pindosata Weidlich, 2008: Entomofauna 29 (27): 481, pl. 2, fig. 2 (Greece, Pindos Mts.: Tringia, 6 km N Kranea/Aheloos). Holotype ♀ (MNHU; two paratypes examined, provided by the courtesy of M. Weidlich).

External characters and abdomen: Male unknown, so far. Wingspan ♂♀ 14.5–16 mm. Ground colour grey with slight brown tinge. Transverse and wavy lines vague. Discal spot conspicuous. Fringe slightly chequered. Hindwing paler than forewing, darker towards

551

t­ermen. Abdomen dark brown, first segment dorsally white. ♀ antennae filiform. Palpi very short, 0.6–0.7 times diameter of eye. Male genitalia: Without significant differences from those of E. conterminata. Vesica with spinulose patch and 2–3 small dentate cornuti and a paired sclerotized ridge. Sternum A8 concave anteriorly, with paired posterior projections, comparatively broad, but for variation in E. conterminata see Mironov 2003: gen.fig. 101). Female genitalia: Apophyses posteriores twice length of apophyses anteriores. Corpus bursae oval, ductus bursae short, sub-quadratic. Colliculum narrow. Lamina dentata anteriorly with small spinules and 7–8 larger spines at centre, a few spinules near posterior margin of corpus bursae. Distribution and abundance: Endemic to northern Greece, described from Pindos mountains (Weidlich 2008), later on found in four localities in a radius of approx. 20 km (Weidlich pers. comm.). Replacing the allopatric sister species E. conterminata (Lienig & Zeller, 1846), the latter having its southernmost distribution at a distance of 700 km in the Carpathian mountains. For unclear taxonomy see Genetic data and Remarks.

Phenology: Univoltine. Scarce data from mid-April to early May. Biology: Larva probably monophagous (1), suggested to feed on the needles of Abies borisii-­ regis (Weidlich 2008). Adults nocturnal, attracted to light. ♀-rate apparently high, the complete type series consisting of females (high ♀-rate is characteristic also for the sister species E. conterminata).

552

Habitat: Sylvicolous, montane. Inhabiting steep slopes and gorges with shady coniferous forest with Abies borisii-regis. All known records are near the type locality at about 1000 m a.s.l. Similar species: Allopatric sister species E. conterminata with longer palpi (length equal to dia­meter of eye), forewings paler, discal spot somewhat smaller and less distinct (cf. Weidlich 2008). In female genitalia lamina dentata larger with more large spines in the centre. Apophyses posteriores shorter. Genetic data: BIN: BOLD:AAD3722 (n=1 from Greece), the BIN, however, including also seven DNA barcodes from southern Germany and Austria (E. conterminata) at a very small distance of 0.3% (2 base pairs) from the Greek specimen, questioning the current taxo­ nomic status as a separate species. Sister species E. conterminata splitting into four, geographically separated DNA clusters (BINs), requiring further study. Remarks: To be listed in the irriguata species-group (numbers 98–101 in GME4: Mironov 2003). Current classification as a separate species doubtful, requiring further integrative study involving nuclear genes, rearings and morphological analysis with larger samples.

Eupithecia mandarinca Mironov, 2001 Eupithecia mandarinca Mironov, 2001: Nota lepid. 24 (1/2): 27, fig. 3, 15 (Ukraine: Crimea, Karadagh). Holotype ♀ (ZISP, examined by V. Mironov). ‡ Eupithecia nachadira: sensu Mironov, 2001 (partim) nec Brandt, 1941: Mitt. münch. ent. Ges. 31 (3): 877, pl. 29: 23 ([Iran]: [Belutschistan], Kouh i Taftan).

Remarks: Recorded for the fauna of Europe under the name of E. nachadira in GME4 (Mironov 2003, number 107), with the taxon Eupithecia mandarinca Mironov, 2001 listed as a junior synonym. The latter, however now again raised from synonymy and validated at species rank (V. Mironov pers. comm.), the European records (Crimea) thus referring to Eupi­thecia mandarinca. For description, data and illustrations for E. mandarinca see Mironov (2001; 2003). In GME3 (Mironov 2003), figs 107a and 107b (S. Iran) referring to E. nachadira, figs 107c and 107d (N. Iran; Crimea) to E. mandarinca (V. Mironov pers. comm.).

200. Eupithecia goossensiata Mabille, 1869 Eupithecia goossensiata Mabille, 1869: Annls Soc. ent. France (4) 9 (1): 78 ([northern? France]). Lectotype ♀ (coll. Herbulot in ZSM, examined).

External characters and abdomen: Very similar to its sister species E. absinthiata (Clerck, 1759) but smaller (wingspan ♂♀ 13–18 mm, females up to 19 mm; 17–22 mm in E. absinthiata), forewings narrower and more pointed, ground colour grey-brown, lacking the warm reddish brown tinge, forewing discal spot usually smaller. Variation: Ground colour varying from pale grey-brown to darker grey-brown but never redbrown in the centre of the forewing. Male genitalia: Without constant differences from those of E. absinthiata (see Mironov 2003). Female 2003).

genitalia:

Without constant differences from those of E. absinthiata (see Mironov

553

Distribution and abundance: European. From France across central Europe to southern Scandinavia, in the north to central Finland (cf. Opheim 1972; Knaben 1977; Weigt 2006; Mikkola et al. 1989; Silvonen et al. 2014) and Baltic countries. Not distributed south of the Alps, but one DNA barcode from central Italy, requiring confirmation. – Usually local, at the sites of occurrence sometimes not rare. – Not distributed outside Europe.

Phenology: Univoltine. In central Europe from mid-July to mid-August, exceptionally 10 days earlier or later. In Finland from mid-June to mid-August (Mikkola et al. 1989; Silvonen et al. 2014), exceptionally earlier. Larva from late July to early October (Weigt 2006). Biology: Larva oligophagous, clearly preferring Calluna. Collected from Calluna vulgaris by net-sweeping, rarely also on other plants, e.g. Solidago and Artemisia (Weigt 2006). ­According to Weigt (1991) and other authors, the larvae of E. goossensiata differ from those of E. absinthiata by their smaller size (14–16 mm; 18–20 mm in E. absinthiata) and by different colour and pattern (more rosy-reddish, with five more strongly contrasting dorsal spots). The former character was suggested to be due to starving, the latter character depending on the colour of the inflorescence of the foodplant (Mironov 2003). Weigt (2006) mentions reddish coloration also when feeding on other plants. Habitat: Hygrophilous. Inhabiting low open habitats, heathlands, peat-bogs, forest-fringes with plentiful Calluna. From sea-level up to about 200 m in the north, up to about 500 m in the south of the distribution, exceptionally up to 1000 m (Weigt 2006). Similar species: E. absinthiata (see above under ‘external characters’). Genetic data: BIN BOLD:AAA4217 (n=15 from United Kingdom, Finland, Germany, Italy). Intraspecific variation comparatively low, 0.6%. Constant genetic distance from the sister species E. absinthiata 1.25% in European populations, the latter with 65 DNA barcodes 554

(>500bp) from eight European countries. Both species with separate BINs. The other species of the European absinthiata species-(sub-)group, E. expallidata Doubleday, 1856, at a genetic distance of 3.5%. Remarks: Species rank and species delimitations have repeatedly been discussed in literature (e.g. Dietze 1913; Schütze 1954; Vojnits 1969a; Fibiger 1979; Kaaber 1980; Mikkola 1982; Skou 1986; Weigt 1991; Mikkola et al. 1989; Silvonen et al. 2014). Downgraded to synonymy of E. absinthiata in Mironov (2003) stressing the lack of differences in genitalia and assuming that the small size of goossensiata is hunger-induced due to feeding on a sub-­optimal food-plant (Calluna). Upgraded from synonymy to species rank again in Hausmann et al. (2011b) because of constant differences in the DNA barcode, correlated with differential features in habitus. To be listed in the satyrata species-group (numbers 117–125 in GME4, Mironov 2003). In Mironov (2003), the taxon Eupithecia absinthiata var. callunae Speyer, 1867 is listed as a valid subspecies of E. absinthiata. Since Calluna heaths (name-giving to ‘callunae’) are typical habitats and food-sources of E. goossensiata, it should be checked if this taxon may be a senior synonym of E. goossensiata. The exact locus typicus of the type specimen of goossensiata (northern France?) is not clear which may question the correct attribution of the type specimen.

Text-figs 218–219. Diagnostic characters (indicated) for wing-pattern-based identification of Eupi­thecia absinthiata (Clerck, 1759) and E. goossensiata Mabille, 1869: Text-fig. 218. E. absinthiata (France, Aisne, Puisieux-et-Clanlieu, 25.VII.1969). Text-fig. 219. E. goossensiata (Ger­ many, Umg. Kassel, Wilhelmshausen, e.o., 2.VII.1950).

555

Eupithecia millefoliata f. wettsteini Vojnits, 1974 Eupithecia wettsteini Vojnits, 1974: Folia Ent. Hung. XXVII (2): 235 (Hungary: Balatonszemes). Holotype ♀ (cf. Fazekas 2017). ­ azekas Unavailable names (incorrect subsequent spellings): wettsetini: Fazekas (2017); wettsteni: F (2017).

Remarks: Fazekas (2017) examined the holotype and correctly concluded that this taxon is just an individual form of E. millefoliata. However, in his article in Hungarian language he consistently uses the name as a binomen “Eupithecia wettsteini” (partly misspelled, see above), which may be misleading and suggesting species rank.

Subfamily Ennominae Tribus Ourapterygini 201. Ourapteryx niveiscythes Zolotuhin, 2017 Ourapteryx niveiscythes Zolotuhin, 2017: Fauna Lepid. Volgo-Uraliensis: 531, pl. 63, figs 11–15 (Russia, lower Volga region: Astrachan Reserve, Obzhorovo). Holotype ♂ (ZISP).

External characters and abdomen: Wingspan 45–52 mm, in Caucasus sometimes up to 57 mm. Forewing apex falcate, especially in females. Hindwing shortly tailed at M3, base of tail with two brown dots. Ground colour white, appearing semi-transparent, without yellowish tinge. Towards termen irrorated with silvery striae. Transverse lines grey, in collections sometimes fading to brownish. Postmedial line almost straight, on hindwing curved. Forewing discal spot elongate. See also description in Zolotuhin (2017). Male genitalia: Uncus short, wide at base, apex strongly sclerotized, pointed, hook-shaped. Gnathos at centre dentate, rather broad. Valva broad, parallel-sided. ‘Furca arms’ (homo­ logy unclear) strongly curved at upper third forming a hook or a loop, apically pointed and densely setose, without wrinkles on its dorsal surface. Aedeagus stout, caecum short. Medial part of vesica long, apex with small lateral diverticulum, with patch of long cornuti (cf. Zolotuhin 2017). Female genitalia: Very similar to those of O. sambucaria (Linnaeus, 1758), and O.  per­sica (Ménétriés, 1832), not diagnostic (Zolotuhin 2017). Corpus bursae with comparatively smooth surface. Distribution and abundance: The species was collected in southernmost European Russia and Caucasus (Zolotuhin 2017), one specimen from north-westernmost Iran probably belongs to this species (see under Genetic data). Phenology, biology, habitat: Uni- or bivoltine, from late May to early June and in early August in flooded valley forest, rocky forest, bushy slopes from lowlands up to 1300 m in the Caucasus. Similar species: Differing from O. sambucaria in the whitish ground colour and the straight forewing termen, often with dark fringe. In male genitalia the ‘furca arm’ (homology unclear) is strongly curved, whilst being slightly curved in O. sambucaria. Zolotuhin (2017) states the following diagnostic features for O. niveiscythes (from all congeners in Europe 556

and Caucasian range): “smaller size, falcate apex of forewing, pure white, slightly hyaline ground color with grey – not brownish – pattern and especially strongly curved furca in male genitalia”. Genetic data: BIN: BOLD:AAE3936, based on one barcoded specimen from north-westernmost Iran, in habitus well matching the type series of O. niveiscythes and genetically clearly diverging from the other known species. Nearest species: O. persica Ménétriés, 1858 from Iran (2.7%), European O. sambucaria (3.5%) and O. malatyensis Wehrli, 1936 from Turkey (3.5%).

Tribus Gnophini 202. Dyscia govii Fiumi & Flamigni, 2013 Dyscia govii Fiumi & Flamigni, 2013: Quad. Studi Nat. Romagna 38: 2, figs. 1, 3a, 4a, 5ab, 6bd ([France]: Corsica, Omessa (Haute-Corse), Francardo). Holotype ♂ (Museo Civico di Storia Naturale ‚G. Doria‘ di Genova, Italy).

External characters and abdomen (cf. Fiumi & Flamigni 2013; Flamigni et al. 2016): Wingspan ♂♀ 33–38 mm. Discal spots larger than in D. innocentaria, postmedial line broken into a row of very conspicuous black dots. See colour plates in this volume and the figured Corsican male in Skou & Sihvonen (2015: fig. 128f). Male genitalia (cf. Fiumi & Flamigni 2013; Flamigni et al. 2016): Morphological differences are quite small, in male genitalia concerning shape of saccus (inward projection shallower in D. govii) and aedeagus (more strongly curved in the centre in D. govii).

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Female genitalia (cf. Fiumi & Flamigni 2013; Flamigni et al. 2016): Ductus bursae somewhat larger than in D. innocentaria. Distribution: Tyrrhenian. Endemic to Corsica and Sardinia. – Very isolated occurrences, sometimes not rare at its localities.

Phenology: Bivoltine. From mid-May to early June, and from early September to early ­October. In Corsica at higher elevation also in August. One specimen collected as early as late March (Flamigni et al. 2016). Habitat: Xerothermophilous. Sun-exposed, warm and dry slopes with (not too dense) Mediterranean vegetation. Vertical distribution from 200 up to approx. 500 m in Sardinia, up to approx. 1050 m in Corsica (P. Skou own data; Flamigni et al. 2016). Biology: Larva found on Teucrium marum (Lamiaceae) (Flamigni et al. 2016). Reared to the third instar on Artemisia alba (Asteraceae), possibly not being an ideal food-plant (M. ­Leipnitz pers. comm.). Genetic data: BIN: BOLD:ACM9547 (n=2 from Corsica and Sardinia). Nearest species: D. innocentaria (2.7%) and D. royaria (2.0%). Remarks: Dyscia govii was described in 2013 from the islands of Corsica and Sardinia and downgraded to subspecies rank (‘Dyscia innocentaria ssp. govii’) in GME5 (Skou & Sihvonen 2015). Flamigni et al. (2016) validate it again at species level. Despite the small differences in morphology and habitus, the correlated genetic distances (2.0–2.7% from D. innocenta­ ria and D. royaria) seem to support and corroborate the species status for this taxon.

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Index to scientific names of the Systematic Account, Vol. 6 The index gives page references to the main treatment. Numbers in semibold refer to the species numbers and figures. abruptaria – 230, 70 abstersaria – 420 achyra – 302, 98 Adactylotis – 448 adriatica – 335 Aethalura – 457 Afriberina – 386 Agriopis – 315 albarinata – 75 albarracina – 528 albosignata – 345 Alcis – 436 alexandrinus – 304 algeriensis – 345 algiricaria – 372, 128 almasa – 222 almensis – 366 alpicolaria – 272 alpina – 290, 93 alpinaria – 509 alpinata – 129, 35 alpmaritima – 147, 43 alticolaria – 148, 44 amani – 403 ambiguata – 103, 26 ambustaria – 408, 141 Amorphogynia – 275 amygdalaria – 366, 125 andalusiaria – 357, 121 andereggaria – 192, 59 andrea – 548 androgynus – 63 ankeraria – 310, 101 annegreteae – 242, 76 Apocheima – 259 Apochima – 53 aragona – 372 aragonensis – 70 aragonis – 432, 150 arenaria – 273 arenaria – 337, 114 argillacearia – 67 Arichanna – 469 Ascotis – 460 assoi – 88, 20

Athroolopha – 341 atlantica – 288 atlanticaria – 341 atlanticaria – 345 atlanticaria – 360 atlanticaria – 375, 129 atomaria – 272, 86 aurantiaria – 320, 106 avilarius – 70 bajaria – 318, 105 baldensis – 154, 46 barbuti – 526, 185 barrassoi – 543, 194 bastelbergeri – 439 bastelicaria – 396, 135 Bellachima – 53 bellieri – 68 belzebuth – 144, 41 benestriata – 122 bentelii – 138, 39 berenicidaria – 234, 72 bernieri – 472 bervaensis – 310 beschkovi – 314, 103 betularia – 303, 99 Biston – 297 Bistonini – 254 bistortata – 466 Boarmiini – 253 bowesi – 463 Brachyglossina – 533 britannica – 288 brunnearia – 400, 137 budashkini – 322, 107 budensis – 122 Bupalini – 254 Bupalus – 471 bureschi – 136 buxicolaria – 420 byssata – 538 bytinskii – 479, 484 caelibaria – 183, 56 Calamodes – 326 calceata – 67 caliginearia – 335, 113

canaliculata – 132, 36 carniolica – 290 carpathica – 142 carpophilata – 545, 195 Cassymini – 40 castiliaria – 509 cataloniae – 277 catharia – 370 cebennaria – 509 certhiatus – 125 chalybaeus – 149 Charissa – 66, 67 Chemerina – 334 Chondrosoma – 56 chrysitaria – 343, 117 cinctaria – 463, 163 cineraria – 481 Cleora – 462 Cleorini – 253 Cleorodes – 244 cleui – 321 cleui – 490 Cnestrognophos – 82 codetaria – 486 coggii – 318 cognataria – 303 Colotoini – 52 confertata – 75 consonaria – 330, 111 constantina – 521, 183 contaminaria – 450, 158 Contropis – 474 coracina – 135, 38 correptaria – 430, 149 corsica – 91, 21 corsicaria – 417 Costignophos – 70 cottei – 259 crenulata – 70, 12 crepuscularia – 466, 164 cretacaria – 235 Crocota – 212 cuencaensis – 363 culminaria – 513, 179 cyrnea – 493, 172

559

dagestanensis – 38 dalmata – 230 dalmatina – 367, 126 danieli – 266, 84 Dasypteroma – 251 davidi – 529 declinans – 312, 102 defloraria – 417 defoliaria – 307, 100 Deileptenia – 444 deversata – 439, 153 dianaria – 460 Dicrognophos – 41 difficilis – 77, 15 dilucescens – 400 dilucidaria – 196, 60 dinicola – 240 diniensis – 276 diniensis – 490 dioszeghyi – 122 dioszeghyi – 135 distinctaria – 244 dodoneata – 548 dognini – 206 dolomitica – 169, 53 donovani – 536, 191 drenowskii – 534, 190 dresnayi – 237 druentiaria – 428 dubiosa – 237 dubiosa – 546, 196 dumetata – 30, 1 Dyscia – 557 Dysgnophos – 77 ebusaria – 427, 147 Ecleora – 383 Ectropis – 465 Ekboarmia – 374 Elophos – 181 Ematurga – 272 Entephria – 538 Erannini – 254 Erannis – 307 erebaria – 410, 142 erubescens – 533 espagnolus – 472 espaniola – 504, 176 estachyi – 215 estrellae – 357 Euchrognophos – 87 Eumannia – 476, 505 Eupithecia – 545

560

Eurranthis – 345 exilaria – 533 extensaria – 407 Fagivorina – 336 fallentaria – 428 faroulti – 36 fascinataria – 378 faucium – 148 fidelensis – 233 fiduciaria – 56, 8 fingalata – 481 flabellaria – 54, 7 florentina – 295, 95 fortunaria – 397, 136 francothuringiaca – 122 fumosarius – 303 furcata – 548, 197 furvata – 61, 9 galliberia – 420 gedrensis – 148 gelbrechti – 531, 188 gerini – 432 gerini – 486 gesticularia – 448, 157 gigantaria – 372 Glacies – 127 glaucinaria – 116, 31 Gnophini – 58 Gnophos – 61 goossensiata – 553, 200 govii – 557, 20 gracilaria – 514, 180 gracilis – 375 graecaria – 125, 34 graecarius – 293, 94 granataria – 246, 78 grisearia – 269 grisolaria – 474, 167 grupchei – 279 gulsensis – 123 gypsophilata – 550, 198 hammamrirha – 388 hanoviensis – 276 Harrisonodes – 275 harterti – 237, 73 herbuloti – 103 herbuloti – 405 herrerai – 380 hibernica – 31 hirtaria – 276, 87 hispanica – 360, 123 hispidaria – 259, 82

hoefneri – 189 holli – 375 hubertcleui – 509 humiliata – 529 Hypomecis – 265 Hyposcotis – 66 ichnusaria – 417 Idaea- 526 idrisarius – 102 iliaria – 272 ilicaria – 434, 151 impectinata – 75 incerta – 244 incisaria – 528 inconspicuata – 550 innuptaria – 166, 51 insignata – 524 intermedia – 122 inversarius – 279 isabellae – 283, 90 isoscelata – 272 istriana – 276 istrianus – 293 italica – 295 italohelveticus – 73, 13 Ithysia – 516 jacobiaria – 434 japygiaria – 232, 71 jaspidaria – 244 jubata – 442, 154 juracolaria – 540, 193 kabylaria – 341 Kemtrognophos – 102 kerstinae – 269 khenchelae – 424 kindermannaria – 339, 115 kovacsi – 75 Kresnaia – 313 Kuchleria – 524 kusdasi – 142 lafauryata – 68 lanaria – 257, 81 lappona – 135 lapponaria – 281, 89 Larerannis – 332 laricaria – 466 lariciaria – 320 latimargo – 341 lepraria – 497, 173 Leucobrephos – 519 leucophaearia – 316, 104 lhommaria – 489, 171

lichenaria – 244, 77 lobaria – 529, 187 lutescentaria – 63 Lycia – 275 Macariini – 49 macguffini – 80 maghrebica – 434 magyarica – 269 malatyaca – 366 mandarinca – 553 manicaria – 533 margaritata – 35, 2 marginaria – 324, 108 marmararia – 54 marsicaria – 63 maura – 230 mauretania – 405 Megalycinia – 248 Megametopon – 474 melaleucaria – 490 melanaria – 328, 110 melanaria – 470, 165 melloulica – 480, 484 menadiara – 524, 184 mendicaria – 201 Menophra – 228 meridionalis – 298 Microbiston – 257 middendorfii – 520, 182 millefoliata – 556 mimeuri – 424 miniaria – 382, 131 minorasiaticus – 125 minuta – 272 modestaria – 474 mongolicum – 520 montserrata – 420 mucidaria – 99, 25 muraria – 436 mutilata – 83, 17 Myinodes – 521 myrtillata – 63 nachadira – 553 necessaria – 279, 88 necopinatus – 189 Neognophina – 115 nigrolineata – 480, 484 nivea – 63 niveata – 220, 67 niveiscythes – 556, 201 noricana – 142, 40 notarioi – 359, 122

Nothocasis – 538 nubilata – 75 nuragica – 485, 169 Nychiodes – 347 nycthemeraria – 240, 75 Oar – 534 obfuscata – 63, 10 obscuraria – 352, 119 obscuraria – 506 obscurata – 67, 11 occidentaria – 428 occitanaria – 327, 109 Odontognophos – 29 odrinarius -293 oliveirata – 405 omalensis – 422, 145 onustaria – 109, 28 operaria – 189, 58 ophthalmicata – 103 oppositaria – 506, 177 oranaria – 508, 178 orientalis – 341 orientaria – 272 orientis – 260 Ortaliella – 49 orthogrammaria – 333, 112 osthelderi – 363 Ourapteryx – 556 pallidaria – 400 pallidata – 324 Paraboarmia – 446 Paradarisa – 330 Parectropis – 455 parenzani – 400, 408 Parietaria – 195 peletieraria – 217, 65 peloponnesiaria – 84, 18 pennigeraria – 341, 116 pentheri – 86, 19 Peribatodes – 412 Perizoma – 540 perlinii – 138 perspersata – 49, 6 perumbraria – 425 perversaria – 420, 144 petrina – 86 pfeifferi – 119, 32 Phaselia – 369 Phaseliini – 254 Phigalia – 262 Phthonandria – 225 Phyllometra – 476, 512

pilosaria – 263, 83 pindosata – 551, 199 piniaria – 472, 166 plumaria – 403, 138 plummistaria – 345, 118 pomonaria – 285, 91 popovi – 260 postdentaria – 410 powelli – 252 powelli – 433 praerecta – 480, 484 pratana – 534, 189 pravata – 517, 181 predotae – 107, 27 pretiosaria – 526 propinquaria – 222 Prosopolophini – 52 Pseudocoremia – 451 pseudonoricana – 135 pseudotinctaria – 219, 66 Psodos – 127 psyloritaria – 500, 174 pullaria – 67 pullata – 75, 14 pullularia – 103 punctinalis – 269, 85 punctinervis – 335 punctipes – 538 punctulata – 458, 161 pusztae – 277 pyrenaea – 157 pyrenaearia – 405, 139 pyrenaica – 146, 42 pyrenaica – 70 quadrifaria – 157, 48 ragusaria – 354, 120 reisseri – 540 relicta – 372 remmi – 112, 29 repandata – 436, 152 retyezatensis – 138 rhomboidaria – 417, 143 Rhopalognophos – 115 ribeata – 444, 155 roboraria – 266, 84 rosariae – 538, 192 rosenbaueri – 300, 97 rossica – 288 rothschildi – 98 saerdabensis – 417 saerdabica – 363 sagnesi – 380, 130

561

salemae – 391, 134 Sardocyrnia – 395 sartata – 41, 4 scandinaviaria – 400 schwingenschussi – 132 Sciadia – 159 Scopula – 534 scotica – 281 secundaria – 428, 148 selenaria – 460, 162 Selidosema – 399 sepiaria – 481, 168 septaria – 165, 50 serotinaria – 206, 63 serotinoides – 207 serraria – 109 serraria – 249, 79 serrularia – 370, 127 sheljuzhkoi – 103 shurensis – 370 siciliana – 44, 5 sicula – 487, 170 similaria – 455, 160 slovenica – 168, 52 sociaria – 222, 68 sodorensium – 436 solieraria – 384, 132 Spartopteryx – 339 spitzi – 156, 47 sproengertsi – 199, 61 spurcaria – 183 staudingeri – 92, 22 strataria – 298, 96 strictaria – 226, 69

562

strobioni – 284 suavis – 453, 159 subflavaria – 420 sublutearia – 417 submelanaria – 327 subscudularia – 328 subtaurica – 94, 23 sudetica – 157 supinaria – 122, 33 Synopsia – 222 syriaca – 310 syriaca – 363 syriacaria – 400 syrirana – 424 syritaurica – 417 taeniolaria – 407, 140 tagana – 474 taurusica – 161 teberdensis – 78 tenebraria – 161, 49 tenietaria – 388, 133 Tephronia – 476, 479 teriolensis – 307 teriolica – 481 terraria – 390 textaria – 528, 186 thaumasia – 252, 80 theophilaria – 501, 175 thuriferaria – 239, 74 thurnerata – 550 tinctaria – 214, 64 tonnara – 493 torrevinagrensis – 361 transcaspia – 363

transiens – 135 Trilobignophos – 125 truschi – 533 trypanaria – 234 tundranoides – 135 turfosaria – 80, 16 tyronensis – 400 umbraria – 424, 146 undulosa – 237 ungemachi – 448 unicoloraria – 177, 55 unitaria – 222 uralaria – 276 urmensis – 78 variegata – 96, 24 vaucheri – 424 viertlii – 446, 156 vintejouxi – 215 vittaria – 201, 62 waltheri – 363, 124 wehrlii – 151, 45 werneri – 134, 37 wettsteini – 556 wockearia – 161 Yezognophos – 194 zacharia – 38, 3 zehrounensis – 318 zeitunaria – 114, 30 zelleraria – 172, 54 Zernyia – 246 zetterstedtaria – 273 zirbitzensis – 187, 57 zonaria – 288, 92

The Geometrid Moths of Europe

Volume 6 Part 2

Bernd Müller, Sven Erlacher, Axel Hausmann, ­Hossein Rajaei, Pasi Sihvonen & Peder Skou

The Geometrid Moths of Europe Axel Hausmann, Pasi Sihvonen, Hossein Rajaei & Peder Skou (eds) Volume 6 Part 2

Bernd Müller, Sven Erlacher, Axel Hausmann, Hossein Rajaei, Pasi Sihvonen & Peder Skou Colour plates, Genitalia plates, Data of microscopic preparations, Checklist to Vols 1–6, References

Subfamily Ennominae II (Boarmiini, Gnophini, additions to previous volumes)

leiden

| boston

Editors: Axel Hausmann, Zoologische Staatssammlung München, Germany; Pasi Sihvonen, Finnish Museum of Natural History, Helsinki, Finland; Hossein Rajaei, State Museum of Natural History Stuttgar, Germany; Peder Skou, Ollerup, Denmark. Linguistic editor: Martin Corley, Faringdon, United Kingdom. Colour plates photographed and compiled by: Igor Kostjuk, Kiev, Ukraine. This publication should be cited as: Müller, B., Erlacher, S., Hausmann, A., Rajaei, H., Sihvonen, P. & Skou, P. (2019): Ennominae II. – In A. Hausmann, P. Sihvonen, H. Rajaei & P. Skou (eds): The Geometrid Moths of Europe 6: 1–906. Brill, Leiden.

Date of publication: 15.05.2019

Authors’ addresses: Bernd Müller, Weissdornallee 13, D-13158 Berlin, Germany, [email protected] Sven Erlacher, Museum of Natural History, Moritzstr. 20, D-09111 Chemnitz, Germany, [email protected] Axel Hausmann, SNSB – Bavarian State Museum of Zoology, Münchhausenstr. 21, D-81247 Munich, Germany, [email protected] Hossein Rajaei, State Museum of Natural History Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany, [email protected] Pasi Sihvonen, Finnish Museum of Natural History, Zoology Unit, P.O. Box 17, FI-00014 University of Helsinki, Finland, [email protected] Peder Skou, Aamosen 1, Ollerup, DK-5762, Vester Skerninge, Denmark, [email protected]

Front cover: Erannis defoliaria (Clerck, 1759). Denmark: Fyn, Aamosen, 31 October 2018. Photo by Peder Skou, Denmark. Numerous photos of differential external features shown in Text-figs modified from Igor Kostjuk’s plate photographs. The work of Axel Hausmann was supported by grants of the Bavarian ministry of science (project BFB) and by the German ministry of Science (BMBF; project GBOL). The genetic analyses have received considerable support from Paul D. N. Hebert and his competent teams at the Centre for Biodiversity Genomics (CBG) and the Canadian Centre for DNA Barcoding (CCDB University of Guelph). The data management & analysis system BOLD was provided by Sujeevan Ratnasingham. The work was financially supported by Genome Canada (­Ontario Genomics Institute) in the framework of the iBOL I and II programs. Library of Congress Control Number: 2019938865 ISBN: 978-90-04-25222-6 (hardback set) ISBN: 978-90-04-40783-1 (part 1) ISBN: 978-90-04-40784-8 (part 2) ISBN: 978-90-04-38748-5 (e-book) Copyright 2019 by Koninklijke Brill NV, Leiden, The Netherlands. Koninklijke Brill NV incorporates the imprints Brill, Brill Hes & De Graaf, Brill Nijhoff, Brill Rodopi, Brill Sense, Hotei Publishing, mentis Verlag, Verlag Ferdinand Schöningh and Wilhelm Fink Verlag. All rights reserved. No part of this publication may be reproduced, translated, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior written permission from the publisher. Authorization to photocopy items for internal or personal use is granted by Koninklijke Brill NV provided that the appropriate fees are paid directly to The Copyright Clearance Center, 222 Rosewood Drive, Suite 910, Danvers, MA 01923, USA. Fees are subject to change. This book is printed on acid-free paper and produced in a sustainable manner.

Colour plates

PLATE 1 (all specimens natural size) 1a. Odontognophos dumetata Tr., ♂, Pálava-Kotel, Moravia, Czech Republic, 25.viii.1997 (coll. Skou) 1b. O. dumetata Tr., ♂, Castel di Ieri, Abruzzo, Italy, 2.x.2010 (coll. Skou) 1c. O. dumetata Tr., ♂, Severinovka, Odesskaya obl., Ukraine, 11.ix.2016 (ZMKU) 1d. O. dumetata Tr., ♂, Gurb by Vic, Prov. Barcelona, Spain, 11.ix.2002 (coll. Skou) 1e. O. dumetata Tr., ♂, Oslje, Dubrovnik-Neretva Distr., Croatia, 5.x.2015 (coll. Skou), dissected 1f. O. dumetata Tr., ♀, Gornja Brela, Split-Dalmatia Distr., Croatia, 4.x.2015 (coll. Skou), dissected 2a. Odontognophos margaritata Zerny, ♂, Albarracin, Prov. Teruel, Spain, 18.ix.2002 (coll. Skou), dissected 2b. O. margaritata Zerny, ♂, Tramacastilla, Prov. Teruel, Spain, 27.viii.2001 (coll. Skou) 2c. O. margaritata Zerny, ♂, Albarracin, Prov. Teruel, Spain, 22.ix.1926 (coll. Skou) 2d. O. margaritata Zerny, ♀, Albarracin, Prov. Teruel, Spain, 30.ix.1926 (coll. Skou) 3a. Odontognophos zacharia Stgr., ♂, Parnassos, Fthiódia, 6.ix.2008 (coll. Skou) 3b. O. zacharia Stgr., ♂, Ideon Andron, Crete, Greece, 28.ix.2006 (coll. Nupponen) 3c. O. zacharia Stgr., ♂, Ismit, Taurus, Prov. Antalya, Turkey, 8.x.1999 (coll. Stadie), dissected 3d. O. zacharia Stgr., ♀, Ismit, Taurus, Prov. Antalya, Turkey, e. o. 8.x.1999 (coll. Stadie), dissected 4a. Dicrognophos sartata Tr., ♂, Agia Paraskevi, Prov. Ioánina, Greece, 26.viii.2008 (coll. Skou) 4b. D. sartata Tr., ♀, Kapesovo, Epirus, Greece, 21.vi.2013 (coll. Skou) 4c. D. sartata Tr., ♀, Kalymvari, Crete, Greece, 1.–3.x.2016 (coll. Skou) 4d. D. sartata Tr., ♂, Pulo di Altamura, Puglia, Italy, 29.v.2016 (coll. Skou) 4e. D. sartata Tr., ♀, Leros, Dodokanisa, Greece, 2.–4.v.2004 (coll. Skou) 4f. D. sartata Tr., ♀, Arachova, Central Greece, Greece, 9.vi.2013 (coll. Skou), dissected 4g. D. sartata Tr., ♀, underside, Arachova, Central Greece, Greece, 9.vi.2013 (coll. Skou), dissected 5a. Dicrognophos siciliana Skou & Sihv., ♂, Rifugio Oristano, Piano Zucchi, Sicily, Italy, 5.vi.2016 (coll. 5b. 5c. 5d. 6a. 6b. 6c. 6d.

568

Skou), paratype, dissected D. siciliana Skou & Sihv., ♂, Santo Stefano Quisquina, Sicily, Italy, 2.vi.2016 (coll. Skou), holotype, dissected D. siciliana Skou & Sihv., ♀, Rifugio Oristano, Piano Zucchi, Sicily, Italy, 17.ix.2014 (coll. Skou), paratype, dissected D. siciliana Skou & Sihv., ♀, underside, Rifugio Oristano, Piano Zucchi, Sicily, Italy, 2.vi.2016 (coll. Skou), paratype, dissected Ortaliella perspersata Tr., ♂, El Pinar, Bedar, Prov. Almeria, Spain, 19.–27.iv.2001 (coll. Skou) O. perspersata Tr., ♀, Güejar-Sierra, Sierra Nevada, Prov. Granada, Spain, 2.vii.2015 (coll. Skou) O. perspersata Tr., ♀, Albarracin, Prov. Teruel, Spain, 1.–15.vii.1926 (ZMKU) O. perspersata Tr., ♀, Albarracin, Prov. Teruel, Spain, 15.vii.1995 (coll. Skou)

569

PLATE 2 (all specimens natural size) 7a. Apochima flabellaria Heeger, ♂, Siv Kladenetz v., Ivailovgrad region, Bulgaria, 30.xi.1996 (coll. Skou) 7b. A. flabellaria Heeger, ♂, Mount Sakar, Dositeevo, Bulgaria, 20.iii.1994 (coll. Skou) 7c. A. flabellaria Heeger, ♀, near Petrich, Struma Valley, Bulgaria, 6.iii.2002 (coll. Skou) 8a. Chondrosoma fiducaria Anker, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 6.x.1976 (coll. 8b. 8c. 8d. 8e. 8f. 9a. 9b. 9c. 9d. 9e. 9f. 10a. 10b. 10c. 10d. 10e. 10f. 10g. 11a. 11b. 11c. 11d. 11e. 11f. 11g. 11h. 11i.

570

B. Müller) C. fiducaria Anker, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 2.x.1976 (coll. B. Müller) C. fiducaria Anker, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 28.x.1978 (coll. Skou) C. fiducaria Anker, ♂, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 31.x.1978 (coll. Skou) C. fiducaria Anker, ♀, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 6.x.1976 (coll. B. Müller) C. fiducaria Anker, ♀, ♂, Zitzmannsdorfer Wiesen, Burgenland, Austria, e. o. 6.xi.1975 (MNHU) Gnophos furvata Den. & Schiff., ♂, Kranichfeld, Thüringen, Germany, 14.vii.1979 (coll. Erlacher), dissected G. furvata Den. & Schiff., ♀, Buchfarter Höhlen, Thüringen, Germany, 15.viii.1993 (coll. Erlacher), barcoded, dissected G. furvata Den. & Schiff., ♂, Waidbruck, South Tyrol, Italy, 25.vii.1914 (MNHU) G. furvata Den. & Schiff., ♀, Visso, Monte Sibellini, Abruzzo, Italy, 19.vii.1979 (coll. Skou) G. furvata Den. & Schiff., ♀, Kapesovo, Epirus, Greece, 30.vi.2016 (coll. Skou) G. furvata Den. & Schiff., ♀, Wien, Austria (ZMKU) Gnophos obfuscata Den. & Schiff., ♂, Rifugio Mandrone, Adamello, Italy, 8.vii.1994 (coll. Erlacher), barcoded, dissected G. obfuscata Den. & Schiff., ♂, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 18.viii.1988 (coll. Skou) G. obfuscata Den. & Schiff., ♂, Campo Imperatore, Abruzzo, Italy, 31.vii.2011 (coll. Skou) G. obfuscata Den. & Schiff., ♂, Ulvö, Småland, Sweden, 5.vii.1977 (coll. Skou) G. obfuscata Den. & Schiff., ♀, Col de la Cayolle, Alpes Maritimes, France, 6.viii.1980 (coll. Skou) G. obfuscata Den. & Schiff., ♀, Schnalstal, Trentino-Alto Adige, Italy, 18.viii.1996 (coll. Skou) G. obfuscata Den. & Schiff., ♀, Rifugio Dibona, Veneto, Italy, 31.vii.2013 (coll. Skou) Charissa obscurata Den. & Schiff., ♂, Leutratal, Jena, Thüringen, Germany, 21.viii.1993 (coll. ­Friedrich) C. obscurata Den. & Schiff., ♀, Boxberg, Oberlausitz, Germany, 23.viii.2008 (coll. Friedrich) C. obscurata Den. & Schiff., ♀, Village Thérmes, Prov. Xanthi, Greece, 2.ix.2008 (coll. Skou) C. obscurata Den. & Schiff., ♀, Horas Fakion, Imbros, Crete, Greece, 2.x.2011 (coll. Friedrich), dissected C. obscurata Den. & Schiff., ♂, Omalos, Crete, Greece, 28.vii.–2.viii.2001 (coll. Skou) C. obscurata Den. & Schiff., ♂, underside, Leutratal, Jena, Thüringen, Germany, 21.viii.1993 (coll. ­Friedrich) C. obscurata bellieri Obth., ♀, Road to Haut Asco, Corsica, France, 13.–14.viii.2012 (coll. Skou) C. obscurata bellieri Obth., ♂, Corsica (ZFMK), lectotype C. obscurata bellieri Obth., ♂, underside Corsica (ZFMK), lectotype

571

PLATE 3 (all specimens natural size) 12a. Charissa crenulata Stgr., ♂, Ruta del Veleta, Sierra Nevada, Prov. Granada, Spain, 23.vii.2003 (coll.

Skou), barcoded 12b. C. crenulata aragonensis Zerny, ♀, Moscardon, Prov. Teruel, Spain, e. o. 13.viii.1998 (coll. Gelbrecht) 12c. C. crenulata aragonensis Zerny, ♀, Moscardon, Prov. Teruel, Spain, e. o. 13.viii.1998 (coll. Gelbrecht) 12d. C. crenulata aragonensis Zerny, ♀, underside, Moscardon, Prov. Teruel, Spain, e. o. 13.viii.1998 (coll. ­ elbrecht) G 12e. C. crenulata aragonensis Zerny, ♀, Albarracin, Prov. Teruel, Spain, 22.viii.2001 (coll. Skou), barcoded 12f. C. crenulata aragonensis Zerny, ♀, Albarracin, Prov. Teruel, Spain, 1.viii.2003 (coll. Skou) 12g. C. crenulata avilarius Reiss., ♂, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 20.–21.vii.2003 (coll. Skou), barcoded 12h. C. crenulata avilarius Reiss., ♀, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 20.–21.vii.2003 (coll. Skou), barcoded 12i. C. crenulata avilarius Reiss., ♀, underside, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 20.–21. vii.2003 (coll. Skou), barcoded 12j. C. crenulata pyrenaica Obth., ♀, Solsona, Prov. Lerida, Spain, 27.vii.1991 (coll. Skou), barcoded 12k. C. crenulata pyrenaica Obth., ♀, Vernet-les-Bains, Pyrénées-Orientales, France (ZFMK) 12l. C. crenulata pyrenaica Obth., ♀, underside, Vernet-les-Bains, Pyrénées-Orientales, France (ZFMK) 13a. Charissa italohelveticus Reser, ♂, Raron, Valais, Switzerland, e. o. 8.iv.1987 (coll. Gelbrecht), barcoded 13b. C. italohelveticus Reser, ♂, Raron, Valais, Switzerland, e. o. 30.iii.1987 e. o. (coll. Skou) 13c. C. italohelveticus Reser, ♀, Raron, Valais, Switzerland, e. o. 17.vii.1987 (ZSM) 13d. C. italohelveticus Reser, ♀, underside, Raron, Valais, Switzerland, e. o. 17.vii.1987 (ZSM) 14a. Charissa pullata Den. & Schiff., ♂, Staumauer Bleiloch, Thüringen, Germany, 26.vi.2006 (coll. ­Erlacher), barcoded 14b. C. pullata Den. & Schiff., ♂, Blankenberg, Thüringen, Germany, 19.vi.1993 (coll. Gelbrecht), barcoded 14c. C. pullata Den. & Schiff., ♂, Parfi, Rila Mountains, Bulgaria, 23.vii.1980 (coll. Gelbrecht), barcoded 14d. C. pullata Den. & Schiff., ♂, Bad Urach, Baden-Württemberg, Germany, e. o. 13.vi.–1.vii.1998 (coll. ­ elbrecht), barcoded G 14e. C. pullata Den. & Schiff., ♀, Fließ am Inn, North Tyrol, Austria, 1.–2.vii.1991 (coll. Skou) 14f. C. pullata Den. & Schiff., ♂, Isola 2000, Alpes Maritimes, France, 1.viii.2005 (coll. Skou) 14g. C. pullata Den. & Schiff., ♀, Col de la Cayolle, Alpes Maritimes, France, 29.–30.vii. 2005 (coll. Skou) 14h. C. pullata Den. & Schiff., ♂, underside, Fließ am Inn, North Tyrol, Austria, 25.vi.2008 (coll. Skou) 15a. Charissa difficilis Alph., ♂, Donsko, Orenburg Region, South Ural, Russia, 1.–4.vi.1998 (coll. Skou), barcoded, dissected 15b. C. difficilis Alph., ♂, Bokaydin-tau, Zailisky Alatau, Prov. Almaty, Kazakhstan, 13.–15.vi.1994 (coll. ­Sommerer), dissected 15c. C. difficilis Alph., ♂, Mount Atzhaylai, Akterek, Prov. Almaty, Kazakhstan, 25.v.1994 (coll. Sommerer), dissected 15d. C. difficilis Alph., ♂, underside, Bokaydin-tau, Zailisky Alatau, Prov. Almaty, Kazakhstan, 13.–15.vi.1994 (coll. Sommerer), dissected 16a. Charissa turfosaria Whli., ♂, Aktasch, Republic Altai, Russia, 6.–15.vi.2010 (coll. Schacht), dissected 16b. C. turfosaria Whli., ♂, Aktasch, Republic Altai, Russia, 16.vi.2012 (coll. Schacht), dissected 16c. C. turfosaria Whli., ♀, Golets Sokhondo, Sokhondinskiy Nat. Res., Transbaikal, Russia, 10.vii.1997, (ZMKU) 16d. C. turfosaria Whli., ♂, underside, Aktasch, Republic Altai, Russia, 16.vi.2012 (coll. Schacht), dissected

572

573

PLATE 4 (all specimens natural size) 17a. Charissa mutilata Stgr., ♂, Oviddağı Geçidi, Doğu Karadeniz Mts., Prov. Trabzon, Turkey, 30.v.2000,

(coll. Gelbrecht), dissected 17b. C. mutilata Stgr., ♀, Köprüköy, Mount Korga, Prov. Ispir, Turkey, 28.vii.2001 (coll. Gelbrecht), barcoded, dissected 17c. C. mutilata Stgr., ♂, Sivas, Prov. Sivas, Turkey, 22.iv.2001 (coll. Erlacher), barcoded, dissected 17d. C. mutilata Stgr., ♂, Ispir, Korga Dağı, Prov. Erzurum, Turkey, 23.vii.2001 (coll. Gelbrecht), barcoded, dissected 17e. C. mutilata Stgr., ♀, underside, Köprüköy, Mount Korga, Prov. Ispir, Turkey, 28.vii.2001 (coll. Gelbrecht), barcoded, dissected 18a. Charissa peloponnesiaria Erl. & Erl., ♂, Kosmas, Parnon Mts., Peloponnesus, Greece, 11.09.2004 (coll. Friedrich), holotype, dissected, barcoded 18b. C. peloponnesiaria Erl. & Erl., ♀, Mount Chelmos, Peloponnesus, Greece, 20.ix.1991 (ZMUC), dissected, barcoded 18c. C. peloponnesiaria Erl. & Erl., ♂, Mount Aenos, Cephalonia, Greece, 26.viii.1996 (ZSM), barcoded, dissected 18d. C. peloponnesiaria Erl. & Erl., ♂, Kosmas, Parnon Mts., Peloponnesus, Greece, 11.09.2004 (coll. ­Friedrich), dissected 18e. C. peloponnesiaria Erl. & Erl., ♂, underside, Kosmas, Parnon Mts., Peloponnesus, Greece, 11.09.2004 (coll. Friedrich), holotype, barcoded, dissected 18f. C. peloponnesiaria Erl. & Erl., ♀, underside, Mount Chelmos, Peloponnesus, Greece, 20.ix.1991 (ZMUC), barcoded, dissected 19a. Charissa pentheri Rebel, ♂, Petrina, Ochrid, Macedonia, 7.–16.vi.1935 (ZSM), barcoded, dissected 19b. C. pentheri Rebel, ♀, Petrina, Ochrid, Macedonia, 6.vi.1935 (ZSM), dissected 19c. C. pentheri Rebel, ♂, Zabljak, Durmitor Nat. Park., Montenegro, 24.vi.2013 (coll. Skou) 19d. C. pentheri Rebel, ♂, Petrina, Ochrid, Macedonia, 7.vi.1935 (ZFMK) 19e. C. pentheri Rebel, ♂, underside, Petrina, Ochrid, Macedonia, 7.–16.vi.1935 (ZSM), barcoded, dissected 20a. Charissa assoi Red. & Gast., ♂, Road to Veleta, Sierra Nevada, Prov. Granada, Spain, 8.vii.1971 (ZSM), dissected 20b. C. assoi Red. & Gast., ♀, Albatera, Sierra de Crevillente, Prov. Alicante, Spain, 24.v.2004 (TLMF), dissected 20c. C. assoi Red. & Gast., ♂, Sedella, Prov. Malaga, Spain, 18.ix.2009 (coll. Skou) 20d. C. assoi Red. & Gast., ♂, Tramacastilla, Prov. Teruel, Spain, 27.viii.2001 (coll. Skou) 20e. C. assoi Red. & Gast., ♂, Totana, Prov. Murcia, Spain, e. o. 4.vi.2003 (coll. Gelbrecht), dissected 20f. C. assoi Red. & Gast., ♂, underside, Totana, Prov. Murcia, Spain, e. o. 4.vi.2003 (coll. Gelbrecht), dissected 21a. Charissa corsica Obth., ♂, San Pantaleo, Sardinia, Italy, e. o. 4.vi.2007 (coll. Gelbrecht), barcoded 21b. C. corsica Obth., ♂, Fluminimaggiore, Sardinia, Italy, 7.–8.iv.2002 (coll. Skou) 21c. C. corsica Obth., ♂, Road to Haut Asco, Corsica, France, 13.–14.viii.2012 (coll. Skou) 21d. C. corsica Obth., ♀, San Pantaleo, Sardinia, Italy, e. o. 4.vi.2007 (coll. Gelbrecht), barcoded 21e. C. corsica Obth., ♂, Ovette Malu, Orgosolo, Sardinia, Italy, e. o. 2.–16.vi.2002 (coll. Gelbrecht), dissected 21f. C. corsica Obth., ♂, underside, Ovette Malu, Orgosolo, Sardinia, Italy, e. o. 2.–16.vi.2002 (coll. Gelbrecht), dissected 22a. Charissa staudingeri Wnuk., ♂, Mochos, Crete, Greece, 11.v.1996 (coll. Friedrich), dissected 22b. C. staudingeri Wnuk., ♂, Kamára, Samos, Greece, e. l. 9.v.2008 (coll. Erlacher), barcoded 22c. C. staudingeri Wnuk., ♀, Kamára, Samos, Greece, e. l. 9.v.2008 (coll. Erlacher), barcoded 22d. C. staudingeri Wnuk., ♂, Stalida, Crete, Greece, e. o. 18.iv.1999 (coll. Gelbrecht) 22e. C. staudingeri Wnuk., ♂, underside, Stalida, Crete, Greece, e. o. 18.iv.1999 (coll. Gelbrecht) 23a. Charissa subtaurica Whli., ♂, Alanya, Taurus Mountains, Prov. Antalya, Turkey, 4.x.2002 (coll. ­Friedrich), dissected 23b. Charissa subtaurica Whli., ♂, Mount Troodos, Paphos, Cyprus, e. o. 2.iv.2000 (coll. Gelbrecht), barcoded 23c. Charissa subtaurica Whli., ♂, Aspros Valley, Paphos, Cyprus, 27.iii.2001 (coll. Friedrich), dissected 23d. Charissa subtaurica Whli., ♀, Mavrokolympos Valley, Paphos, Cyprus, 25.iii.2001 (coll. Friedrich), dissected 23e. Charissa subtaurica Whli., ♀, underside, Mavrokolympos Valley, Paphos, Cyprus, 25.iii.2001 (coll. ­Friedrich), dissected 24a. Charissa variegata Dup., ♀, Ahtopol, Black Sea Coast, Bulgaria, e. o. 28.vi.1985 (coll. Gelbrecht) 24b. C. variegata Dup., ♂, Gignod, Aosta, Italy, 14.vii. 1990 (coll. Gelbrecht), barcoded 24c. C. variegata Dup., ♂, Dimitsana, Peloponnesus, Greece, 13.ix.2004 (coll. Friedrich), barcoded, dissected 24d. C. variegata Dup., ♂, Monodendri, Epirus, Greece, 24.v.2002 (coll. Friedrich), barcoded 24e. C. variegata Dup., ♂, underside, Monodendri, Epirus, Greece, 24.v.2002 (coll. Friedrich), barcoded 24f. C. variegata Dup., ♀, Ahtopol, Black Sea Coast, Bulgaria, e. o. 28.vi.1985 (coll. Gelbrecht), barcoded 24g. C. variegata Dup., ♂, Varna, Pobiti Kameni, Bulgaria, e. o. 29.vi.1989 (coll. Gelbrecht), barcoded 24h. C. variegata Dup., ♂, Kalavrita, Peloponnesus, Greece, 24.ix.1997 (coll. Friedrich), barcoded 24i. C. variegata Dup., ♀, Varna, Pobiti Kameni, Bulgaria, e. o. 29.vi.1989 (coll. Gelbrecht) 24j. C. variegata Dup., ♀, underside, Varna, Pobiti Kameni, Bulgaria, e. o. 29.vi.1989 (coll. Gelbrecht) 25a. Charissa mucidaria Hbn., ♂, Axarquia, Prov. Malaga, Spain, 8.iv.2002 (coll. Friedrich) 25b. C. mucidaria Hbn., ♂, Sant Rafel, Ibiza, Spain 30.iii.–3.iv.2016 (coll. Skou) 25c. C. mucidaria Hbn., ♂, Almunecar, Prov. Granada, Spain, 6.–12.v.2000 (coll. Skou) 25d. C. mucidaria Hbn., ♀, Cabo de Gata, Prov. Almeria, Spain, e.o. ii.1992 (coll. Gelbrecht), dissected 25e. C. mucidaria Hbn., ♀, underside, Cabo de Gata, Prov. Almeria, Spain, e.o. ii.1992 (coll. Gelbrecht), dissected

574

575

PLATE 5 (all specimens natural size) 26a. Charissa ambiguata Dup., ♀, Col des Champs, Colmar, Alpes Hautes Provence, France, e. o. 2.viii.2004 (coll. Gelbrecht), barcoded, dissected 26b. C. ambiguata Dup., ♂, North Umhausen, North Tyrol, Austria, 23.vi.1994 (coll. Erlacher), barcoded, dissected 26c. C. ambiguata Dup., ♂, Kleinbreitenbach, Thüringen, Germany, 20.vi.1983 (coll. Erlacher), dissected 26d. C. ambiguata Dup., ♀, underside, Col des Champs, Colmar, Alpes Hautes Provence, France, e. o. 2.viii.2004 (coll. Gelbrecht), barcoded, dissected 26e. C. ambiguata sheljuzhkoi Schaw., ♂, Russia, Aksakovo, Gub. Ufa, 20.vi.1915 (ZMKU), syntype 27a. Charissa predotae Schaw., ♂, Bronchales, Prov. Teruel, Spain, 29.vi.1962 (ZSM), dissected 27b. C. predotae Schaw., ♀, Miraflores, Prov. Madrid, Spain, e. o. 2.–3.vi.1996 (coll. Gelbrecht) 27c. C. predotae Schaw., ♀, Bayarcál, Prov. Almeria, Spain, 19.v.2007 (coll. Gelbrecht), dissected 27d. C. predotae Schaw., ♂, Bayarcál, Prov. Almeria, Spain, 19.v.2007 (coll. Gelbrecht), barcoded, dissected 27e. C. predotae Schaw., ♀, underside, Miraflores, Prov. Madrid, Spain, e. o. 2.–3.vi.1996 (coll. Gelbrecht), dissected 28a. Charissa onustaria H.-S., ♀, Liljanovo, Pirin Mountains, Bulgaria, 11.v.1982 (coll. Gelbrecht), barcoded 28b. C. onustaria H.-S., ♂, Liljanovo, Pirin Mountains, Bulgaria, e. o. 15.ix.1982 (coll. B. Müller), dissected 28c. C. onustaria H.-S., ♀, Liljanovo, Pirin Mountains, Bulgaria, e. o. 25.viii.1982 (coll. B. Müller), dissected 28d. C. onustaria H.-S., ♀, underside, Liljanovo, Pirin Mountains, Bulgaria, e. o. 25.viii.1982 (coll. B. Müller) 28e. C. onustaria serraria Gn., ♂, Vizzavona, Corsica, France, 11.vii.1951 (ZSM), 29a. Charissa remmi Viid., ♂, Ust-Nukzka, Tynda Distr., Amur Region, Russia, 22.–24.vi.1996 (coll. Skou), barcoded 29b. C. remmi Viid., ♂, Kulu, Magadan Distr., Russia, 11.vii.1980 (coll. Skou), paratype, barcoded, dissected 29c. C. remmi Viid., ♂, Irkuta, Sajan Mountains, Burjatia, Russia, 16.vi.–13.vii.2013 (coll. R. Müller), dissected 29d. C. remmi Viid., ♀, by river Zeja, Tukuringra Mountains, Amur region, Russia, 6.vi.1914 (ZMKU) 29e. C. remmi Viid., ♂, underside, Irkuta, Sajan Mountains, Burjatia, Russia, 16.vi.–13.vii.2013 (coll. R. Müller), dissected 30a. Charissa zeitunaria Stgr., ♂, Ispir, Doğu Karadeniz, Prov. Erzurum, Turkey 4.viii.2001 (coll. ­Gelbrecht), dissected 30b. C. zeitunaria Stgr., ♂, Baklatepe, Prov. Van, Turkey, 29.vii.2001 (coll. Gelbrecht), dissected 30c. C. zeitunaria Stgr., ♂, Erzurum, Palandöken, Turkey, 30.viii.1997 (coll. Löbel), dissected 30d. C. zeitunaria Stgr., ♂, Aktepe, Cappadokia, Prov. Nevsehir, Turkey, 14.–17.viii.2009 (coll. Fiebig), dissected 30e. C. zeitunaria Stgr., ♂, underside, Baklatepe, Prov. Van, Turkey, 29.vii.2001 (coll. Gelbrecht), dissected 31a. Charissa glaucinaria Hbn., ♂, Rhemes-Notre-Dame, Aosta, Italy, 12.–14.vii.1997 (coll. Gelbrecht), barcoded 31b. C. glaucinaria Hbn., ♂, Rhemes-Notre-Dame, Aosta, Italy, 12.–14.vii.1997 (coll. Gelbrecht), barcoded 31c. C. glaucinaria Hbn., ♀, Col de Champs, Alpes de Hautes Provence, France, 9.vii.1997 (coll. Gelbrecht), barcoded 31d. C. glaucinaria Hbn., ♀, Leonessa, Monte Terminillo, Lazio, Italy, 24.vi.2014 (coll. Skou) 31e. C. glaucinaria Hbn., ♂, underside, Villar d’Arene, La Grave, Dept. Hautes Alpes, France, 26.vii.2000 (coll. Skou) 32a. Charissa pfeifferi Whli., ♂, Saklikent, Bei Dağları, Antalya, Turkey, 21.v.1999 (coll. Gelbrecht), dissected 32b. C. pfeifferi Whli., ♂, Arachova, Parnassos, Greece, 1.–2.v.1998 (coll. Gelbrecht), barcoded 32c. C. pfeifferi Whli., ♂, Arachova, Parnassos, Greece, 1.–2.v.1998 (coll. Gelbrecht), barcoded 32d. C. pfeifferi Whli., ♀, Sertavu Pass, Taurus Mountains, Turkey, e. o. 15.vi.1996 (coll. Gelbrecht), barcoded 32e. C. pfeifferi Whli., ♂, underside, Saklikent, Bei Dağları, Antalya, Turkey, 21.v.1999 (coll. Gelbrecht), dissected 33a. Charissa supinaria Mann, ♂, Croatia (ZFMK), syntype 33b. C. supinaria Mann, ♂, underside, Croatia (ZFMK), syntype 33c. C. supinaria Mann, ♀, Popina Laka, Liljanova, Pirin Mountains, Bulgaria, e. o. 11.ix.1981 (coll. B. Müller), barcoded 33d. C. supinaria Mann, ♀, Koberfels near Burgk, Thüringen, Germany, e. o. 5.v.1993 (coll. Gelbrecht) 33e. C. supinaria Mann, ♂, underside, Zirl, North Tyrol, Austria, 26.vi.1961 (ZSM) 34a. Charissa graecaria Stgr., ♂, Kirilowa Poljana, Rila Mountains, Bulgaria, e. o. 14.iv.1982 (coll. G ­ elbrecht), dissected 34b. C. graecaria Stgr., ♂, Chelmos, Peloponnesus, Greece, e. o. 16.–22.vii.1994 (coll. Gelbrecht), dissected 34c. C. graecaria Stgr., ♀, Chelmos, Peloponnesus, Greece, e. o. 16.–22.vii.1994 (coll. Gelbrecht), dissected 34d. C. graecaria Stgr., ♀, Kirilowa Poljana, Rila Mountains, Bulgaria, e. o. 15.xi.1981 (coll. Gelbrecht), dissected 34e. C. graecaria Stgr., ♀, underside, Kirilowa Poljana, Rila Mountains, Bulgaria, e. o. 15.xi.1981 (coll. Gelbrecht), dissected

576

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PLATE 6 (all specimens natural size) 35a. 35b. 35c. 35d. 35e. 35f. 36a. 36b. 36c. 36d. 36e. 36f. 37a. 37b. 37c. 38a. 38b. 38c. 38d. 38e. 38f. 38g. 38h. 38i. 39a. 39b. 39c. 39d. 39e. 39f. 40a. 40b. 40c. 40d. 40e. 40f. 41a. 41b. 41c. 41d. 41e. 41f. 42a. 42b. 42c. 43a. 43b. 43c. 44a. 44b. 44c. 44d. 44e. 44f. 44g. 44h. 45a. 45b. 45c. 45d.

578

Psodos alpinata Scop., ♂, Vennatal, North Tyrol, Austria, 28.vi.1947 (ZSM) P. alpinata Scop., ♂, Innsbruck, North Tyrol, Austria, 24.vii.1994 (coll. Erlacher), barcoded, dissected P. alpinata Scop., ♂, Hochvogel, Hinterhornbach, Tyrol, Austria, 1.vii.1994 (coll. Erlacher), barcoded, dissected P. alpinata Scop., ♂, Hochtor, Großglockner, Kärnten, Austria, 23.vii.1985 (coll. Erlacher) P. alpinata Scop., ♀, Dümlerhütte, Warscheneck, Oberösterreich, Austria, 23.vii.1928 (ZSM) P. alpinata Scop., ♀, underside, Vennatal, North Tyrol, Austria, 28.vi.1947 (ZSM) Psodos canaliculata Hochw., ♂, Hoher Burgstall, North Tyrol, Austria, 16.vii.1941 (ZSM) P. canaliculata Hochw., ♂, Hochtor, Großglockner, Kärnten, Austria, 19.vii.1988 (coll. Erlacher), dissected P. canaliculata Hochw., ♂, Tierser Alp, Dolomites, Veneto, South Tyrol, Italy, 21.–26.vii.1991 (coll. Friedrich), barcoded P. canaliculata Hochw., ♀, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, late vii.1967 (coll. TLMF) P. canaliculata Hochw., ♀, underside, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, late vii.1967 (coll. TLMF) P. canaliculata schwingenschussi Whli., ♂, Tatra Mountains, Poland, 15.vii.1927 (ZSM) Psodos werneri Schaw., ♂, Bosnia-Herzegovina, Balkan: Volujak, (NHMW), paralectotype, dissected P. werneri Schaw., ♂, Bosnia-Herzegovina, Balkan: Volujak, (NHMW), lectotype, dissected P. werneri Schaw., ♂, underside, (NHMW),Bosnia-Herzegovina, Balkan: Volujak, (NHMW), paralectotype, dissected Psodos coracina Esp., ♂, Valparolo Pass, Dolomites, Veneto, South Tyrol, Italy, 13.vii.1994 (coll. E ­ rlacher), dissected P. coracina Esp., ♂, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, early viii.1967 (ZSM) P. coracina Esp., ♀, Valparolo Pass, Dolomites, Veneto, South Tyrol, Italy, 13.vii.1994 (coll. Erlacher), dissected P. coracina Esp., ♀, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, late vii.1962 (ZSM) P. coracina Esp., ♀, underside, Valparolo Pass, Dolomites, Veneto, South Tyrol, Italy, 13.vii.1994 (coll. E ­ rlacher), dissected P. coracina bureschi Varga, ♂, Viehren, Pirin Mountains, Bulgaria 17.–24.vii.1972 (ZSM), paratype P. coracina dioszeghyi Schmidt, ♂, Romania, Retyezat, Aragyes-teto, 18.–19.vii.1914 (NHMW), lectotype, dissected P. coracina lappona Whli., ♂, Enontekiö, Finland, 27.vi.1936 (ZSM) P. coracina pseudonoricana Whli., ♂, Triglav, Slovenia, 25.vii.1906 (ZSM) Psodos bentelii Rätzer, ♂, Zermatt, Valais, Switzerland, 7.viii.1912 (ZSM) P. bentelii Rätzer, ♂, Breiul-Cervinia, Matterhorn, Aosta, Italy, 15.vi.1967 (coll. Gelbrecht), barcoded, dissected P. bentelii perlini Trti., ♂, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, early viii.1967 (ZSM) P. bentelii perlini Trti., ♂, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, 7.–10.vii.1964 (coll. TLMF) P. bentelii perlini Trti., ♂, underside, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, 7.–10.vii.1964 (coll. TLMF) P. bentelii retyezatensis Bartha, ♂, Retyezot Mountains, Zenoga, Transylvania, Romania, 28.vii.1932 (ZSM) Psodos noricana Wagner, ♂, Obergurgel, Ötztaler Alpen, North Tyrol, Austria, 17.vii.1951 (ZSM) P. noricana Wagner, ♂, Nordkette, Innsbruck, North Tyrol, Austria, 24.vii.1994 (coll. Erlacher), dissected P. noricana Wagner, ♂, Bovec, Monte Kanin, Julian Alps, Slovenia, 27.vi.2003 (coll. Friedrich), barcoded, dissected P. noricana Wagner, ♀, Obergurgel, Ötztaler Alpen, North Tyrol, Austria, 17.vii.1951 (ZSM) P. noricana Wagner, ♂, underside, Nordkette, Innsbruck, North Tyrol, Austria, 24.vii.1994 (coll. Erlacher), dissected P. noricana kusdasi Whli., ♂, Furtschaglhaus, Zillertal Alps, North Tyrol, Austria, 30.vii.1927 (ZSM) Psodos belzebuth Praviel, ♂, Mont Demaut, Alpes-Maritimes, France, 9.viii.1936 (ZSM), paratype P. belzebuth Praviel, ♂, Colmars, Alpes-de-Haute-Provence, France, 29.vi.1924 (ZSM) P. belzebuth Praviel, ♂, Col de la Cayolle, Alpes Maritimes, France, vii.1953 (ZSM) P. belzebuth Praviel, ♂, Colmars, Alpes-de-Haute-Provence, France, 29.vi.1924 (ZSM) P. belzebuth Praviel, ♀, Cima Fauniera, Caráglio, Piemonte, Italy, 28.vii.2009 (coll. Skou) P. belzebuth Praviel, ♂, underside, Colmars, Alpes-de-Haute-Provence, France, 29.vi.1924 (ZSM) Psodos pyrenaica Schaw., ♂, Formiguères, Pyrénées-Orientales, France, vii.1922 (ZSM) P. pyrenaica Schaw., ♂, Formiguères, Pyrénées-Orientales, France, vii.1922 (ZSM) P. pyrenaica Schaw., ♂, underside, Formiguères, Pyrénées-Orientales, France, vii.1922 (ZSM) Psodos alpmaritima Schaw., ♂, Tête de la Roubine, Isola 2000, Alpes-Maritimes, France, 6.vii.1997 (coll. Tautel) P. alpmaritima Schaw., ♂, Mont Gelas, Alpes-Martimes, France, 31.vii.1935 (MNHN), dissected P. alpmaritima Schaw., ♂, underside, Mont Gelas, Alpes-Martimes, France, 31.vii.1935 (MNHN), dissected Psodos alticolaria Mann, ♂, Gaisbergferner, Obergurgl, North Tyrol, Austria, 2.viii.2002 (coll. ­Friedrich), barcoded P. alticolaria Mann, ♂, Rotmoostal, Obergurgl, North Tyrol, Austria, 1.viii.1994 (coll. Erlacher) P. alticolaria Mann, ♂, underside, Rotmoostal, Obergurgl, North Tyrol, Austria, 1.viii.1994 (coll. Erlacher) P. alticolaria chalybaeus Zerny, ♂, Stilfserjoch, South Tyrol, Italy, 21.vii.1896 (NHMW), lectotype, dissected P. alticolaria chalybaeus Zerny, ♂, Muttekopf, North Tyrol, Austria, 25.viii.1953 (coll. TLMF) P. alticolaria chalybaeus Zerny, ♂, underside, Stilfserjoch, South Tyrol, Italy, 21.vii.1896 (NHMW), lectotype, dissected P. alticolaria faucium Favre, ♂, Mettelhorn, Zermatt, Valais, Switzerland, 20.vii.2003 (coll. Erlacher), dissected, barcoded P. alticolaria gedrensis Rondou, ♀, Gèdre, Hautes-Pyrénées, France, vii.1908 (ZSM) Psodos wehrlii Vorbr., ♂, Mettelhorn, Valais, Switzerland, 11.vii.1921 (ZSM) P. wehrlii Vorbr., ♂, Täsch Alp, Valais, Switzerland, 3.viii.1987 (coll. TLMF), barcoded P. wehrlii Vorbr., ♀, Täsch Alp, Valais, Switzerland, 13.viii.1988 (coll. TLMF), barcoded P. wehrlii Vorbr., ♂, underside, Täsch Alp, Valais, Switzerland, 3.viii.1987 (coll. TLMF), barcoded

579

PLATE 7 (all specimens natural size) 46a. Psodos baldensis Wolfsb., ♂, Cima Valdritta, Monte Baldo, Verona, Italy, mid vii.1969 (ZSM) 46b. P. baldensis Wolfsb., ♂, Monte Baldo, Verona, Italy, mid vii.1966 (TLMF), paratype 46c. P. baldensis Wolfsb., ♀, Monte Baldo Verona, Italy, mid vii.1966 (TLMF), paratype 46d. P. baldensis Wolfsb., ♀, Cima Valdritta, Monte Baldo, Verona, Italy, mid vii.1969 (ZSM) 46e. P. baldensis Wolfsb., ♂, underside, Cima Valdritta, Monte Baldo, Verona, Italy, mid vii.1969 (ZSM) 46f. P. baldensis Wolfsb., ♀, underside, Cima Valdritta, Monte Baldo, Verona, Italy, mid vii.1969 (ZSM) 47a. Psodos spitzi Rbl., ♂, Triglav, Slovenia, 3.viii.1921 (coll. TLMF), cotype 47b. P. spitzi Rbl., ♂, Bovec, Mount Kanin, Slovenia, 2.viii.2001 (coll. Erlacher), barcoded, dissected 47c. P. spitzi Rbl., ♂, Triglav, Julian Alps, Slovenia, late vii.1934 (ZSM) 47d. P. spitzi Rbl., ♀, Mount Kanin, Slovenia, 4,viii.1995 (TLMF) 47e. P. spitzi Rbl., ♂, underside, Triglav, Slovenia, 3.viii.1921 (coll. TLMF), cotype 47f. P. spitzi Rbl., ♀, underside, Mount Kanin, Slovenia, 4.viii.1995 (TLMF) 48a. Psodos quadrifaria Sulzer, ♂, Nordkette, Innsbruck, North Tyrol, Austria, 24.vii.1994 (coll. Erlacher),

barcoded, dissected 48b. P. quadrifaria Sulzer, ♂, underside, Nordkette, Innsbruck, North Tyrol, Austria, 24.vii.1994 (coll. Erlacher), barcoded, dissected 48c. P. quadrifaria pyrenaea Obth., ♂, Gèdre, Hautes-Pyrénées, France, vii.1921 (ZSM) 48d. P. quadrifaria pyrenaea Obth., ♀, Gèdre, Hautes-Pyrénées, France, 28.vii.1925 (ZSM) 48e. P. quadrifaria sudetica Stern., ♂, Riesengebirge, Poland/Czech Republic, 15.vi.1930 (ZSM) 48f. P. quadrifaria sudetica Stern., ♀, Riesengebirge, Poland/Czech Republic, 4.vii.1932 (ZSM) 49a. Sciadia tenebraria Esp., ♂, Nordkette, North Tyrol, Austria, 14.viii.1970 (TLMF), dissected 49b. S. tenebraria Esp., ♀, Nordkette, North Tyrol, Austria, 27.vii.1939 (TLMF) 49c. S. tenebraria Esp., ♂, underside, Nordkette, North Tyrol, Austria, 14.viii.1970 (TLMF), dissected 49d. S. tenebraria Esp., ♀, underside, Nordkette, North Tyrol, Austria, 27.vii.1939 (TLMF) 49e. S. tenebraria taurusica Huem. & Hausm., ♂, Gartlsee, Schober-Gruppe, East Tyrol, Austria, 17.viii.1991 (TLMF) 49f. S. tenebraria taurusica Huem. & Hausm., ♀, Hochtor, Glockner, Kärnten, Austria, 19.vii.1988 (TLMF), paratype 49g. S. tenebraria taurusica Huem. & Hausm., ♂, underside, Gartlsee, Schober-Gruppe, East Tyrol, Austria, 17.viii.1991 (TLMF) 49h. S. tenebraria taurusica Huem. & Hausm., ♀, underside, Hochtor, Glockner, Kärnten, Austria, 19.vii.1988 (TLMF), paratype 49i. S. tenebraria wockearia Stgr., ♂, Rifugio Mandrone, Adamello, Italy, mid viii.1958 (TLMF) 49j. S. tenebraria wockearia Stgr., ♀, Rifugio Mandrone, Adamello, Italy, mid viii.1958 (TLMF) 49k. S. tenebraria wockearia Stgr., ♂, underside, Rifugio Mandrone, Adamello, Italy, mid viii.1958 (TLMF) 49l. S. tenebraria wockearia Stgr., ♀, underside, Rifugio Mandrone, Adamello, Italy, mid viii.1958 (TLMF) 50a. Sciadia septaria Gn., ♂, Vall Ferrera, Prov. Prov. Lerida, Spain, 30.vii.1957 (coll. Skou), dissected 50b. S. septaria Gn., ♂, Ibon de Sabocos, Panticosa, Prov. Huesca, Spain, 26.vi.1999 (coll. Gaston) 50c. S. septaria Gn., ♀, Gèdre, Hautes-Pyrénées, France (ZSM), dissected 50d. S. septaria Gn., ♀, Ibon de Sabocos, Panticosa, Prov. Huesca, Spain, 26.vi.1999 (coll. Gaston) 50e. S. septaria Gn., ♂, underside, Vall Ferrera, Prov. Prov. Lerida, Spain, 30.vii.1957 (coll. Skou), dissected 51a. Sciadia innuptaria H.-S., ♂, Simonyhütte, Dachstein, Salzburg, Austria (TLMF), dissected 51b. S. innuptaria H.-S., ♂, Simonyhütte, Dachstein, Salzburg, Austria, 17.vii.2011 (coll. Skou) 51c. S. innuptaria H.-S., ♀, Simonyhütte, Dachstein, Salzburg, Austria, 15.vii.1928 (TLMF), dissected 51d. S. innuptaria H.-S., ♂, underside, Simonyhütte, Dachstein, Salzburg, Austria (TLMF), dissected 52a. Sciadia slovenica Ler., ♂, Triglav, Krain, Slovenia, 31.vii.1906, (TLMF), dissected 52b. S. slovenica Ler., ♀, Dolič mountain pass, Triglav, Julian Alps, Slovenia, 29.vii.1909 (TLMF) 52c. S. slovenica Ler., ♀, Bovec, Mount Canin, Slovenia, 15.viii.2002, (TLMF), dissected 52d. S. slovenica Ler., ♂, underside, Triglav, Krain, Slovenia, 31.vii.1906, (TLMF), dissected 52e. S. slovenica Ler., ♀, underside, Dolič mountain pass, Triglav, Julian Alps, Slovenia, 29.vii.1909 (TLMF), dissected 53a. Sciadia dolomitica Huem. & Hausm., ♂, Rifugio Pedrotti alle Rosetta, Trento, Italy, 6.viii.2013 (TLMF), dissected 53b. S. dolomitica Huem. & Hausm., ♀, Antermojasee, South Tyrol, Italy, 15.vii.1924 (ZSM) 53c. S. dolomitica Huem. &  Hausm., ♂, underside, Rifugio Pedrotti alle Rosetta,Trento, Italy, 6.viii.2013 (TLMF), dissected

580

581

PLATE 8 (all specimens natural size) 54a. Sciadia zelleraria Frr., ♂, Völs am Schlern, South Tyrol, Italy, 27.vii.2007 (TLMF) 54b. S. zelleraria Frr., ♂, Rifugio Dibona, Veneto, Italy, 31.vii.2013 (coll. Skou) 54c. S. zelleraria Frr., ♂, Sonnenlagant, Brandnertal, Voralberg, Austria, 27.–28.vii.1983 (coll. Skou) 54d. S. zelleraria Frr., ♂, Rifugio Novezza, Monte Baldo, Trento, Italy, 27.vi.1981 (coll. Skou) 54e. S. zelleraria Frr., ♀, Cima Valdritta, Monte Baldo, Trento, Italy, late vii.1966 (TLMF), dissected 54f. S. zelleraria Frr., ♂, underside, Telegrafo, Monte Baldo, Trento, Italy, mid vii.1969, (TLMF) 54g. S. zelleraria robusta Whli., ♂, Umbrail, Graubünden, Switzerland, 5.viii.1977 (TLMF), dissected 54h. S. zelleraria robusta Whli., ♀, Umbrail, Graubünden, Switzerland, 6.viii.1986 (TLMF), dissected 55a. Sciadia unicolaria Stgr., ♂, Caserne du Restfond, Hautes-Alpes, France, 23.vii.2017 (coll. Skou) 55b. S. unicolaria Stgr., ♂, Cima Fauniera, Caráglio, Piemonte, Italy, 28.vii.2009 (coll. Skou) 55c. S. unicolaria Stgr., ♂, Col d’Izoard, Hautes Alpes, France, 2.viii.2006 (coll. Skou) 55d. S. unicolaria Stgr., ♂, Col de Salèse, le Boréon, Alpes Maritimes, France (coll. Skou) 55e. S. unicolaria Stgr., ♂, Grand St. Bernard, Valle d’Aosta, Italy, 15.–16.vii.2015 (coll. Skou) 55f. S. unicolaria Stgr., ♀, Lago Sommeiller, Bardonecchia, Piemonte, Italy, 7.viii.1998, (coll. Flamigni) 55g. S. unicolaria Stgr., ♂, underside, Col de la Cayolle, Hautes-Alpes, France, 29.–30.vii.2009 (coll. Skou), 55h. 56a. 56b. 56c. 56d. 56e. 56f. 56g. 57a. 57b. 57c. 57d. 58a. 58b. 58c. 58d. 59a. 59b. 59c. 59d.

582

dissected S. unicolaria Stgr., ♂, Navela, Marguareis, Alpes Maritimes, France, 21.–23.vii.1990, (TLMF), dissected Elophos caelibaria Hdr., ♂, Hochtor, Glockner, Kärnten, Austria, e. p. 27.vi.1988 (TLMF), dissected E. caelibaria Hdr., ♂, Nordkette, North Tyrol, Austria, vii.1963 (TLMF) E. caelibaria Hdr., ♂, Pasterze, Kärnten, Austria, 22.vii.1920 (ZMKU) E. caelibaria Hdr., ♀, Hochtor, Glockner, Kärnten, Austria, e. l. 27.vi.1988 (TLMF), dissected E. caelibaria Hdr., ♂, underside, Nordkette, North Tyrol, Austria, vii.1963 (TLMF) E. caelibaria spurcaria Harpe, ♂, Timmelsjoch, North Tyrol, Austria, mid vii. 1974 (TLMF), dissected E. caelibaria spurcaria Harpe, ♀, Timmelsjoch, North Tyrol, Austria, mid vii. 1974 (TLMF), dissected Elophos zirbizensis Pieszcek, ♂, Zirbitzkogel, Steiermark, Austria, (TLMF), dissected E. zirbizensis Pieszcek, ♂, Zirbitzkogel, Steiermark, Austria, 1909 (ZMKU) E. zirbizensis Pieszcek, ♀, Zirbitzkogel, Steiermark, Austria, e. p. 30.vi.1987 (TLMF), dissected E. zirbizensis Pieszcek, ♂, underside, Zirbitzkogel, Steiermark, Austria, 27.vi.1949 (TLMF), dissected Elophos operaria Hbn., ♂, Rax, Steiermark, Austria, 13.vi.1967 (TLMF), dissected E. operaria Hbn., ♀, Eisenerz, Steiermark, 14.vii.1938 (TLMF), dissected E. operaria hoefneri Rbl., ♂, Zirbitzkogel, Steiermark, Austria, 24.–26.vi.1987 (TLMF), dissected E. operaria necopinatus Reser, ♂, Pilatus, Luzern, Switzerland, 4.vii.1977 (TLMF), dissected Elophos andereggaria Harpe, ♂, Valais, Grand St. Bernard, Valle d’Aosta, Italy, 18.vii.2017 (coll. Skou) E. andereggaria Harpe, ♂, Grand St. Bernard, Valle d’Aosta, Italy, 15.–16.vii.2015 (coll. Skou) E. andereggaria Harpe, ♂, Petit St. Bernard, Valle d’Aosta, Italy, 19.vii.2017 (coll. Skou) E. andereggaria Harpe, ♀, Valais, Switzerland, 1879 (TLMF), dissected

583

PLATE 9 (all specimens natural size) 60a. Yezognophos dilucidaria Den. & Schiff., ♂, Wolkenstein, Gröden, South Tyrol, Italy, 11.vii.2015 (coll. B. Müller), dissected 60b. Y. dilucidaria Den. & Schiff., ♂, Zabljak, Durmitor Nat. Park, Montenegro, 20.–23.vii.2014 (coll. Skou) 60c. Y. dilucidaria Den. & Schiff., ♂, Saas-Amlagell, Valais, Switzerland, 13.–14.vii.2015 (coll. Skou) 60d. Y. dilucidaria Den. & Schiff., ♀, Vens, Valle d’Aosta, Italy, 19.viii.2000 (coll. Skou) 60e. Y. dilucidaria Den. & Schiff., ♀, Großglockner, Kärnten, Austria, 15.viii.1999 (coll. Skou) 60f. Y. dilucidaria Den. & Schiff., ♀, Rauschbrunnen, Innsbruck, North Tyrol, Austria, 5.vii.2004 (TLMF) 60g. Y. dilucidaria Den. & Schiff., ♀, underside, Rauschbrunnen, Innsbruck, North Tyrol, Austria, 5.vii.2004 (TLMF) 60h. Y. dilucidaria carpathica Soffn., ♂, Lastun, Black Tcheremosh River, Carpathians, Ukraine 22.vi.2003 (ZMKU) 61a. Yezognophos sproengertsi Püng., ♂, Col de la Boucharde, Alpes-Maritimes, France, 7.vii.2012 (TLMF), dissected 61b. Y. sproengertsi Püng., ♂, Colle di Lombarda, Piemonte, Italy, 27.vii.2009 (coll. Skou) 61c. Y. sproengertsi Püng., ♀, Col de la Boucharde, Alpes-Maritimes, France, 7.vii.2012 (TLMF), dissected 61d. Y. sproengertsi Püng., ♀, Colle di Lombarda, Piemonte, Italy, 31.vii.2005 (coll. Skou) 61e. Y. sproengertsi Püng., ♂, underside, Col de la Boucharde, Alpes-Maritimes, France, 7.vii.2012 (TLMF), dissected 62a. Yezognophos vittaria Thnbg., ♂, Enonkoski, Finland, 6.–7.vii.1996 (coll. Sihvonen) 62b. Y. vittaria Thnbg., ♂, Trosa, Södermanland, Sweden, 9.vi.1973 (coll. Skou) 62c. Y. vittaria Thnbg., ♂, Abisko, Torne Lapmark, Sweden, 29.vi.1974 (coll. Skou) 62d. Y. vittaria Thnbg., ♂, West Sajan Mts., Chakasija, Russia, 11.–15.vi.2012 (coll. Skou) 62e. Y. vittaria Thnbg., ♀, Burfjord, Finmark, Norway, 20.vii.1998 (coll. Sihvonen), dissected 62f. Y. vittaria mendicaria H.-S., ♂, Ritten, Obergrünwald, Bolzano, Italy, 1.vii.2010 (TLMF), dissected 62g. Y. vittaria mendicaria H.-S., ♂, Rifugio Mandrone, Adamello, Trentino-Alto Adige, Italy, 26.–27.vi.2014 (coll. Skou) 62h. Y. vittaria mendicaria H.-S., ♀, Ritten, Obergrünwald, Bolzano, Italy, 1.vii.2010 (TLMF), dissected 63a. Yezognophos serotinaria Den. & Schiff., ♂, Umhausen, North Tyrol, Austria, 4.vii.1945 (TLMF), dissected 63b. Y. serotinaria Den. & Schiff., ♀, East Tyrol, Austria, 18.vii.1985 (TLMF), dissected 63c. Y. serotinaria Den. & Schiff., ♀, Ötz, North Tyrol, Austria (TLMF), dissected 63d. Y. serotinaria Den. & Schiff., ♂, underside, Umhausen, North Tyrol, Austria, 4.vii.1945 (TLMF), dissected 63e. Y. serotinaria dognini Th.-Mieg, ♂, Lac d’Orédon, Hautes Pyrénées, 24.vii.2014, (coll. Skou), dissected 63f. Y. serotinaria dognini Th.-Mieg, ♀, Lac d’Orédon, Hautes Pyrénées, 24.vii.2014, (coll. Skou), dissected 63g. Y. serotinaria serotinoides Whli., ♂, Leuk/Guttet, Valais, Switzerland, e. o. 1.viii.1991 (TLMF), dissected 63h. Y. serotinaria serotinoides Whli., ♂, Vallée des Marveilles, Alpes Maritimes, France, 14.vii.1964 (TLMF), dissected 63i. Y. serotinaria serotinoides Whli., ♂, Estenc, Alpes Maritimes, France, 29.vii.2005 (coll. Skou) 63j. Y. serotinaria serotinoides Whli., ♀, Thorens, Savoie, France, 18.vii.1934 (TLMF), dissected 63k. Y. serotinaria serotinoides Whli., ♀, Isola 2000, Alpes Maritimes, France, 1.viii.2005 (coll. Skou) 63l. Y. serotinaria serotinoides Whli., ♀, Pont du Fau, Alpes de Haute Provence, France, 9.vii.1994 (coll. Skou)

584

585

.

PLATE 10 (all specimens natural size) 64a. Crocota tinctaria Hbn., ♂, Monte Rondinaio, Lombardia, Italy, 5.vii.1999 (coll. Sihvonen), dissected 64b. C. tinctaria Hbn., ♂, Furkapaß, Valais, Switzerland, 10.vii.1979 (coll. Skou) 64c. C. tinctaria Hbn., ♂, Monte Rondinaio, Lombardia, Italy, 5.vii.1999 (coll. Sihvonen) 64d. C. tinctaria Hbn., ♂, Mikulichyn, Iv.-Frankovskaya obl., Ukraine (SMNH) 64e. C. tinctaria Hbn., ♀, Demonte, Alpi Cozie, Piemonte, Italy, 2.viii.2010 (TLMF), dissected 65a. Crocota pelietaria Dup., ♂, L’Hospitalet, Ariège, France, 13.vii.1946 (TLMF), dissected 65b. C. pelietaria Dup., ♂, Port de la Bonaigua, Prov. Lerida, Spain, 31.vii.2007 (coll. Skou) 65c. C. pelietaria Dup., ♂, Port de la Bonaigua, Prov. Lerida, Spain, 31.vii.2007 (coll. Skou) 65d. C. pelietaria Dup., ♀, Val Ferrera, Prov. Lerida, Spain, 25.vii.1957 (coll. Skou), dissected 65e. C. pelietaria Dup., ♀, Font Romeu, Pyrénées Orientales, France, 31.vii.1958 (coll. Skou) 66a. Crocota pseudotinctaria Ler., ♂, Leukerbad, Valais, Switzerland, e. o. 24.vii.1993 (coll. Sihvonen), dis66b. 66c. 66d. 66e. 67a. 67b. 67c. 67d. 67e. 68a. 68b. 68c. 68d. 68e. 68f. 68g. 68h. 69a. 69b. 69c. 69d. 69e. 69f. 69g. 69h.

586

sected C. pseudotinctaria Ler., ♂, Gran St. Bernard, Aosta, 20.viii.2000 (coll. Skou) C. pseudotinctaria Ler., ♂, Argentera, Piemonte, Italy, 3.vii.1952 (coll. Skou) C. pseudotinctaria Ler., ♂, le Boréon, Col de Salèse, Alpes Maritimes, 3.viii.2005 (coll. Skou) C. pseudotinctaria Ler., ♀, Leukerbad, Valais, Switzerland, e. o. 24.vii.1993 (coll. Sihvonen), dissected Crocota niveata Scop., ♂, Saualpe, Kärnten, Austria, 12.vii.2011 (coll. Sihvonen), dissected C. niveata Scop., ♂, Brett Höhe, Nockgebiet, Kärnten, Austria, 10.vii.1993 (coll. Skou) C. niveata Scop., ♂, Saualpe, Kärnten, Austria, (ZMKU) C. niveata Scop., ♀, Saualpe, Kärnten, Austria, 14.vii.2011 (coll. Sihvonen), dissected C. niveata Scop., ♀, Obertauern, Salzburg, Austria, 2.–3.viii.2013 (coll. Skou) Synopsia sociaria Hbn., ♂, Albarracin, Prov. Teruel, Spain, 18.vi.2007 (coll. B. Müller) S. sociaria Hbn., ♂, Liljanovo, Pirin Mountains, Bulgaria, 1.vi.1981 (coll. B. Müller) S. sociaria Hbn., ♂, Kranevo, Varna, Black Sea Coast, Bulgaria, e. o. early vi.1984 (coll. Gelbrecht) S. sociaria Hbn., ♂, Liljanovo, Pirin Mountains, Bulgaria, 2.vi.1981 (coll. B. Müller) S. sociaria Hbn., ♀, Liljanovo, Pirin Mountains, Bulgaria,29.v.1981 (coll. B. Müller), barcoded S. sociaria Hbn., ♀, Kranevo, Varna,Black Sea Coast, Bulgaria, e. o. 5.viii.1984 (coll. Gelbrecht) S. sociaria Hbn., ♀, Budaörs, Budapest, Hungary, e. l. 3.v.1986 (coll. B. Müller), barcoded S. sociaria Hbn., ♀, Geroplatanos, Epirus, Greece, 2.vi.2010 (coll. Skou) Phthonandria strictaria Led., ♂, Jantyshevo village, Bashkiria Sakmara river, South Ural, Russia, 21.vi.1996 (coll. Nupponen) P. strictaria Led., ♂, Kuvandyk village, South Ural, Russia, 1.–7.vi.1996 (coll. Skou) P. strictaria Led., ♀, Jantyshevo village, Bashkiria Sakmara river, South Ural, Russia, 21.vi.1996 (coll. ­Nupponen) P. strictaria Led., ♀, Kuvandyk village, South Ural, Russia, 13.–13.vi.1997 (coll. Skou) P. strictaria Led., ♂, Kuraiskaja steppe, Altai Mountains, Russia 25.vi.2000 (coll. Nupponen) P. strictaria Led., ♀, Ust-Sema village, Katun valley, Altai Mountains, Russia, 23.vi.2000 (coll. Nupponen) P. strictaria Led., ♂, Tyrma, Bureinskyi mountains, Amur, Russia 1.vii.1997 (coll. Skou), dissected P. strictaria Led., ♀, Tyrma, Bureinskyi mountains, Amur, Russia 15.vi.1997 (coll. Skou), dissected

587

PLATE 11 (all specimens natural size) 70a. Menophra abruptaria Thnbg., ♂, Krk island, Croatia, 11.v.1982 (coll. B. Müller), barcoded 70b. M. abruptaria Thnbg., ♂, Podgora, Dalmatia, Croatia, 23.iv.2017 (coll. B. Müller) 70c. M. abruptaria Thnbg., ♂, Drosia, Réthimno, Crete, Greece, 8.iv.2005 (coll. B. Müller), barcoded 70d. M. abruptaria Thnbg., ♂, Ghisoni, Corsica, France, 1.vi.2008 (coll. B. Müller), barcoded 70e. M. abruptaria Thnbg., ♂, Aiguafreda, Prov. Gerona, Spain, e. o. 17.vii.1991 (coll. B. Müller) 70f. M. abruptaria Thnbg., ♂, Salema Camping, Algarve, Portugal, 21.iv.2017 (coll. Skou) 70g. M. abruptaria Thnbg., ♀, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.iii.–3.iv.2016 (coll. Skou) 70h. M. abruptaria Thnbg., ♀, Krk island, Croatia, 11.v.1982 (coll. B. Müller) 71a. Menophra japygiaria Costa, ♂, San Pantaleo, Sardinia, Italy 16.iv.2007 (coll. B. Müller) 71b. M. japygiaria Costa, ♂, Tabernas, Prov. Almeria, Spain, 24.iii.2006 (coll. B. Müller) 71c. M. japygiaria Costa, ♀, San Pantaleo, Sardinia, Italy, 9.iv.2007 (coll. B. Müller) 71d. M. japygiaria Costa, ♀, Sorbas, Prov. Almeria, Spain, 26.iii.2006 (coll. B. Müller), barcoded 71e. M. japygiaria Costa, ♀, Cabo Pecora, Sardinia, Italy, 16.v.2004 (coll. Skou) 71f. M. japygiaria Costa, ♀, Lido di Pietrapaola, Calabria, Italy, 2.vi.2005 (coll. Skou) 71g. M. japygiaria Costa, ♂, Agadir, Morocco, 11.iv.2015 (coll. Skou) 71h. M. japygiaria Costa, ♂, Aransis, Tremp Valley, Lerida, Spain, 8.vii.1993 (coll. Skou) 72a. Menophra berenicidaria Trti., ♂, Drosia, Réthimno, Crete, Greece, 7.iv.2005 (coll. B. Müller) 72b. M. berenicidaria Trti., ♀, Antalya, Turkey, e. o. 27.iv.1995 (coll. B. Müller) 72c. M. berenicidaria Trti., ♀, Stalida, Iraklion, Crete, Greece, e. o. 6.iv.1999 (coll. B. Müller) 72d. M. berenicidaria Trti., ♀, Stalida, Iraklion, Crete, Greece, e. o. 6.iv.1999 (coll. B. Müller) 73a. Menophra harterti Roths., ♂, Guelt ès Stel, Algeria, 11.x.1931 (ZSM) 73b. M. harterti Roths., ♂, Guelt ès Stel, Algeria, x.1929 (ZSM) 73c. M. harterti Roths., ♂, Algeria, vii.1907 (ZSM) 73d. M. harterti Roths., ♂, Douéra, Algeria, 15.x.1930 (ZSM) 73e. M. harterti Roths., ♀, Guelt ès Stel, Algeria, 16.–23.ix.1933 (ZSM), 74a. Menophra thuriferaria Zerny, ♂, Albarracin, Prov. Teruel, Spain, 22.viii.2001 (coll. Skou) 74b. M. thuriferaria Zerny, ♂, Tramacastilla, Prov. Teruel, Spain, 24.viii.2001 (coll. Skou) 74c. M. thuriferaria Zerny, ♂, Albarracin, Prov. Teruel, Spain, 18.vii.2007 (coll. B. Müller) 74d. M. thuriferaria Zerny, ♀, Albarracin, Prov. Teruel, Spain, 30.ix.2009 (coll. Skou) 74e. M. thuriferaria Zerny, ♀, Albarracin, Prov. Teruel, Spain, e. o. 26.vi.2006 (coll. B. Müller) 75a. Menophra nycthemeraria Geyer, ♂, Chaudon-Noranthe, Provence, France, e. o. 2.vi.1995 (coll.

B. Müller) M. nycthemeraria Geyer, ♂, Chaudon-Noranthe, Provence, France, e. o. 2.vi.1995 (coll. B. Müller) M. nycthemeraria Geyer, ♀, Alhama, Prov. Murcia, Spain, 26.v.1998 (coll. Skou) M. nycthemeraria Geyer, ♀, Albarracin, Prov. Teruel, Spain, 3.vi.2007 (coll. B. Müller) Menophra annegreteae Skou, ♂, Tabernas, Prov. Almeria, Spain, 24.–25.v.2006 (coll. Skou), paratype M. annegreteae Skou, ♂, Tabernas, Prov. Almeria, Spain, 24.–25.v.2006 (coll. Skou), paratype M. annegreteae Skou, ♂, Cabo de Gata village, Prov. Almeria, Spain, 16.–17.iv.2001 (coll. Skou), paratype 76d. M. annegreteae Skou, ♀, Cabo de Gata village, Prov. Almeria, Spain, 16.–17.iv.2001 (coll. Skou), paratype 76e. M. annegreteae Skou, ♀, Tabernas, Prov. Almeria, Spain, 15.v.2006 (coll. Skou), paratype

75b. 75c. 75d. 76a. 76b. 76c.

588

589

PLATE 12 (all specimens natural size) 77a. Cleorodes lichenaria Hufn., ♂, St. Barnabé, Alpes Maritimes, France, 12.vi.1937 (ZSM) 77b. C. lichenaria Hufn., ♂, Amfreville, Eure, France, 10.vii.1954 (ZSM) 77c. C. lichenaria Hufn., ♂, Amfreville, Eure, France, 11.vii.1954 (ZSM) 77d. C. lichenaria Hufn., ♂, Vizzavona, Corsica, France, 6.vi.1951 (ZSM) 77e. C. lichenaria Hufn., ♀, Fazèt de Jugny, Cote d’Or, France, 26.–27.vi.1965 (ZSM) 78a. Zernyia granataria Stgr., ♂, Albarracin, Prov. Teruel, Spain, 4.ix.2008 (coll. B. Müller) 78b. Z. granataria Stgr., ♂, Sedella, Prov. Malaga, Spain, 18.ix.2009 (coll. Skou) 78c. Z. granataria Stgr., ♂, Tosos, Prov. Zaragoza, Spain, 20.vii.2001 (coll. Skou) 78d. Z. granataria Stgr., ♀, Candasnos, Prov. Huesca, Spain, 28.viii.2001 (coll. Skou), dissected 78e. Z. granataria Stgr., ♂, Albarracin, Prov. Teruel, Spain, 7.viii.1988 (coll. Skou) 78f. Z. granataria Stgr., ♂, Candasnos, Prov. Huesca, Spain, 13.–14.ix.2002 (coll. Skou) 78g. Z. granataria Stgr., ♀, Tosos, Prov. Zaragoza, Spain, 15.ix.2002 (coll. Skou) 78h. Z. granataria Stgr., ♀, Calar Alto, Sierra de los Filabres, Prov. Almeria, Spain, 24.viii.2001 (coll. Skou) 79a. Megalycinia serraria Costa, ♂, Terranovo, Monto Pollino, Calabria, Italy, 10.–16.viii.1980 (coll. Skou) 79b. M. serraria Costa, ♂, Lagonegro, Monte Sirino, Basilicata, Italy, 31.viii.2018 (ZSM) 79c. M. serraria Costa, ♀, Silvana Mansio, Calabria, Italy, 31.viii.2000 (ZSM), dissected 79d. M. serraria Costa, ♀, Lagonegro, Basilicata, Italy, 31.viii.2018 (ZSM) 80a. Dasypteroma thaumasia Stgr., ♂, Aranjuez, Prov. Madrid, Spain, 7.x.2008 (coll. B. Müller), dissected 80b. D. thaumasia Stgr., ♂, Aranjuez, Prov. Madrid, Spain, 7.x.2008 (coll. B. Müller), 80c. D. thaumasia Stgr., ♂, Aranjuez, Prov. Madrid, Spain, 7.x.2008 (coll. B. Müller), 80d. D. thaumasia Stgr., ♂, Candasnos, Prov. Huesca, Spain, 13.–14.ix.2002 (coll. Skou) 80e. D. thaumasia Stgr., ♀, San Ildefonso, Castilia, Spain, undated (coll. MNHU) 81a. Microbiston lanaria Ev., ♂, Kapustin Yar, Astrakhanskaya obl., Russia, 28.iii.1975 (ZMKU) 81b. M. lanaria Ev., ♂, Uralsk, Russia, 1913 (ZMKU) 81c. M. lanaria Ev., ♂, Kapustin Yar, Astrakhanskaya obl., Russia, 16.iii.1977 (ZMKU) 81d. M. lanaria Ev., ♂, Kapustin Yar, Astrakhanskaya obl., Russia, 22.iii.1974 (ZMKU) 81e. M. lanaria Ev., ♀, Uralsk, Russia, 1913 (ZMKU) 81f. M. lanaria Ev., ♀, Uralsk, Russia, 1913 (ZMKU) 82a. Apocheima hispidaria Den. & Schiff., ♂, Königs Wusterhausen, Mark Brandenburg, Germany, 1.iv.1980 82b. 82c. 82d. 82e. 83a. 83b. 83c. 83d. 83e. 83f. 83g. 83h.

590

(coll. B. Müller) A. hispidaria Den. & Schiff., ♂, Nauen, Brandenburg, Germany, 23.iii.1981 (coll. B. Müller) A. hispidaria Den. & Schiff., ♂, Egebjerg bakker, Fyn, Denmark, 19.iv.1976 (coll. Skou) A. hispidaria Den. & Schiff., ♂, Retzau, Sachsen-Anhalt, Germany, 16.iv.1980 (coll. B. Müller) A. hispidaria Den. & Schiff., ♀, Königs Wusterhausen, Mark Brandenburg, Germany, 28.iii.1980 (coll. ­ elbrecht) G Phigalia pilosaria Den. & Schiff., ♂, St. Gangloff, Thüringen, Germany, 25.ii.1974 (coll. B. Müller) P. pilosaria Den. & Schiff., ♂, Pankow, Rosenthal, Berlin, Germany, 25.ii.2012 (coll. B. Müller) P. pilosaria Den. & Schiff., ♂, Erfurt, Thüringen, Germany 26.i.1974 (coll. B. Müller) P. pilosaria Den. & Schiff., ♀, Pottenstein, Bayern, Germany, 11.iii.1994 (coll. B. Müller) P. pilosaria Den. & Schiff., ♂, Nauen, Brandenburg, Germany, 26.ii.1978 (coll. B. Müller) P. pilosaria Den. & Schiff., ♂, Kirkeby Sand, Fyn, Denmark, 20.iii.2010 (coll. Skou) P. pilosaria Den. & Schiff., ♂, Erfurt, Thüringen, Germany, 26.i.1974 (coll. B. Müller) P. pilosaria Den. & Schiff., ♀, Nauen, Brandenburg, Germany, 23.iii.1981 (coll. B. Müller)

591

PLATE 13 (all specimens natural size) 84a. Hypomecis roboraria Den. & Schiff., ♂, Waren, Mecklenburg-Vorpommern, Germany, 3.vii.1973 84b. 84c. 84d. 84e. 84f. 85a. 85b. 85c. 85d. 85e. 85f. 86a. 86b. 86c. 86d. 86e. 86f. 86g. 86h. 86i. 86j. 87a. 87b. 87c. 87d. 87e. 87f.

592

(MNHU) H. roboraria Den. & Schiff., ♂, Soca, Julian Alps, Slovenia, 10.vii.2002 (coll. B. Müller), barcoded H. roboraria Den. & Schiff., ♂, Snezhinsk, South Ural, Russia, 18.vi.2015 (coll. Skou) H. roboraria Den. & Schiff., ♀, Maryanovka, Kievskaya obl., Ukraine, 14.viii.2010 (ZMKU) H. roboraria Den. & Schiff., ♀, Summt, Brandenburg, Germany, 12.vi.2015 (coll. B. Müller) H. roboraria Den. & Schiff., ♀, Grano, Brandenburg, Germany, e. o. 11.v.1989 (coll. B. Müller Hypomecis punctinalis Scop., ♂, Liljanovo, Pirin Mountains, Bulgaria, 1.vi.1981 (coll. B. Müller), barcoded H. punctinalis Scop., ♂, Nebbeskov, Langeland, Denmark, 23.vi.1973 (coll. Skou) H. punctinalis kerstinae Fisch. & Lew., ♂, Platres, Troodos Mountains, Cyprus, 15.v.1999 (coll. Skou) H. punctinalis Scop., ♀, Garda-Marchiaga, Trento, Italy, 28.vii.1990 (coll. B. Müller) H. punctinalis Scop., ♀, Vidreras, Prov. Girona, Spain, 5.vii.1993 (coll. B. Müller), barcoded H. punctinalis Scop., ♀, Mühlenbeck, Brandenburg, Germany, e. o. 8.iv.1987 (coll. B. Müller) Ematurga atomaria L., ♂, Ilmenau, Thüringen, Germany, 9.v.1975 (coll. B. Müller) E. atomaria L., ♂, Popina Laka, Liljanovo, Pirin Mountains, Bulgaria, 16.vi.1981 (coll. B. Müller) E. atomaria L., ♂, Gornje, Tučepi, Dalmatia, Croatia, 30.iv.2017 (coll. B. Müller) E. atomaria L., ♂, Lorry-Mardigny, Lorraine, France, 23.iv.2006, (coll. B. Müller) E. atomaria L., ♂, Liljanovo, Pirin Mountains, Bulgaria, 20.v.1982 (coll. B. Müller) E. atomaria L., ♂, Garda-Marchiaga, Trento, Italy, 5.viii.1990 (coll. B. Müller), barcoded E. atomaria L., ♂, Oberhaverbeck, Niedersachsen, Germany, 19.v.1978 (coll. Wegner) E. atomaria L., ♂, Oberhaverbeck, Niedersachsen, Germany, 19.v.1978 (coll. Wegner) E. atomaria L., ♀, Summt, Brandenburg, Germany, 14.vi.2005 (coll. B. Müller) E. atomaria L., ♀, Sejs, East Jutland, Denmark, 15.v.1982 (coll. Skou) Lycia hirtaria Cl., ♂, Val d’Isone, Medeglia, Ticino, Switzerland, 21.–31.iii.2001 (coll. B. Müller) L. hirtaria Cl., ♂, Sórgono, Sardinia, Italy, 18.v.2004 (coll. Skou) L. hirtaria Cl., ♂, Asserbo, Northeast Zealand, Denmark, 16.v.1980 (coll. Skou) L. hirtaria Cl., ♂, Spiazzi, Veneto, Italy, 22.iv.2009 (coll. Skou) L. hirtaria Cl., ♀, Ahrensfelde, Berlin, Germany, 19.iv.1970 (coll. B. Müller) L. hirtaria Cl., ♀, Allerød, Northeast Zealand, Denmark, 30.iv.1980 (coll. Skou)

593

PLATE 14 (all specimens natural size) 88a. Lycia necessaria Zell., ♂, Mtatzminda, Tbilissi, Georgia, e. l. 26.iii.1989 (coll. B. Müller), dissected 88b. L. necessaria Zell., ♂, Babuna, Izvor, Macedonia, 21.iv.1985 (coll. Skou) 88c. L. necessaria Zell., ♂, Kas, Antalya, Turkey, 25.v.1982 (SMNS) 88d. L. necessaria Zell., ♀, Mtatzminda, Tbilissi, Georgia, e. l. 17.–21.vi.1988 (coll. Gelbrecht) 89a. Lycia lapponaria Bsd., ♂, Dammossen, Västmanland, Sweden, 30.iv.1973 (coll. Skou) 89b. L. lapponaria Bsd., ♂, Tervajoki, Etelä-Pohjanmaa, Finland, 15.v.1941 (MNHU) 89c. L. lapponaria Bsd., ♂, Dammossen, Västmanland, Sweden, 30.iv.1973 (coll. Skou) 89d. L. lapponaria Bsd., ♂, Scotland, United Kingdom, e. o. 16.ii.1937 (MNHU) 89e. L. lapponaria Bsd., ♀, Damtorpsmossen, Västmanland, Sweden, 27.iv.1973 (coll. Skou) 89f. L. lapponaria Bsd., ♀, Heidal, Northern Oppland, Norway, 1987 (coll. Skou) 90a. Lycia isabellae Harr., ♂, Kyjov, Moravia, Czech Republic, 30.iii.1987 (coll. B. Müller) 90b. L. isabellae Harr., ♂, Kyjov, Moravia, Czech Republic, 30.iii.1987 (coll. B. Müller) 90c. L. isabellae Harr., ♂, Studce, Bohemia, Czech Republic, iii.1995 (coll. Skou) 90d. L. isabellae Harr., ♂, St. Aegyd am Neuwalde, Lower Austria, Austria, e. p. 11.ii.1978 (coll. Skou) 90e. L. isabellae Harr., ♀, Kyjov, Moravia, Czech Republic, 30.iii.1987 (coll. B. Müller) 90f. L. isabellae Harr., ♀, Kyjov, Moravia, Czech Republic, 30.iii.1987 (coll. B. Müller) 91a. Lycia pomonaria Hbn., ♂, Paulinenaue, Brandenburg, Germany, 12.iii.2007 (coll. B. Müller) 91b. L. pomonaria Hbn., ♂, Brieselang, Brandenburg, Germany, 13.iii.2007 (coll. B. Müller) 91c. L. pomonaria Hbn., ♂, Hollola, Etelä-Häme, Finland, 21.iv.1973 (coll. Skou) 91d. L. pomonaria Hbn., ♂, Kisdobsza, Somogy, Hungary, 9.iii.1983 (coll. Skou) 91e. L. pomonaria Hbn., ♀, Vicinity of Berlin, Germany, e. l. 9.ii.1927 (coll. B. Müller) 91f. L. pomonaria Hbn., ♀, Las Wolski, Krakow, Poland, e. o. 21.iii.1958 (coll. Skou) 92a. Lycia zonaria Den. & Schiff., ♂, Odvashegy, Budaörs, Hungary, 11.iv.1985 (coll. B. Müller) 92b. L. zonaria Den. & Schiff., ♂, Odvashegy, Budaörs, Hungary, 4.iv.1985 (coll. B. Müller) 92c. L. zonaria Den. & Schiff., ♂, Billund, West Jutland, Denmark, 21.iv.1996 (coll. Skou) 92d. L. zonaria Den. & Schiff., ♂, Prignitz, Brandenburg, Germany, 6.iv.1920 (coll. B. Müller) 92e. L. zonaria Den. & Schiff., ♀, Dreetz, Brandenburg, Germany, 22.iv.1991 (coll. Gelbrecht) 92f. L. zonaria Den. & Schiff., ♀, Dreetz, Brandenburg, Germany, 22.iv.1991 (coll. Gelbrecht), 93a. Lycia alpina Sulz., ♂, Zermatt, Valais, Switzerland, 20.vi.1909 (MNHU) 93b. L. alpina Sulz., ♂, Crosedomini Pass, Lombardy, Italy, 10.iv.2003 (coll. Skou) 93c. L. alpina Sulz., ♂, Mangart, Julian Alps, Slovenia, 28.v.2005 (coll. Skou) 93d. L. alpina Sulz., ♀, Vallarsa, South Tyrol, Italy (ZSM) 93e. L. alpina Sulz., ♀, Mangart, Julian Alps, Slovenia, 28.v.2005 (coll. Skou) 94a. Lycia graecarius Stgr., ♂, Punat, Krk Island, Croatia, 31.iii.2003 (coll. B. Müller) 94b. L. graecarius Stgr., ♂, Vivaro, Friuli-Venezia Giuglia, Italy, 17.iv.2009 (coll. Skou) 94c. L. graecarius Stgr., ♂, Kristalopigi, Florini, Greece, 17.iv.1987 (coll. Skou) 94d. L. graecarius Stgr., ♀, Liljanovo, Pirin Mountains, Bulgaria, 6.v.1982 (coll. B. Müller) 94e. L. graecarius Stgr., ♀, Liljanovo, Pirin Mountains, Bulgaria, e. l. 20.ii.1982 (coll. B. Müller) 95a. Lycia florentina Stefanelli, ♂, Forlì, Emilia-Romagna, Italy, e. o. 2.iii.1992 (coll. Skou) 95b. L. florentina Stefanelli, ♂, Malagrotta, Rome, Italy, 9.iii.1959 (MNHU) 95c. L. florentina Stefanelli, ♂, Modena, Emilia-Romagna, Italy, 1892 (MNHU) 95d. L. florentina Stefanelli, ♀, Napoli, Campania, Italy, v.1905 (MNHU) 95e. L. florentina Stefanelli, ♀, Acquatraversa, Rome, Italy, 26.iii.1940 (coll. Skou)

594

595

PLATE 15 (all specimens natural size) 96a. Biston strataria Hufn., ♂, Dannenreich, Brandenburg, Germany, 12.iv.1980 (coll. B. Müller) 96b. B. strataria Hufn., ♂, Dannenreich, Brandenburg, Germany, 11.iv.1980 (coll. B. Müller) 96c. B. strataria Hufn., ♂, Ziegelrodaer Forst, Sachsen-Anhalt, Germany, e. l. 25.iii.1961 (MNHU) 96d. B. strataria Hufn., ♂, San Ildefonso, Prov. Segovia, Spain, 1910 (MNHU) 96e. B. strataria Hufn., ♀, Potsdam, Brandenburg, Germany 16.iv.1986 (coll. B. Müller) 96f. B. strataria Hufn., ♀, Arnstadt, Thüringen, Germany, 3.iv.1971 (coll. B. Müller) 97a. Biston rosenbaueri Müller, ♂, Punat, Krk Island, Croatia, 27.iii.2005 (coll. Friedrich) 97b. B. rosenbaueri Müller, ♂, Vrbnik, Krk Island, Croatia, 26.iii.2005 (coll. Friedrich) 97c. B. rosenbaueri Müller, ♂, Kristalopigi, Florini, Greece, 17.iv.1987 (coll. Skou), dissected 98a. Biston achyra Whli., ♂, Kokkari, Samos, Greece, 9.iii.2010 (coll. Fritsch), barcoded 98b. B. achyra Whli., ♂, Kokkari, Samos, Greece, 9.iii.2010 (coll. Skou) 98c. B. achyra Whli., ♂, Arslanli, Mersin, Turkey, 19.–21.iv.1987 (coll. Skou) 99a. Biston betularia L., ♂, Ghisoni, Corsica, France, 7.vi.2008 (coll. B. Müller) 99b. B. betularia L., ♂, Horní Maršov, Hradec Králové, Czech Republic, 29.vi.1983 (coll. B. Müller) 99c. B. betularia L., ♀, Leukerbad, Valais, Switzerland, 14.vii.1993 (coll. B. Müller) 99d. B. betularia L., ♂, Zehdenick, Brandenburg, Germany, 6.vi.1954 (MNHU) 99e. B. betularia L., ♂, Horní Maršov, Hradec Králové, Czech Republic, 30.vi.1983 (coll. B. Müller) 99f. B. betularia L., ♀, Horní Maršov, Hradec Králové, Czech Republic, 29.vi.1983 (coll. B. Müller) 100a. Erannis defoliaria Cl., ♂, Ilmenau, Thüringen, Germany, 22.x.1969 (coll. B. Müller) 100b. E. defoliaria Cl., ♂, Tiefensee, Brandenburg, Germany, 13.xi.1980 (coll. B. Müller) 100c. E. defoliaria Cl., ♂, Ilmenau, Thüringen, Germany, 24.x.1969 (coll. B. Müller) 100d. E. defoliaria Cl., ♂, Chemnitz, Sachsen, Germany, 31.x.1979 (coll. B. Müller) 100e. E. defoliaria Cl., ♂, Weimar, Thüringen, Germany, 15.xi.2014 (coll. B. Müller) 100f. E. defoliaria Cl., ♂, Kresna Gorge, Pirin Mountains, Bulgaria, e. l. 28.xii.1982 (coll. B. Müller) 100g. E. defoliaria Cl., ♂, Arnstadt, Thüringen, Germany, e. l. 24.x.1969 (coll. B. Müller) 100h. E. defoliaria Cl., ♀, Tiefensee, Brandenburg, Germany, 13.xi.1980 (coll. B. Müller) 100i. E. defoliaria Cl., ♀, Tiefensee, Brandenburg, Germany, 13.xi.1980 (coll. B. Müller)

596

597

PLATE 16 (all specimens natural size) 101a. Erannis ankeraria Stgr., ♂, Podgorski, Kras Brkini, Slovenia, 20.iii.2006 (coll. Leipnitz) 101b. E. ankeraria Stgr., ♂, Podgorski, Kras Brkini, Slovenia, 20.iii.2006 (coll. Skou) 101c. E. ankeraria Stgr., ♂, Podgorski, Kras Brkini, Slovenia, 20.iii.2006 (coll. Skou) 101d. E. ankeraria Stgr., ♀, Podgorje, Sezana, Slovenia, e. o. 24.iii.2012 (coll. May) 102a. Erannis declinans Stgr., ♂, Gölyurt Geçidi, Mescit Dağları, Erzurum, Turkey, 7.x.2000 (coll. B. Müller) 102b. E. declinans Stgr., ♂, Arda Valley, Eastern Rhodopi Mountains, Bulgaria, 8.–9.xii.2000 (coll. Skou) 102c. E. declinans Stgr., ♂, Kayluka Park, Pleven, Bulgaria, 16.xi.1987 (coll. Skou) 102d. E. declinans Stgr., ♀, Gölyurt Geçidi, Mescit Dağları, Erzurum, Turkey, larva 4.vii.2002 (coll. Gelbrecht) 103a. Kresnaia beschkovi Ganev, ♂, Kresna Gorge, Blagoevgrad, Bulgaria, 3.xii.1988 (coll. Skou) 103b. K. beschkovi Ganev, ♂, Kresna Gorge, Blagoevgrad, Bulgaria, 3.xii.1988 (coll. Skou) 103c. K. beschkovi Ganev, ♂, Vlahi Village, Pirin-Kresna Gorge, Blagoevgrad, Bulgaria, 17.xi.2014 (coll. ­Beshkov) 103d. K. beschkovi Ganev, ♂, Vlahi Village, Kresna Gorge, Blagoevgrad, Bulgaria, 17.xi.2014 (coll. Beshkov) 103e. K. beschkovi Ganev, ♀, Kresna Gorge, Blagoevgrad, Bulgaria, 26.xi.2009 (coll. Beshkov) 104a. Agriopis leucophaearia Den. & Schiff., ♂, Rippersroda, Thüringen, Germany, 16.ii.1972 (coll.

B. Müller) 104b. A. leucophaearia Den. & Schiff., ♂, Glienicke, Brandenburg, Germany, 28.ii.1978 (coll. B. Müller) 104c. A. leucophaearia Den. & Schiff., ♂, Ahrensfelde, Brandenburg, Germany, 28.iii.1976 (coll. B. Müller) 104d. A. leucophaearia Den. & Schiff., ♂, Brieselang, Brandenburg, Germany, 26.ii.1978 (coll. B. Müller) 104e. A. leucophaearia Den. & Schiff., ♀, Zeuthen, Brandenburg, Germany, 1.iv.1980 (coll. B. Müller) 105a. Agriopis bajaria Den. & Schiff., ♂, Road to Pico Veleta, Sierra Nevada, Prov. Granada, Spain, 20.11.1980 (coll. Skou) 105b. A. bajaria Den. & Schiff., ♂, Arnstadt, Thüringen, Germany, 23.x.1977 (coll. B. Müller) 105c. A. bajaria Den. & Schiff., ♂, Kresna Gorge, Blagoevgrad, Bulgaria, e. l. 17.xii.1982 (coll. B. Müller) 105d. A. bajaria Den. & Schiff., ♂, Arnstadt, Thüringen, Germany, 23.x.1977 (coll. B. Müller) 105e. A. bajaria Den. & Schiff., ♀, Palau, Sardinia, Italy, e. l. 10.iv.2007 (coll. B. Müller) 106a. Agriopis aurantiaria Hbn., ♂, Fiskartorpet, Upland, Sweden, 26.x.1973, (coll. Skou) 106b. A. aurantiaria Hbn., ♂, Tiefensee, Brandenburg, Germany, 16.x.1980 (coll. B. Müller) 106c. A. aurantiaria Hbn., ♂, Tiefensee, Brandenburg, Germany, 16.x.1980 (coll. B. Müller) 106d. A. aurantiaria Hbn., ♂, Arnstadt, Thüringen, Germany, e. l. 26.x.1969 (coll. B. Müller) 106e. A. aurantiaria Hbn., ♂, Aleksandrovska, Odesskaya obl., Ukraine, 5.xi.2010 (ZMKU) 106f. A. aurantiaria Hbn., ♂, Melchow, Brandenburg, Germany, 24.x.1980 (coll. B. Müller) 106g. A. aurantiaria Hbn., ♂, Telfes, North Tyrol, Austria, 9.xi.1965 (MNHU) 106h. A. aurantiaria Hbn., ♀, Arnstadt, Thüringen, Germany e. l. 2.xi.1969 (coll. B. Müller) 107a. Agriopis budashkini Kostjuk, ♂, Karadagh, Crimea, Ukraine, 1.xii.1987 (ZMKU), paratype 107b. A. budashkini Kostjuk, ♂, Asfendos, Crete, Greece, e. o. 8.xi.2015 (coll. Leipnitz) 107c. A. budashkini Kostjuk, ♂, Asfendos, Crete, Greece, e. o. 28.xii.2014 (coll. B. Müller) 107d. A. budashkini Kostjuk, ♀, Krasnolesye, Crimea, Ukraine, 17.xi.2010 (ZMKU) 107e. A. budashkini Kostjuk, ♀, Asfendos, Crete, Greece, e. o. 14.xi.2015 (coll. B. Müller) 108a. Agriopis marginaria F., ♂, Prignitz, Brandenburg, Germany, 10.ii.2008 (coll. B. Müller) 108b. A. marginaria F., ♂, Alt Schadow, Brandenburg, Germany, 9.ii.2008 (coll. B. Müller) 108c. A. marginaria F., ♂, Grande Pevero, Sardinia, Italy, larva 28.iv.2007 (coll. B. Müller) 108d. A. marginaria F., ♀, Glienicke, Brandenburg, Germany, 27.iii.1978 (coll. B. Müller) 108e. A. marginaria F., ♀, Zeuthen, Brandenburg, Germany, 1.iv.1980 (coll. B. Müller)

598

599

PLATE 17 (all specimens natural size) 109a. Calamodes occitanaria Dup., ♂, St. Julien, Alpes-de-Haute-Provence, France, 30.ix.1996 (coll. B. Müller) 109b. C. occitanaria Dup., ♂, Albarracin, Prov. Teruel, Spain, 15.ix.2008 (coll. B. Müller), barcoded 109c. C. occitanaria Dup., ♂, Lagoa de Santo André, Prov. Alentejo, Portugal, 25.ix.2009 (coll. Skou) 109d. C. occitanaria Dup., ♀, Bezas, Prov. Teruel, Spain, 9.ix.2008 (coll. B. Müller) 109e. C. occitanaria Dup., ♀, Mallemoisson, Alpes-de-Haute Provence, France, 3.x.1996 (coll. B. Müller) 110a. Calamodes melanaria Obth., ♂, Fiumara Trionto, Cropalati, Calabria, Italy, 4.x.2010 (coll. Skou) 110b. C. melanaria Obth., ♂, Fiumara Trionto, Calábria, Italy, 21.x.1999 (ZSM) 110c. C. melanaria Obth., ♂, Jebel Chambi, Al Qasrayn, Tunisia, 11.x.2006 (coll. B. Müller) 110d. C. melanaria Obth., ♀, Fiumara Trionto, Calábria, Italy, 15.x.1999 (ZSM) 110e. C. melanaria Obth., ♀, Francavilla in Sinni, Basilicata, Italy, 10.x.1982 (coll. Skou) 111a. Paradarisa consonaria Hbn., ♂, Klueß, Güstrow, Mecklenburg-Vorpommern, Germany, 16.v.1980 (coll. B. Müller) 111b. P. consonaria Hbn., ♂, Martinroda, Thüringen, Germany, 30.iv.1971 (coll. B. Müller) 111c. P. consonaria Hbn., ♀, Martinroda, Thüringen, Germany, 10.v.1969 (coll. B. Müller) 111d. P. consonaria Hbn., ♀, Martinroda, Thüringen, Germany, 2.v.1972 (coll. B. Müller) 112a. Larerannis orthogrammaria Whli., ♂, [Russia], 1.x.2006 (coll. ZISP) 112b. L. orthogrammaria Whli., ♂, Izmailov Park, Moscow, Russia, 1.x.2002 (ZISP) 112c. L. orthogrammaria Whli., ♂, Sad-Gorod, Vladivostok, Russia, 23.x.1996 (coll. Skou) 112d. L. orthogrammaria Whli., ♀, Izmajlovskij Park, Moscow, Russia, 3.x.2005 (ZISP), 113a. Chemerina caliginearia Rmb., ♂, Plage de la Marana, Corsica, France, 27.iii.2002 (coll. Skou) 113b. C. caliginearia Rmb., ♂, Gordes, Vaucluse, France, 1.i.1973 (ZSM) 113c. C. caliginearia Rmb., ♀, Plage de la Marana, Corsica, France, 27.iii.2002 (coll. Skou) 113d. C. caliginearia Rmb., ♀, Roche, Prov. Cadiz, Spain, 6.iii.2005 (coll. Skou) 114a. Fagivorina arenaria Hufn., ♂, Pisoderi Ski Resort, Macedonia, Greece, 4.vii.2016 (coll. Skou) 114b. F. arenaria Hufn., ♂, Heringsdorf, Mecklenburg-Vorpommern, Germany, 5.vii.1958, (MNHU) 114c. F. arenaria Hufn., ♀, Heringsdorf, Mecklenburg-Vorpommern, Germany, 11.vii.1956, (MNHU) 114d. F. arenaria Hufn., ♀, Göhrde, Niedersachsen, Germany, 3.vii.1987 (coll. Wegner) 115a. Spartopteryx kindermannaria Stgr., ♂, Nizhnij Tsasutchej, Dahuria, Russia, 24.vi.1988 (coll. B. Müller) 115b. S. kindermannaria Stgr., ♂, Ortatau Mountains, Dshungarian Alatau, Kazakhstan, 9.vi.1997 (coll. Skou) 115c. S. kindermannaria Stgr., ♂, Aktash, Altai Mountains, Russia, 13.vi.2015 (coll. B. Müller) 115d. S. kindermannaria Stgr., ♀, Minussinsk, Krasnojarsk, Russia, 17.vi.1933 (ZFMK) 115e. S. kindermannaria Stgr., ♀, Kuku-Noor, Tibet, vi.1898 (MNHU) 116a. Athroolopha pennigeraria Hbn., ♂, Albarracin, Prov. Teruel, Spain, 8.vi.2007 (coll. B. Müller) 116b. A. pennigeraria Hbn., ♂, Calomarde, Prov. Teruel, Spain, 9.vi.2007 (coll. B. Müller) 116c. A. pennigeraria Hbn., ♂, Albarracin, Prov. Teruel, Spain, 27.vi.1992 (coll. Skou) 116d. A. pennigeraria Hbn., ♀, Sierra de los Filabres, Prov. Almeria, Spain, e. l. 26.iv.2004 (coll. B. Müller) 117a. Athroolopha chrysitaria Hbn., ♂, Palermo, Sicily, Italy, 20.vi.1905 (MNHU), dissected 117b. A. chrysitaria Hbn., ♂, Palermo, Sicily, Italy (MNHU) 117c. A. chrysitaria Hbn., ♀, Alpe Cucco, Ficuzza, Sicily, Italy, 9.vi.1990 (coll. Skou) 117d. A. chrysitaria Hbn., ♀, Sicily, Italy (MNHU)

600

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PLATE 18 (all specimens natural size) 118a. Eurranthis plummistaria Vill., ♂, Digne, Alpes-de-Haute-Provence, France, 18.–19.iv.1992 (coll.

B. Müller) 118b. E. plummistaria Vill., ♂, Chemin de La Sarrée, Alpes Maritimes, France, 9.iv.1930 (ZSM) 118c. E. plummistaria Vill., ♀, Chemin de La Sarrée, Alpes Maritimes, France, 25.iv.1929 (ZSM) 118d. E. plummistaria Vill., ♀, Lucainena de la Torres, Prov. Almeria, Spain, 23.iii.2003 (coll. B. Müller) 119a. Nychiodes obscuraria Vill., ♂, Mèounes, Var, France, 3.vii.1967 (ZSM) 119b. N. obscuraria Vill., ♂,Latsch, South Tyrol, Italy, e. l. 19vii.1983 (coll. B. Müller), dissected 119c. N. obscuraria Vill., ♂, Bidsca, Tessin, Switzerland, (ZSM), dissected 119d. N. obscuraria Vill., ♂, Eisak Valley, South Tyrol, Italy (ZSM), dissected 119e. N. obscuraria Vill., ♂, Monti Simbruini, Lazio, Italy, 21.vi.1927 (ZSM) 119f. N. obscuraria Vill., ♀, Latsch, South Tyrol, Italy, e. p. 16.vii.1976 (coll. B. Müller), dissected 120a. Nychiodes ragusaria Mill., ♂, Cività, Monte Pollino, Calabria, Italy, 31.v.–1.vi.2005 (coll. Skou), dissected 120b. N. ragusaria Mill., ♂, Ski Centre, Monte Sirino, Basilicata, Italy, 29.vii.2011 (coll. Skou) 120c. N. ragusaria Mill., ♂, Coccorino, Calabria, Italy, e. o. 15.vi.2008 (ZSM), dissected 120d. N. ragusaria Mill., ♂, Sicily, Italy (MNHN) 120e. N. ragusaria Mill., ♂, Civita, Monte Pollino, Calabria, Italy, 31.v.–1.vi.2005 (coll. Skou) 120f. N. ragusaria Mill., ♀, Coccorino, Calabria, Italy, e. o. 15.vi.2008 (coll. B. Müller), dissected 121a. Nychiodes andalusiaria Mill., ♂, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 21.vii.2003 (coll. Skou), dissected 121b. N. andalusiaria Mill., ♂, Serra da Estrela, Beira, Portugal (ZSM), dissected 121c. N. andalusiaria Mill., ♂, Sierra Estrela, ‘Spain’, e. l. 20.x.1931 (ZSM), dissected 121d. N. andalusiaria Mill., ♂, Serra da Estrela, Beira, Portugal (ZSM), dissected 121e. N. andalusiaria Mill., ♂, Serra da Estrela, Beira, Portugal (ZSM), dissected 121f. N. andalusiaria Mill., ♂, underside, La Plataforma, Sierra de Gredos, Prov. Avila, Spain, 21.vii.2003 (coll. Skou), dissected

602

603

PLATE 19 (all specimens natural size) 122a. Nychiodes notarioi Exp., ♂, Sierra del Montseny, Prov. Girona, Spain, e. o. 8.vii.1993 (coll. B. Müller),

dissected 122b. N. notarioi Exp., ♂, Col del Canto, Prov. Lerida, Spain, 16.vii.2003 (coll. Skou), dissected 122c. N. notarioi Exp., ♂, Sierra de Javalambre, Prov. Teruel, Spain, 25.vii.1978 (ZSM) 122d. N. notarioi Exp., ♂, Uña, Prov. Cuenca, Spain, 8.–9.vii.2002 (coll. Skou), barcoded, dissected 122e. N. notarioi Exp., ♂, Salvacanete, Prov. Cuenca, Spain, 7.vii.2014 (ZSM) 122f. N. notarioi Exp., ♀, Noguera, Prov. Teruel, Spain, e. o. 8.x.1990 (ZSM), dissected 122g. N. notarioi Exp., ♀, Sierra del Montseny, Prov. Girona, Spain, e. o. 8.vii.1993 (coll. B. Müller), dissected 122h. N. notarioi Exp., ♀, Albarracin, Prov. Teruel, Spain, e. o. vi.1953, (ZSM), dissected 122i. N. notarioi Exp., ♀, Sort, Prov. Lerida, Spain, 7.vii.1993 (coll. Skou) 123a. Nychiodes hispanica Whli., ♂, Sierra Segura, Prov. Granada, Spain 27.vii.1979 (SMNS), dissected 123b. N. hispanica Whli., ♂, Puerto de la Ragua, Prov. Granada, Spain, e. o. 12.vii.2009 (coll. B. Müller), dissected 123c. N. hispanica Whli., ♂, Algatocin, Prov. Malaga, Spain, e. o. 28.vii.1998 (coll. B. Müller), dissected 123d. N. hispanica Whli., ♂, Algatocin, Prov. Malaga, Spain, e. o. 28.vii.1998 (coll. B. Müller), barcoded, dissected 123e. N. hispanica Whli., ♂, Ifrane, Middle Atlas, Morocco, e. o. 30.vi.1994 (coll. B. Müller), dissected 123f. N. hispanica Whli., ♂, Road to Veleta, Sierra Nevada, Prov. Granada, Spain, 20.vi.1960 (ZSM), dissected 123g. N. hispanica Whli., ♂, Otivar, Prov. Granada, Spain, 24.vii.2003 (coll. Skou), dissected 123h. N. hispanica Whli., ♂, Prov. Granada, Spain (MNHU), dissected 123i. N. hispanica Whli., ♂, Puerto de la Ragua, Prov. Granada, Spain, e. o. 16.vii.2009 (coll. B. Müller), dissected 123j. N. hispanica Whli., ♂, Almoraima, Prov. Cadiz, Spain, 24.ix.1987 (coll. Skou), dissected 123k. N. hispanica Whli., ♀, Algatocin,Prov. Malaga, Spain, e. o. 28.vii.1998 (coll. Leipnitz) 123l. N. hispanica Whli., ♀, Sierra Nevada, Prov. Granada, Spain, 6.vii.1926 (ZFMK), lectotype

604

605

PLATE 20 (all specimens natural size) 124a. Nychiodes waltheri Wagn., ♂, Studen Kladenets, Rhodopi Mountains, Bulgaria, 23.v.1990 (coll. Skou),

barcoded, dissected 124b. N. waltheri Wagn., ♀, Madzhorovo, Ostrhodope, Bulgaria, e. o. 17.–24.v.2004 (ZSM), dissected 124c. N. waltheri Wagn., ♂, Sozopol, Black Sea Coast, Bulgaria, e. o. 16.xi.1987 (coll. B. Müller) 124d. N. waltheri Wagn., ♂, Varagöz, Sat Dağları, Hakkari, Turkey, 21.–24.vii.1983 (SMNS), dissected 124e. N. waltheri Wagn, ♂, Marasch, Taurus Mountains, Turkey, e. o. 29.vii.1928 (ZSM) 124f. N. waltheri Wagn, ♀, Marasch, Taurus Mountains, Turkey, 21.vii.1929 (ZSM) 125a. Nychiodes amygdalaria H.-S., ♂, Aksehir, Konya, Turkey, 20.vii.–23.viii.1967 (SMNS) 125b N. amygdalaria H.-S., ♂, Liljanovo, Pirin Mountains, Bulgaria, 16.vi.1981 (coll. B. Müller) 125c. N. amygdalaria H.-S., ♀, Altıparmak, Artvin, Turkey, 31.vii.–3.viii.1983 (SMNS) 125d. N. amygdalaria H.-S., ♀, Esimi, Thrakia, Greece, 20.ix.2017 (coll. Gelbrecht) 125e. N. amygdalaria H.-S., ♂, Tecde, Malatya, Turkey, ix.1917 (ZSM) 125f. N. amygdalaria H.-S., ♂, Süldüf Dagh, Amanus Mountains, Hatay, 30.viii. (ZSM) 126a. Nychiodes dalmatina Wagn., ♂, Liljanovo, Pirin Mountains, Bulgaria, e. o. 14.viii.1981 (coll. B. Müller), barcoded, dissected 126b. N. dalmatina Wagn., ♂, Aghiokambos, Thessaly, Greece, 31.v-1.vi.2013 (coll. Skou), dissected 126c. N. dalmatina Wagn., ♂, Monemvasia, Peloponnese, Greece, 30.vi.–3.vii.1985 (coll. Skou), dissected 126d. N. dalmatina Wagn., ♂, Monastir, Serbia [without date] (ZSM) 126e. N. dalmatina Wagn., ♂, Chelmos, Peloponnese, Greece, e. o. 22.viii.1995 (ZSM), dissected 126f. N. dalmatina Wagn., ♂, Dubrovnik, Croatia, (ZSM), dissected 126g. N. dalmatina Wagn., ♂, Zadar, Dalmatia Croatia, 17.vi.1975 (coll. Skou) 126h. N. dalmatina Wagn., ♀, Liljanovo, Pirin Mountains, Bulgaria, e. o. 21.viii.1981 (coll. B. Müller), barcoded, dissected 126i. N. dalmatina Wagn., ♀, by river Sarandaporos, Aghia Paraskevi, Epirus, Greece, 3.vi.2010 (coll. Skou)

606

607

PLATE 21 (all specimens natural size) 127a. Phaselia serrularia Ev., ♂, Pokrovka village, South Ural, Russia, 21.vi.1999 (coll. Skou), dissected 127b. P. serrularia Ev., ♂, Sarepta, Volgograd, Russia, vii.2009 (ZSM) 127c. P. serrularia Ev., ♂, Sarepta, Volgograd, Russia (MNHU), dissected 127d. P. serrularia Ev., ♀, Krasnodar, Adygeja, Russia, 1888 (MNHU), dissected 128a. Phaselia algiricaria aragona Whli., ♂, Albarracin, Prov. Teruel, Spain, 11.vi.2007 (coll. B. Müller) 128b. P. algiricaria aragona Whli., ♂, Turillas, Prov. Almeria, Spain, 28.v.2006 (coll. Skou) 128c. P. algiricaria gigantaria Schwingens., ♂, Aqesri, High Atlas, Morocco, 17.iv.2012 (coll. B. Müller) 128d. P. algiricaria aragona Whli., ♀, El Pozo del Esparto, Prov. Almeria, Spain, 27.v.2006 (coll. Skou) 129a. Ekboarmia atlanticaria Stgr., ♂, Punta Paloma, Prov. Cádiz, Spain, 22.–24.iv.2007 (coll. Skou) 129b. E. atlanticaria Stgr., ♂, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.iii.–3.iv.2016 (coll. Skou) 129c. E. atlanticaria Stgr., ♂, Salema Camping, Algarve, Portugal, 22.iv.2017 (coll. Skou) 129d. E. atlanticaria Stgr., ♀, Punta Paloma, Prov. Cádiz, Spain, 12.–13.iv.2007 (coll. Skou) 129e. E. atlanticaria Stgr., ♀, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.iii.–3.iv.2016 (coll. Skou) 130a. Ekboarmia sagnesi Dufay, ♂, Romanche, Usine, Hautes-Alpes, France, e. l. 15.viii.2009 (coll. Tautel) 130b. E. sagnesi Dufay, ♂, Le Fréaux, Hautes-Alpes, France, 20.vi.1979 (coll. Tautel), paratype 130c. E. sagnesi Dufay, ♂, Valle de Luna, Prov. Leon, Spain, 22.vii.2011 (coll. Skou), dissected 130d. E. sagnesi Dufay, ♀, Valle de Luna, Prov. Leon, Spain, 22.vii.2011 (coll. Skou) 130e. E. sagnesi Dufay, ♀, Romanche, Usine, Hautes-Alpes, France, 24.vi.2009 (coll. Tautel) 131a. Ekboarmia miniaria Skou et al., ♂, Ameiras de Baixo, Grandola, Portugal, 20.vi.2011 (coll. Skou),

paratype, dissected 131b. E. miniaria Skou et al., ♂, Lagoa de Santo André, Prov. Alentejo, Portugal, 16.ix.–10.x.1995 (coll. Skou), paratype, dissected 131c. E. miniaria Skou et al., ♂, Ameiras de Baixo, Grandola, Portugal, 20.vi.2011 (coll. Skou), holotype, dissected 131d. E. miniaria Skou et al., ♀, Ameiras de Baixo, Grandola, Baixo Alentejo, Portugal, 15.vi.2009 (ZFMK), para­ type, dissected 131e. E. miniaria Skou et al., ♀, Ameiras, de Baixo, Grandola, Baixo Alentejo, Portugal, 15.vi.2009 (coll. Skou), paratype, dissected 132a. Ecleora solieraria Rmb., ♂, Albarracin, Prov. Teruel, Spain, 6.vi.2007 (coll. B. Müller) 132b. E. solieraria Rmb., ♂, Albarracin, Prov. Teruel, Spain, 11.vi.2007 (coll. B. Müller) 132c. E. solieraria Rmb., ♂, Albarracin, Prov. Teruel, Spain, 4.ix.2008 (coll. B. Müller) 132d. E. solieraria Rmb., ♀, Denia, Prov. Alicante, Spain, e. o. 16.vi.2002 (coll. B. Müller) 132e. E. solieraria Rmb., ♀, Albarracin, Prov. Teruel, Spain, 18.vi.2007 (coll. B. Müller) 133a. Afriberina tenietaria Stgr., ♂, Albarracin, Prov. Teruel, Spain, 7.viii.2007 (coll. Skou), dissected 133b. A. tenietaria Stgr., ♂, Albarracin, Prov. Teruel, Spain, 25.–26.vi.1992 (coll. Skou) 133c. A. tenietaria Stgr., ♂, La Toba, Prov. Cuenca, Spain, 27.vi.2006 (coll. Skou) 133d. A. tenietaria Stgr., ♀, Albarracin, Prov. Teruel, Spain, 27.vi.1992 (coll. Skou) 133e. A. tenietaria Stgr., ♂, near Savournon, Hautes-Alpes, France, 9.vii.2016 (coll. Colomb) 133f. A. tenietaria Stgr., ♀, near Savournon, Hautes-Alpes, France, 9.vii.2016 (coll. Colomb) 133g. A. tenietaria Stgr., ♂, Playa de Rompeculos, Prov. Huelva, Spain, 20.iv.2007 (coll. Skou) 133h. A. tenietaria Stgr., ♂, Matalascañas, Prov. Huelva, Spain, 16.iv.2007 (coll. Skou) 133i. A. tenietaria Stgr., ♀, Matalascañas, Prov. Huelva, Spain, 18.iv.2017 (coll. Skou) 133j. A. tenietaria Stgr., ♀, Matalascañas, Prov. Huelva, Spain, 18.iv.2017 (coll. Skou) 134a. Afriberina salemae Skou & Sihv., ♂, Salema Camping, Algarve, Portugal, 21.iv.2017 (coll. Skou), holo­ type 134b. A. salemae Skou & Sihv., ♂, Salema Camping, Algarve, Portugal, 19.iv.2007 (coll. Skou), paratype 134c. A. salemae Skou & Sihv., ♂, Salema Camping, Algarve, Portugal, 22.iv.2017 (coll. Skou), paratype 134d. A. salemae Skou & Sihv., ♀, Salema Camping, Algarve, Portugal, 22.iv.2017 (coll. Skou), paratype 134e. A. salemae Skou & Sihv., ♀, Salema Camping, Algarve, Portugal, 22.iv.2017 (coll. Skou), paratype, dissected

608

609

PLATE 22 (all specimens natural size) 135a. Sardocyrnia bastelicaria Bell., ♂, Ghisoni, Corsica, France, 1.vi.2008 (coll. B. Müller), barcoded 135b. S. bastelicaria Bell., ♂, Bois de Barbaggio, Corsica, France, vii.1923 (ZSM) 135c. S. bastelicaria Bell., ♂, Corte, Corsica, France, e. o. early vi.1970 (MNHU) 135d. S. bastelicaria Bell., ♂, Bonifacia, Corsica, France, 9.–10.v.2004 (coll. Skou) 135e. S. bastelicaria Bell., ♀, Castirla, Corsica, France, 26.ix.2003 (coll. Skou), dissected 136a. Sardocyrnia fortunaria Vazq., ♂, Albarracin, Prov. Teruel, Spain, 1.vi.2007 (coll. B. Müller) 136b. S. fortunaria Vazq., ♂, Rodalquilar, Prov. Almeria, Spain, 26.iii.2003 (coll. B. Müller), barcoded 136c. S. fortunaria Vazq., ♀, Albarracin, Prov. Teruel, Spain, 10.vi.2007 (coll. B. Müller) 136d. S. fortunaria Vazq., ♀, Albarracin, Prov. Teruel, Spain, 3.vi.2007 (coll. B. Müller) 136e. S. fortunaria Vazq., ♀, Rodalquilar, Prov. Almeria, Spain, 25.iii.2003 (coll. B. Müller) 137a. Selidosema brunnearia Vill., ♂, Monte Paradiso, L’Aquila, Italy (ZFMK) 137b. S. brunnearia Vill., ♂, Madonie, Sicily, Italy (ZFMK) 137c. S. brunnearia Vill., ♂, Hoyerswerda, Sachsen, Germany, 10.viii.1985 (coll. B. Müller) 137d. S. brunnearia Vill., ♂, Vicklerby, Öland, Sweden, 23.vii.1971 (coll. Skou) 137e. S. brunnearia Vill., ♂, Möckelmossen, Öland, Sweden, 2.viii.1973 (coll. Skou) 137f. S. brunnearia Vill., ♀, Madonie, Sicily, Italy (ZFMK) 137g. S. brunnearia Vill., ♀, Bornholm, Denmark, e. o. 22.vii.1990 (coll. B. Müller) 137h. S. brunnearia Vill., ♀, Staakow, Brandenburg, Germany, e. o. 16.viii.1991 (coll. B. Müller) 138a. Selidosema plumaria Den. & Schiff., ♂, Budaörs, Pest, Hungary, e. o. 26.v.1984 (coll. B. Müller) 138b. S. plumaria Den. & Schiff., ♂, Geroplatanos, Ioánina, Greece, 31.viii.1983 (coll. Skou) 138c. S. plumaria Den. & Schiff., ♂, Garda, Verona, Italy, 2.–5.x.1983 (coll. Skou) 138d. S. plumaria Den. & Schiff., ♀, Katara Pass, Ioánina, Greece, 23.viii.2008 (coll. Skou) 138e. S. plumaria Den. & Schiff., ♀, Puerto de Orihuela, Prov. Teruel, Spain, 23.viii.2001 (coll. Skou) 138f. S. plumaria Den. & Schiff., ♀, Hundsheim, Niederösterreich, Austria, 29.viii.1975 (coll. Skou) 138g. S. plumaria Den. & Schiff., ♀, Budaörs, Pest, Hungary, 30.viii.1984 (coll. B. Müller) 138h. S. plumaria Den. & Schiff., ♀, Aghia Paraskevi, Ioánina, Greece, 26.viii.2008 (coll. Skou) 139a. Selidosema pyrenaearia Bsd., ♂, Montalegre, Trás-os-Montes, Portugal (ZSM) 139b. S. pyrenaearia Bsd., ♂, Granada, Prov. Granada, Spain, (MNHU) 139c. S. pyrenaearia Bsd., ♀, Lagoa de Santo André, Alentejo, Portugal, 24.ix.2009 (coll. Skou), barcoded 139d. S. pyrenaearia Bsd., ♀, Lagoa de Santo André, Alentejo, Portugal, 24.ix.2009 (coll. Skou), barcoded 140a. Selidosema taeniolaria Hbn., ♂, Albarracin, Prov. Teruel, Spain, 9.ix.2008 (coll. B. Müller) 140b. S. taeniolaria Hbn., ♀, Albarracin, Prov. Teruel, Spain, 9.ix.2008 (coll. B. Müller) 140c. S. taeniolaria Hbn., ♀, Barranco de Valcuerna, Prov. Huesca, Spain, larva 15.iv.2001 (coll. Skou) 140d. S. taeniolaria Hbn., ♀, San Marti de Tous, Prov. Barcelona, Spain, 16.viii.2001 (coll. Skou) 141a. Selidosema ambustaria Geyer, ♂, Agrigento, Sicily, Italy e. o. 29.vi.1910 (MNHU), 141b. S. ambustaria Geyer, ♂, Ficuzza, Sicily, Italy, viii.1904 (MNHU) 141c. S. ambustaria Geyer, ♂, Madonie, Sicily, Italy (ZSM) 141d. S. ambustaria Geyer, ♀, Madonie, Sicily, Italy (ZSM)

610

611

PLATE 23 (all specimens natural size) 142a. Selidosema erebaria postdentaria Roths., ♂, Montagna Grande, Pantelleria, Italy, 8.–13.x.2007 (coll. Fischer) 142b. S. erebaria postdentaria Roths., ♂, Montagna Grande, Pantelleria, Italy, 8.–13.x.2007 (coll. Fischer) 142c. S. erebaria postdentaria Roths., ♂, Montagna Grande, Pantelleria, Italy, 8.–13.x.2007 (coll. Fischer) 142d. S. erebaria postdentaria Roths., ♀, Montagna Grande, Pantelleria, Italy, 8.–13.x.2007 (coll. Fischer) 143a. Peribatodes rhomboidaria Den. & Schiff., ♂, Rosenthal, Berlin, Germany, 27.v.2002, (coll. B. Müller), dissected 143b. P. rhomboidaria Den. & Schiff., ♂, Bad Mergentheim, Baden-Württemberg, Germany, 30.vi.1956 (SMNS) 143c. P. rhomboidaria Den. & Schiff., ♂, Tréccina, Basilicata, Italy, 17.x.2017 (coll. Skou) 143d. P. rhomboidaria Den. & Schiff., ♂, Odense, Fyn, Denmark, 21.vii.1971 (coll. Skou) 143e. P. rhomboidaria Den. & Schiff., ♀, Fluminimaggiore, Sardinia, Italy, 7.–8.iv.2002 (coll. Skou) 143f. P. rhomboidaria Den. & Schiff., ♀, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.iii-3.iv.2016 (coll. Skou) 143g. P. rhomboidaria Den. & Schiff., ♀, Lake Garda, Verona, Italy, e. l. 10.v.1980 (SMNS) 143h. P. rhomboidaria Den. & Schiff., ♂, Marasch, ’Syria’ [Turkey], 26.v.–15.vi.1929 (ZFMK) 144a. Peribatodes perversaria Bsd., ♂, Visperterminen, Valais, Switzerland, e. o. 15.vi.1995 (coll. B. Müller), dissected 144b. P. perversaria Bsd., ♂, Norante, Haute Provence, France, e. o. 21.vii.2002 (coll. B. Müller), dissected 144c. P. perversaria Bsd., ♂, Gèdre, Haute Pyrénées, France, vii.1898 (ZSM), dissected 144d. P. perversaria Bsd., ♂, Barbaira, Aude, France, 10.ix.1988 (ZSM), dissected 144e. P. perversaria Bsd., ♂, Vallee du Miroux, Digne, 13.ix.1969 (ZSM), dissected 144f. P. perversaria Bsd., ♂, Moscardon, Prov. Teruel, Spain, 30.vii.1973 (SMNS), dissected 144g. P. perversaria Bsd., ♂, Porté, Pyrénées-Orientales, France (ZSM), dissected 144h. P. perversaria Bsd., ♀, Aragon, Spain, 18.–31.viii.1927 (ZSM), dissected 145a. Peribatodes omalensis Rajaei et al., ♂, Omalos Plateau, Crete, Greece, 18.vi.2014 (coll. Skou), paratype, barcoded, dissected 145b. P. omalensis Rajaei et al., ♀, Nea Roumata, Crete, Greece, 9.vi.1998 (ZSM), holotype, barcoded 145c. P. omalensis Rajaei et al., ♀, Omalos Plateau, Crete, Greece, 27.vi.2012 (SMNS), paratype, barcoded, dissected 145d. P. omalensis Rajaei et al., ♀, Omalos Plateau, Crete, Greece, 15.–20.vi.2014 (coll. Skou), paratype barcoded, dissected 146a. Peribatodes umbraria Hbn., ♂, Ghisoni, Corsica, France, 2.vi.2008 (coll. B. Müller), dissected 146b. P. umbraria Hbn., ♀, Karadagh, Crimea, Ukraine, 3.vi.1985 (ZMKU) 146c. P. umbraria Hbn., ♂, Lambèse, Algeria, viii.1912 (ZFMK) 146d. P. umbraria Hbn., ♀, Mischliffen, Middle Atlas, Morocco, 17.vii.1998 (coll. B. Müller), dissected 146e. P. umbraria Hbn., ♂, Palermo, Sicily, Italy (ZFMK) 146f. P. umbraria Hbn., ♂, Kasrun, Iran, late iv.1938 (ZFMK), dissected

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PLATE 24 (all specimens natural size) 147a. Peribatodes ebusaria Vallh., ♂, Sant Llorenc de Bàlàfia, Ibiza, Spain, 4.iv.2016 (coll. Skou) 147b. P. ebusaria Vallh., ♂, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.iii.–3.iv2016 (coll. Skou), dissected 147c. P. ebusaria Vallh., ♂, Coll de sa Creu, Ibiza, Spain, 1.x.2018 (coll. Skou) 147d. P. ebusaria Vallh., ♀, Puig de Fornàs, Sant Rafel, Ibiza, Spain, 30.ix.2018 (coll. Skou), dissected 147e. P. ebusaria Vallh., ♀, Cala d’Hort, Ibiza, Spain, 3.–4.x.2018 (coll. Skou) 148a. Peribatodes secundaria Den. & Schiff., ♂, Weinheim, Baden-Württemberg, Germany, 8.vii.1979

(SMNS) 148b. P. secundaria Den. & Schiff., ♂, Katzenbacher-Hof, Baden-Württemberg, Germany, 27.vii.1962 (SMNS) 148c. P. secundaria Den. & Schiff., ♂, Saualpe, Kärnten, Austria, 6.vii.2011 (coll. B. Müller), dissected 148d. P. secundaria Den. & Schiff., ♂, Agabeyli, Maras, Turkey, 27.–28.v.1981 (SMNS) 148e. P. secundaria Den. & Schiff., ♂, Zastler, Baden-Württemberg, Germany, 1.viii.1969 (SMNS) 148f. P. secundaria Den. & Schiff., ♀, Neuhütten, Baden-Württemberg, Germany, 2.vii.1976, (SMNS) 148g. P. secundaria Den. & Schiff., ♀, Saualpe, Kärnten, Austria, 9.vii.2011 (coll. B. Müller), dissected 148h. P. secundaria Den. & Schiff., ♀, Marash, Taurus Mountains, Turkey, 19.v.1928 (ZSM), dissected 149a. Peribatodes correptaria Zell., ♂, Agios Nikolaos, Crete, Greece, e. o. late v.1993 (coll. B. Müller), dissected 149b. P. correptaria Zell., ♂, Amasya, Turkey, 8.–9.vi.1980 (SMNS) 149c. P. correptaria Zell., ♂, Akdeniz, Cyprus, 3.iv.2008, (coll. Skou) 149d. P. correptaria Zell., ♂, Pirgos, Peloponnese, Greece, 13.x.1984 (coll. Skou) 149e. P. correptaria Zell., ♀, Dalmatia, Croatia [without dates] (ZSM) 149f. P. correptaria Zell., ♀, Agios Nikolaos, Crete, Greece, e. o. mid v.1993 (coll. B. Müller), dissected 149g. P. correptaria Zell., ♀, Batiliman, Crimea, Ukraine, 26.ix.2002 (ZMKU) 149h. P. correptaria Zell., ♀, Omalos, Crete, Greece, 26.vi.2012 (coll. Skou) 150a. Peribatodes aragonis Whli., ♂, Albarracin, Prov. Teruel, Spain, e. o. 4.ix.2008 (coll. B. Müller), dissected 150b. P. aragonis Whli., ♂, Ifrane, Middle Atlas, Morocco, 16.–31.vii.1953 (MNHN) 150c. P. aragonis Whli., ♀, Albarracin, Prov. Teruel, Spain, 5.viii.1930 (ZFMK), dissected 150d. P. aragonis Whli., ♀, Albarracin, Prov. Teruel, Spain, 5.ix.2008 (coll. B. Müller), dissected 151a. Peribatodes ilicaria Geyer, ♂, Aledo, Prov. Murcia, Spain, e. o. 4.vi.2003 (coll. B. Müller), dissected 151b. P. ilicaria Geyer, ♂, Oraison, Alpes-de-Haute-Provence, France, 8.vi.1967 (ZSM) 151c. P. ilicaria Geyer, ♀, Mazagón, Prov. Huelva, Spain, e. o. 15.x.2005 (coll. B. Müller), dissected 151d. P. ilicaria Geyer, ♀, Aledo, Prov. Murcia, Spain, e. o. 4.vi.2003 (coll. B. Müller), dissected 151e. P. ilicaria Geyer, ♀, La-Garde-Freinet, Var, France, 13.–14.ix.1987 (ZSM), dissected 151f. P. ilicaria Geyer, ♂, Prov. Cuenca, Spain, 1906 (ZSM), dissected 151g. P. ilicaria Geyer, ♀, Albarracin, Prov. Teruel, Spain, 30.vii.1928 (ZSM), dissected 151h. P. ilicaria Geyer, ♂, Kenitra, Morocco, 10.vi.1976 (SMNS)

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PLATE 25 (all specimens natural size) 152a. Alcis repandata L., ♂, Reschen, South Tyrol, Italy, 5.vii.2002 (coll. B. Müller) 152b. A. repandata L., ♂, Degioz, Aosta, Italy, 15.vii.1999 (coll. B. Müller), barcoded 152c. A. repandata L., ♂, Svanninge Bakker, Fyn, Denmark, 25.vi.1971 (coll. Skou) 152d. A. repandata L., ♂, Schnalstal, Trentino-Alto Adige, Italy, 18.viii.1996 (coll. Skou) 152e. A. repandata L., ♀, Vradeto, Epirus, Greece, 9.–10.vii.2016 (coll. Skou) 152f. A. repandata L., ♀, Wilhelmshagen, Berlin, Germany, 1.vii.1970 (coll. B. Müller) 152g. A. repandata L., ♀, Ilmenau, Thüringen, Germany, e. l. 10.vi.1973 (coll. B. Müller) 152h. A. repandata L., ♀, Stade, Niedersachsen, Germany, 11.vii.1985 (coll. Wegner) 152i. A. repandata L., ♀, Slotslyngen, Bornholm, Denmark, 11.vii.1981 (coll. Skou) 153a. Alcis deversata Stgr., ♂, Mnichova-Lehota,Trencin, Slovak Republic, e. o. 3.iv.196 (coll. B. Müller) 153b. A. deversata Stgr., ♂, Sauerland, Nordrhein-Westfalen, Germany, e. o. 27.iv.1982 (coll. B. Müller) 153c. A. deversata Stgr., ♀, Sauerland, Nordrhein-Westfalen, Germany, e. o. 22.iv.1982 (coll. B. Müller) 153d. A. deversata Stgr., ♀, Mnichova-Lehota,Trencin, Slovak Republic, e. o. 3.iv.1986 (coll. B. Müller) 154a. Alcis jubata Thnbg., ♂, Märkisch Buchholz, Brandenburg, Germany, e. l. 12.vii.1902 (MNHU) 154b. A. jubata Thnbg., ♂, Zanadvorovska, Ussuri, Russia, viii.1991 (coll. Skou) 154c. A. jubata Thnbg., ♂, Saualpe, Kärnten, Austria, 9.vii.2011 (coll. B. Müller) 154d. A. jubata Thnbg., ♀, Åminsby, Uusimaa, Finland, 4.viii.1953 (MNHU) 154e. A. jubata Thnbg., ♀, Kamenz, Sachsen, Germany e. l. vii.1901 (MNHU) 155a. Deileptenia ribeata Cl., ♂, Schortetal, Thüringen, Germany, 16.vii.1969 (coll. B. Müller) 155b. D. ribeata Cl., ♂, Sauwald, Sachsen, Germany, 3.vii.2014 (coll. B. Müller) 155c. D. ribeata Cl., ♂, Coll del Portillón, Prov. Catalonia, Spain, 22.vii.2014 (coll. Skou) 155d. D. ribeata Cl., ♀, Uutela, Uusima, Finland, 6.–10.vii.2002 (coll. Skou) 155e. D. ribeata Cl., ♀, Coll del Portillón, Prov. Catalonia, Spain, 22.vii.2014 (coll. Skou) 155f. D. ribeata Cl., ♀, Manebach, Ilmenau, Thüringen, Germany, e. o. 3.viii.1972 (coll. B. Müller) 156a. Paraboarmia viertlii Boh., ♂, Valtice, Moravia, Czech Republic, 15.vii.1981 (coll. B. Müller) 156b. P. viertlii Boh., ♂, Hungary (ZFMK) 156c. P. viertlii Boh., ♂, Valtice, Moravia, Czech Republic, 15.vii.1981 (coll. Skou) 156d. P. viertlii Boh., ♂, Fayence, Var, France, 2.–7.vii.1987 (coll. Skou) 156e. P. viertlii Boh., ♀, Hungary (ZFMK)

616

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PLATE 26 (all specimens natural size) 157a. Adactylotis gesticularia Hbn., ♂, Eslida, Prov. Castellon de la Plana, Spain, 1.v.1997 (coll. Skou) 157b. A. gesticularia Hbn., ♂, Moscardón, Prov. Teruel, Spain, 17.vi.2007 (coll. B. Müller) 157c. A. gesticularia Hbn., ♂, Lagoa de Santo André, Prov. Alentejo, Portugal, 24.ix.2009 (coll. Skou) 157d. A. gesticularia Hbn., ♂, Ruta del Veleta, Sierra Nevada, Prov. Granada, Spain, 19.ix.1987 (coll. Skou) 157e. A. gesticularia Hbn., ♂, Embid, Prov. Guadalajara, Spain, 8.viii.2007 (coll. Skou), 157f. A. gesticularia Hbn., ♂, El Curaton, Prov. Cádiz, Spain, 22.–24.iv.2007 (coll. Skou) 157g. A. gesticularia Hbn., ♀, Besalu, Prov. Girona, Spain, l4.vii.1995 (coll. Skou) 157h. A. gesticularia Hbn., ♀, Lagoa de Santo André, Prov. Alentejo, Portugal, 24.ix.2009 (coll. Skou) 157i. A. gesticularia Hbn., ♀, El Curaton, Prov. Cádiz, Spain, 22.–24.iv.2007 (coll. Skou) 158a. Adactylotis contaminaria Hbn., ♂, Forêt de Chizè, Deux-Sèvres, France, 27.v.1929 (ZSM) 158b. A. contaminaria Hbn., ♂, Corsavu, Pyrénées Orientales, France, 26.iii.2002 (coll. Skou) 158c. A. contaminaria Hbn., ♂, Pruit, Prov. Catalonia, Spain, 8.vii.1993 (coll. B. Müller) 158d. A. contaminaria Hbn., ♀, Presles, Val-d’Oise, France, 18.v.1944 (ZSM) 159a. Pseudocoremia suavis Btl., ♂, Godolphin Cross, Cornwall, United Kingdom, 27.ix.2016 (coll. Clarke) 159b. P. suavis Btl., ♂, Godolphin Cross, Cornwall, United Kingdom, 27.ix.2016 (coll. Clarke) 159c. P. suavis Btl., ♀, Godolphin Cross, Cornwall, United Kingdom, 27.ix.2016 (coll. Clarke) 159d. P. suavis Btl., ♀, Godolphin Cross, Cornwall, United Kingdom, 27.ix.2016 (coll. Clarke) 160a. Parectropis similaria Hufn., ♂, Białowieza, Wojewodztwo Podlaskie, Poland, 31.v.2000 (coll. B. Müller) 160b. P. similaria Hufn., ♂, Behringen, Thüringen, Germany, 8.vi.1969 (coll. B. Müller) 160c. P. similaria Hufn., ♂, Berkenbrück, Brandenburg, Germany, 6.vi.1980 (coll. B. Müller) 160d. P. similaria Hufn., ♀, Maciejów, Poland, 1.vi.1935 (MNHU), 161a. Aethalura punctulata Den. & Schiff., ♂, Güstrow, Mecklenburg, Germany, 10.v.1980 (coll. B. Müller) 161b. A. punctulata Den. & Schiff., ♀, Lanke, Brandenburg, Germany, 16.iv.2008 (coll. B. Müller) 161c. A. punctulata Den. & Schiff., ♂, Ilmenau, Thüringen, Germany, 10.v.1975 (coll. B. Müller) 161d. A. punctulata Den. & Schiff., ♂, Güstrow, Mecklenburg, Germany, 9.v.1980 (coll. B. Müller) 161e. A. punctulata Den. & Schiff., ♀, Güstrow, Mecklenburg, Germany, 7.v.1980 (coll. B. Müller) 162a. Ascotis selenaria Den. & Schiff., ♂, Liljanovo, Pirin Mountains, Bulgaria, 21.v.1982 (coll. B. Müller) 162b. A. selenaria Den. & Schiff., ♂, Liljanovo, Pirin Mountains, Bulgaria, 31.v.1989 (coll. B. Müller) 162c. A. selenaria Den. & Schiff., ♂, Coltesti, Rimetea, Alba, Romania, 29.v.2009 (coll. B. Müller) 162d. A. selenaria Den. & Schiff., ♂, Alt Schadow, Brandenburg, Germany, 22.vi.2008 (coll. B. Müller) 162e. A. selenaria Den. & Schiff., ♂, Ghisoni, Corsica, France, 1.vi.2008 (coll. B. Müller) 162f. A. selenaria Den. & Schiff., ♀, Budaörs, Budapest, Hungary, e. l. 25.vii.1985 (coll. B. Müller)

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PLATE 27 (all specimens natural size) 163a. Cleora cinctaria Den. & Schiff., ♂, Ilmenau, Thüringen, Germany, 11.v.1974 (coll. B. Müller) 163b. C. cinctaria Den. & Schiff., ♂, Martinroda, Thüringen, Germany, 20.v.1973 (coll. B. Müller) 163c. C. cinctaria Den. & Schiff., ♀, Siegelbach, Thüringen, Germany, 18.v.2002 (coll. B. Müller) 163d. C. cinctaria Den. & Schiff., ♀, Budaörs, Budapest, Hungary, 10.iv.1986 (coll. B. Müller) 164a. Ectropis crepuscularia Den. & Schiff., ♂, Elgersburg, Thüringen, Germany, 30.iii.1974 (coll. B. Müller) 164b. E. crepuscularia Den. & Schiff., ♂, Kesztölc, Komárom-Esztergom, Hungary, 16.vii.1985 (coll. B. Müller) 164c. E. crepuscularia Den. & Schiff., ♂, Puig Véntos, Prov. Girona, Spain, 12.vii.1991 (coll. B. Müller) 164d. E. crepuscularia Den. & Schiff., ♀, Gehren, Thüringen, Germany, 8.iv.1973 (coll. B. Müller) 164e. E. crepuscularia Den. & Schiff., ♂, Vivaro, Friuli-Venezia Giulia, Italy, 28.vi.2008 (coll. Skou) 164f. E. crepuscularia Den. & Schiff., ♂, Sibhult, Skåne, Sweden, 9.v.1981 (coll. Skou) 164g. E. crepuscularia Den. & Schiff., ♂, Asserbo, Sjælland, Denmark, 19.vi.1982 (coll. Skou) 164h. E. crepuscularia Den. & Schiff., ♀, Pecs, Pecs County, Hungary, 13.vii.1999 (coll. Skou) 164i. E. crepuscularia Den. & Schiff., ♀, Zühlsdorf, Brandenburg, Germany, e. l. 23.iii.1983 (coll. B. Müller) 164j. E. crepuscularia Den. & Schiff., ♀, Ilfeld, Harz, Sachsen, Germany, 18.iv.2010 (coll. B. Müller) 164k. E. crepuscularia Den. & Schiff., ♀, Snarup Mose, Fyn, Denmark, 27.v.1934 (coll. Skou) 164l. E. crepuscularia Den. & Schiff., ♀, Snarup Mose, Fyn, Denmark, 22.v.1978 (coll. Skou) 165a. Arichanna melanaria Den. & Schiff., ♂, Melchow, Brandenburg, Germany, 10.vii.1982 (coll. B. Müller) 165b. A. melanaria Den. & Schiff., ♂, Melchow, Brandenburg, Germany, 8.vii.1981 (coll. B. Müller) 165c. A. melanaria Den. & Schiff., ♀, Melchow, Brandenburg, Germany, 10.vii.1982 (coll. B. Müller) 165d. A. melanaria Den. & Schiff., ♀, Melchow, Brandenburg, Germany, 8.vii.1981 (coll. B. Müller) 166a. Bupalus piniaria L., ♂, Moscardón, Prov. Teruel, Spain, 17.vi.2007 (coll. B. Müller) 166b. B. piniaria L., ♂, Zühlsdorf, Brandenburg, Germany, e. p. 13.iv.1988 (coll. B. Müller) 166c. B. piniaria L., ♂, Ilmenau, Thüringen, Germany, 14.vi.1976 (coll. B. Müller) 166d. B. piniaria L., ♂, Eptachori, Macedonia, Greece, 13.vii.1998 (coll. Skou) 166e. B. piniaria L., ♂, Villenave d’Ornon, Gironde, France, 20.iv.1957 (MNHU) 166f. B. piniaria L., ♂, Zühlsdorf, Brandenburg, Germany, e. p. 18.v.1983 (coll. B Müller) 166g. B. piniaria L., ♀, Zühlsdorf, Brandenburg, Germany, e. p. 2.iv.1986 (coll. B. Müller) 166h. B. piniaria L., ♀, Skallingen, Vestjylland, Denmark, 27.vi.1977 (coll. Skou) 167a. Megametopon grisolaria Ev., ♂, Aiyrkyzyl Sands, Igniz River basin, Kasakhstan, 9,ix.2013 (coll. Skou) 167b. M. grisolaria Ev., ♂, Saikhin, European part of Kazakhstan, 16.–25.ix.2000 (coll. Skou) 167c. M. grisolaria Ev., ♂, Urda, Ryn-kum steppe, European part of Kazakhstan, 12.–15.ix.2000 (coll. Skou) 167d. M. grisolaria Ev., ♂, Aiyrkyzyl Sands, Igniz River basin, Kasakhstan, 9,ix.2013 (coll. Skou) 167e. M. grisolaria Ev., ♂, Kapustin Yar, Astrakhanskaya obl., Russia, 22.ix.1982 (ZMKU)

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PLATE 28 (all specimens 1.3 x natural size) 168a. Tephronia sepiaria Hufn., ♂, Gumpoldskirchen, Niederösterreich, Austria, viii.1947 (ZSM), barcoded 168b. T. sepiaria Hufn., ♀, Austria, 1887 (ZSM) 168c. T. sepiaria Hufn., ♂, Marciaga, Lago di Garda, Verona, Italy, 27.vii.1990 (coll. B. Müller) 168d. T. sepiaria Hufn., ♂, Naturns, South Tyrol, Italy, 24.vi.1970 (ZSM) 168e. T. sepiaria Hufn., ♂, Puyvert, Vaucluse, France, 8.–10.ix.1988 (ZSM) 168f. T. sepiaria Hufn., ♀, Vendrell, Prov. Tarragona, Spain, 3.vi.1981 (ZSM), barcoded 168g. T. sepiaria Hufn., ♂, Çamlıyayla, Bolkar Dağları, Mersin, Turkey, 1.vii.1996 (coll. B. Müller), barcoded 169a. Tephronia nuragica Fiumi et al., ♂, Aritzo, Sardinia, Italy, 13.viii.1934 (coll. Sommerer) 169b. T. nuragica Fiumi et al., ♂, Camping Lozzi, Corsica, France, 5.–8.viii.2012 (coll. Skou), barcoded 169c. T. nuragica Fiumi et al., ♂, Camping Lozzi, Corsica, France, 5.–8.viii.2012 (coll. Skou), barcoded 170a. Tephronia codetaria sicula Whli., ♀, Madonie, Sicily, Italy (ZFMK), barcoded 170b. T. codetaria sicula Whli., ♀, Madonie, Sicily, Italy 24.viii. [1927] (ZSM), dissected 170c. T. codetaria sicula Whli., ♂, Castel di Ieri, Abruzzo, Italy, 30.vii.2011 (coll. Skou), barcoded 170d. T. codetaria sicula Whli., ♂, Monti Sabini, Lazio, Italy, 11.viii. [1927] (ZSM) 170e. T. codetaria Obth., ♂, Ifrane, Middle Atlas, Morocco, 13.vii.1998 (coll. B. Müller) 171a. Tephronia lhommaria Cleu, ♂, Raposeira, Algarve, Portugal, e. l. 18.v.2006 (ZSM), barcoded 171b. T. lhommaria Cleu, ♀, Albarracin, Prov. Teruel, Spain, 26.vii.1974 (ZSM) 171c. T. lhommaria Cleu, ♀, les Mées, Alpes-de-Haute-Provence, France, 27.vii.1990 (ZSM), barcoded 171d. T. lhommaria Cleu, ♀, St. Colomban, Alpes-Maritimes, France, 27.vii.1973 (ZSM) 171e. T. lhommaria Cleu, ♂, Lagorce, Ardèche, France, e. l. 19.vii.1932 (ZSM) 172a. Tephronia cyrnea Schaw., ♂, Stagno de Platamona, Sardinia, Italy, 23.v.2004 (coll. Skou), barcoded, dissected 172b. T. cyrnea Schaw., ♀, Bonifacio, Corsica, France, 29.v.2014 (ZSM), barcoded, dissected 172c. T. cyrnea Schaw., ♂, Francardo, Corsica, France, 4.viii.1988 (ZSM), dissected 172d. T. cyrnea Schaw., ♀, Su Cologone, Sardinia, Italy, 8.ix.1998 (coll. Pöll), barcoded 172e. T. cyrnea Schaw., ♀, Ciamanacce, Corsica, France, 7.–14.viii.1977 (ZSM), dissected 173a. Tephronia lepraria Rbl., ♂, Oberweiden, Niederösterreich, Austria, 25.vii.1936 (ZSM) 173b. T. lepraria Rbl., ♀, Bugac, Bács-Kiskun, Hungary, 4.viii.1980 (HNHM), barcoded, dissected 173c. T. lepraria Rbl., ♀, Skopje, Macedonia, 14.viii.1973 (coll. Skou), barcoded 173d. T. lepraria Rbl., ♂, Kosmas, Peloponnese, Greece, 4.ix.2004 (coll. Friedrich), 173e. T. lepraria Rbl., ♀, Konya, Turkey, 23.vii.1990 (ZSM), barcoded 174a. Tephronia psyloritaria Reiss., ♂, Omalos, Crete, Greece, 28.vii.–2.viii.2001 (coll. Skou), barcoded 174b. T. psyloritaria Reiss., ♀, Omalos, Crete, Greece, 28.vii.–2.viii.2001 (coll. Skou), barcoded 174c. T. psyloritaria Reiss., ♀, Kares, Crete, Greece, e. l. 28.v.2012 (coll. Friedrich), barcoded, dissected 174d. T. psyloritaria Reiss., ♂, Vafes, Crete, Greece, 1.x.2011 (coll. Friedrich), barcoded 174e. T. psyloritaria Reiss., ♀, Idneon, Crete, Greece, 17.viii.1982 (ZSM), barcoded, dissected 175a. Tephronia theophilaria Hausm., ♂, Bale, Istria, Croatia, late viii.1968 (ZSM) 175b. T. theophilaria Hausm., ♀, Bale, Istria, Croatia, early ix.1969 (ZSM) 175c. T. theophilaria Hausm., ♂, Pizzola, Abruzzo, Italy, 4.viii.1970 (ZSM) 175d. T. theophilaria Hausm., ♂, Pizzola, Abruzzo, Italy, 4.viii.1970 (ZSM) 175e. T. theophilaria Hausm., ♂, Camping Lozzi, Corsica, France, 5.–8.viii.2012 (coll. Skou), barcoded 176a. Tephronia espaniola Schaw., ♂, La Granja, Prov. Segovia, Spain (ZSM), barcoded, dissected 176b. T. espaniola Schaw., ♀, Burgos, Spain,14.viii.1900 (ZSM), dissected 176c. T. espaniola Schaw., ♀, Moscardón, Prov. Teruel, Spain, 28.vii.2014 (coll. Skou), barcoded 176d. T. espaniola Schaw., ♂, Moscardón, Prov. Teruel, Spain, 28.vii.2014 (coll. Skou), barcoded 176e. T. espaniola Schaw., ♀, Estaña, Prov. Huesca, Spain, 25.–26.vii.2014 (coll. Skou), barcoded 177a. Eumannia oppositaria Mann, ♂, Sweti Wratsch, Macedonia, 10.v.1933 (ZSM) 177b. E. oppositaria Mann, ♂, Stavros, Macedonia, early viii.1977 (ZSM) 177c. E. oppositaria Mann, ♀, Rasmun, Ochrid, Macedonia, 1.–31.vii.1936 (ZSM) 177d. E. oppositaria Mann, ♂, Fascomillia, Thesprotia, Epirus, Greece, 30.vi.1998 (ZSM) 177e. E. oppositaria Mann, ♂, Bolkar Dağları, Mercin, Turkey, 5.vii.1996 (coll. B. Müller) 178a. Eumannia oranaria Stgr., ♂, Ifrane, Middle Atlas, Morocco, 17.vii.1998 (coll. B. Müller) 178b. E. oranaria castiliaria Stgr., ♂, Aiguefreda, Prov. Girona, Spain, 17.vii.1991 (coll. B. Müller) 178c. E. oranaria castiliaria Stgr., ♂, St. Tropez, Var, France, 27.vi.1948 (ZSM) 178d. E. oranaria castiliaria Stgr., ♂, Rompon, Ardèche, France, e. l. 23.vi.1897 (ZSM) 178e. E. oranaria castiliaria Stgr., ♂, Mougins, Alpes-Maritimes, France, 8.–30.vi.1963 (ZSM)

622

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PLATE 29 ( most specimens natural size, but species no. 179–181: 1.3 x magnified, 185–187: 1.5 x magnified) 179a. Phyllometra culminaria Ev., ♂, Pilis-hegység, Hungary, 17.v.1956 (ZSM), dissected 179b P. culminaria Ev., ♂, Pilisvörösvá, Hungary, 24.v.1931 (ZSM) 179c. P. culminaria Ev., ♂, Ural, Russia (ZSM) 179d. P. culminaria Ev., ♂, Kapustin Yar, Astrakhanskaya obl., Russia, 12.v.1982 (ZMKU) 179e. P. culminaria Ev., ♂, Germob, Koped-Dagh Mountains, Turkmenistan, 20.iv.1993 (ZSM) 179f. P. culminaria Ev., ♀, Pilisvörösvár, Hungary, 4.v.1934 (ZSM) 180a. Phyllometra gracilaria Bsd., ♂, Albarracin, Prov. Teruel, Spain, 26.–30.v.2003 (ZSM), barcoded 180b. P. gracilaria Bsd., ♂, ♂, Albarracin, Prov. Teruel, Spain, 26.–30.v.2003 (ZSM), barcoded 180c. P. gracilaria Bsd., ♂, Tragacete, Prov. Cuenca, Spain, 15.vi.1982 (ZSM) 180d. P. gracilaria Bsd., ♂, Albarracin, Prov. Teruel, Spain, 25.v.1952 (ZSM) 180e. P. gracilaria Bsd., ♀, Albarracin, Prov. Teruel, Spain, 15.vi.1933 (ZSM) 181a. Ithysia pravata Hbn., ♂, Zolotoje, Crimea, Ukraine, 6.xi.2013 (ZMKU) 181b. I. pravata Hbn., ♂, Zolotoje, Crimea, Ukraine, 6.xi.2013 (ZMKU) 181c. I. pravata Hbn., ♂, Askania Nova nature reserve, Ukraine, 30.x.1984 (ZMKU) 181d. I. pravata Hbn., ♂, Kamenskoe, Crimea, Ukraine, 5.xi.2010 (ZSM), barcoded 181e. I. pravata Hbn., ♂, underside, Zolotoje, Crimea, Ukraine, 6.xi.2013 (ZMKU) 181f. I. pravata Hbn., ♀, Syuyur-Tash, Zolotoje, Crimea, Ukraine,19.xi.2010 (ZMKU) 182a. Leucobrephos middendorfii Mén., ♂, Shuryskary village, Polar Ural, Russia, 7.v.1997 (coll. Nupponen) 182b. L. middendorfii Mén., ♂, Shuryskary village, Polar Ural, Russia, 7.v.1997 (coll. Nupponen) 182c. L. middendorfii Mén., ♂, By river Dzhelinda, Dzhugdzhur mountans, Khabarovskij kraj, Russia, 20.iv.1915

(ZMKU) 182d. L. middendorfii Mén., ♀, By river Dzhelinda, Dzhugdzhur mountans, Khabarovskij kraj, Russia, 30.iv.1915 (ZMKU) 183a. Myinodes constantina Hausm., ♂, El Aiounet, Road to Jerada, East Morocco, 10.iv.1972 (ZSM) 183b. M. constantina Hausm., ♂, Thala, Tunisia, vi.1911 (ZSM) 183c. M. constantina Hausm., ♀, Djebel Penalba, Tunisia, 23.iii.1933 (ZSM) 183d. M. constantina Hausm., ♀, Ifrane, Middle Atlas, Morocco,10.–15.iv.1950 (ZSM) 184a. Kuchleria menadiara Th.-M., ♂, Pozallo, Sicily, Italy, 20.–21.ix.2014 (coll. Skou) 184b. K. menadiara Th.-M., ♂, Pozallo, Sicily, Italy, 20.–21.ix.2014 (coll. Skou) 184c. K. menadiara Th.-M., ♂, Pozallo, Sicily, Italy, 20.–21.ix.2014 (ZSM) 184d. K. menadiara Th.-M., ♂, Pozallo, Sicily, Italy, 20.–21.ix.2014 (ZSM) 185a. Idaea barbuti Taut. & Lev., ♂, Tebosa, Minho, Portugal, 5.viii.2009 (MNHN), holotype, barcoded, dissected 185b. I. barbuti Taut. & Lev., ♀, Tebosa, Minho, Portugal, 5.viii.2009 (MNHN), paratype, barcoded, dissected 186a. Idaea textaria Led., ♂, Marrasch, Taurus Mountains, Turkey, 28.xi.1929 (ZSM) 186b. I. textaria Led., ♀, Marrasch, Taurus Mountains, Turkey, 11.xi.1929 (ZSM) 187a. Idaea lobaria Chrét., ♂, Tozeur, Tunisia, vi.1907 (ZSM) 187b. I. lobaria Chrét., ♂, En Gedi, Neot hakikkar, Israel, 18.iv.1990 (ZSM), barcoded 187c. I. lobaria Chrét., ♂, Tozeur, Tunisia, vi.1907 (ZSM) 187d. I. lobaria Chrét., ♀, Biskra, Algeria, vi.1907 (ZSM) 187e. I. lobaria Chrét., ♂, Tozeur, Tunisia, vi.1907 (ZSM) 188a. Idaea gelbrechti Hausm., ♂, Ifrane, Middle Atlas, Morocco, e. o. 13.vii.1998 (ZSM), holotype, barcoded 188b. I. gelbrechti Hausm., ♂, Ifrane, Middle Atlas, Morocco, 23.vii.–1.viii.1988 (ZSM), paratype 188c. I. gelbrechti Hausm., ♀, Puerto de la Ragua, Sierra Nevada, Prov. Almeria, Spain, e. o. 24.vii.2009 (coll. Skou) 188d. I. gelbrechti Hausm., ♀, Puerto de la Ragua, Sierra Nevada, Prov. Almeria, Spain, e. o. 24.vii.2009 (coll. Skou) 189a. Oar pratana F., ♂, Tunis, Tunisia, 12.v.1911 (ZSM) 189b. O. pratana F., ♂, [North Africa], 21.x.1911 (ZSM), 190a. Scopula drenowskii Stern., ♂, Göle, Erzurum, Turkey, 11.viii.1976 (ZSM), dissected 190b. S. drenowskii Stern., ♂, Kızılcahamam, Turkey, 6.vi.1995 (ZSM), barcoded 190c. S. drenowskii Stern., ♂, Gürün, Turkey, 20.vi.1995 (ZSM), barcoded 190d. S. drenowskii Stern., ♂, Tashdere, Sivas, Turkey, 24.vi.2005 (coll. Skou), barcoded, dissected 190e. S. drenowskii Stern., ♂, Tashdere, Sivas, Turkey, 24.vi.2005 (coll. Skou), barcoded, dissected 191a. Scopula donovani Dist., ♂, Taroudannt, Morocco, iv.1928 (ZSM) 191b. S. donovani Dist., ♂, Aït Melloul, Morocco, 13.iv.1969 (ZSM) 191c. S. donovani Dist., ♂, Ouesso, Peoples Republic of Congo, 22.vi.1984 (ZSM) 191d. S. donovani Dist., ♀, Aït Melloul, Morocco, 13.iv.1969 (ZSM) 191e. S. donovani Dist., ♀, Lamto, Ivory Coast, 24.v.1983 (ZSM)

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625

PLATE 30 (most specimens 1.5 x magnified, but species 192, 193, 201 and 202 natural size) 192a. Nothocasis rosariae Scalercio et al., ♂, Timpone Magara, Saracena, Calabria, Italy ix.–x.1915

(­Crea-Sam) 192b. N. rosariae Scalercio et al., ♀, Timpone Magara, Saracena, Calabria, Italy ix.–x.1915 (Crea-Sam) 192c. N. rosariae Scalercio et al., ♂, Timpone Magara, Saracena, Calabria, Italy ix.–x.1915 (Crea-Sam) 192d. N. rosariae Scalercio et al., ♂, Timpone Magara, Saracena, Calabria, Italy ix.–x.1915 (Crea-Sam) 192e. N. rosariae Scalercio et al., ♀, Vall. Lupara, Alessandria, Calabria, Italy, 12.xi.2017 (coll. Skou) 193a. Perizoma juracolaria Whli., ♂, Le Lautaret, Hautes Alpes, France, 30.vi.1952 (ZSM) 193b. P. juracolaria Whli., ♂, Jägerkamp, Bayern, Germany, mid vii.1975 (ZSM) 193c. P. juracolaria Whli., ♂, Le Lautaret, Hautes Alpes, France, 30.vi.1952 (ZSM) 193d. P. juracolaria Whli., ♂, Le Lautaret, Hautes Alpes, France, 30.vi.1952 (ZSM) 194. Perizoma barrassoi Zahm et al., ♂, Valle Cannella, Maiella, Abruzzo, Italy, 23.vii.1988 (ZSM), holotype, barcoded, dissected 195a. Eupithecia carpophilata Püng., ♂, Ricorda Island, Primorye, Russia, 15.–31.viii.2012 (ZSM) 195b. E. carpophilata Püng., ♂, Nakhodka, Lozovoy range, Primorye, Russia, 23.vii.1999 (ZISP) 195c. E. carpophilata Püng., ♂, Inilchek, Kaindy mountain range, Kirgizia, vii.1993 (ZSM) 195d. E. carpophilata Püng., ♂, Inilchek, Kaindy mountain range, Kirgizia, vii.1993 (ZSM) 195e. E. carpophilata Püng., ♂, Inilchek, Kaindy mountain range, Kirgizia, vii.1993 (ZSM), barcoded 196a. Eupithecia dubiosa Dietze, ♂, N. Ammud, Israel, 18.iii.1991 (ZSM) 196b. E. dubiosa Dietze, ♂, Eilon, Israel, 25.iii.2000 (ZSM) 196c. E. dubiosa Dietze, ♂, Nahal Kziv, Israel, 24.iii.1987 (ZSM), dissected 196d. E. dubiosa Dietze, ♂, Tarbaja beach, Beirut, Lebanon, 3.–16.iv.1968 (ZSM) 196e. E. dubiosa Dietze, ♂, Paphos, Cyprus, 13.–19.iii.1994 (ZSM), dissected 197a. Eupithecia furcata Stgr., ♂, Kirikkale, Ankara, Turkey, 29.v.1964 (ZSM) 197b. E. furcata Stgr., ♂, Kirikkale, Ankara, Turkey, 29.v.1964 (ZSM) 197c. E. furcata Stgr., ♂, Kirikkale, Ankara, Turkey, 29.v.1964 (ZSM) 197d. E. furcata Stgr., ♂, Beysehir gölü, Konya, Turkey, 4.vi.1964 (ZSM) 197e. E. furcata Stgr., ♂, Beysehir gölü, Konya, Turkey, 4.vi.1964 (ZSM) 198a. Eupithecia gypsophilata Skou et al., ♂, Bujaraloz, Prov. Huesca, Spain, 8.viii.1962 (ZSM), paratype, dissected 198b. E. gypsophilata Skou et al., ♀, Bujaraloz, Prov. Huesca, Spain, 31.vii.1962 (ZSM), paratype 198c. E. gypsophilata Skou et al., ♂, Peñalba, Prov. Huesca, Spain, 28.vii.1991 (coll. Skou), holotype 198d. E. gypsophilata Skou et al., ♂, Peñalba, Prov. Huesca, Spain, 28.vii.1991 (coll. Skou), paratype 198e. E. gypsophilata Skou et al., ♀, Peñalba, Prov. Huesca, Spain, 28.vii.1991 (coll. Skou), paratype 199a. Eupithecia pindosata Weidlich, ♀, Krania, Thessalia, Greece 2.v.2002 (coll. Weidlich), paratype 199b. E. pindosata Weidlich, ♀, Krania, Thessalia, Greece 5.v.2003 (coll. Weidlich), paratype 199c. E. pindosata Weidlich, ♂, Krania, Thessalia, Greece, 19.iv.2018 (coll. Skou) 199d. E. pindosata Weidlich, ♂, Krania, Thessalia, Greece, 19.iv.2018 (coll. Skou) 199e. E. pindosata Weidlich, ♀, Krania, Thessalia, Greece, 19.iv.2018 (coll. Skou) 200a. Eupithecia goossensiata Mab., ♂, Steiermark, Austria (ZSM) 200b. E. goossensiata Mab., ♂, Wilhelmshausen, Hessen, Germany, e. o. 2.vii.1950 (ZSM) 200c. E. goossensiata Mab., ♂, Vennermoor, Niedersachsen, Germany, e. p. 5.viii.1985 (ZSM) 200d. E. goossensiata Mab., ♀, Poland, vii.1901 (ZSM) 200e. E. goossensiata Mab., ♂, [Niedersachsen, Germany] (ZSM) 201a. Ourapteryx niveiscythes Zol., ♂, Gunib, Dagestan, Russia, 18.–20.vii.2002 (coll. Skou) 201b. O. niveiscythes Zol., ♂, Gunib, Dagestan, Russia, 18.–20.vii.2002 (coll. Skou) 201c. O. niveiscythes Zol., ♂, Kislovodsk, Northern Caucasus, Russia, 18.vii.1905 (ZMKU) 202a. Dyscia govii Fiumi & Flam., ♂, Francardo, Corsica, France, 26.v.2012 (coll. Govi), paratype 202b. D. govii Fiumi & Flam., ♂, Francardo, Corsica, France, 26.v.2012 (coll. Govi), paratype 202c. D. govii Fiumi & Flam., ♂, Francardo, Corsica, France, 17.vi.2012 (coll. Govi), paratype

626

627

Genitalia plates

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 1. Odontognophos dumetata (Treitschke). a. genitalia, b. aedeagus, c. vesica, d. valva apex. 2. Odontognophos margaritata (Zerny). a. genitalia, b. aedeagus, c. vesica, d. valva apex. 3. Odontognophos zacharia (Costa). a. genitalia, b. aedeagus, c. vesica, d. valva apex.

630

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 4. Dicrognophos sartata (Treitschke). a. genitalia, b. aedeagus, c. vesica, d. cornutus. 5. Dicrognophos siciliana Skou & Sihvonen. a. genitalia, b. aedeagus, c. vesica, d. cornutus. 6. Ortaliella perspersata (Treitschke). a. genitalia, b. aedeagus (lateral view), c. aedeagus (ventral view). d. vesica, e. uncus.

631

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 7. Apochima flabellaria (Hübner). a. genitalia, b. aedeagus, c. vesica. 8. Chondrosoma fiducaria Anker. a. genitalia, b. aedeagus, c. vesica. 9. Gnophos furvata (Denis & Schiffermüller). a. genitalia, b. aedeagus. 10. Gnophos obfuscata (Denis & Schiffermüller). a. genitalia, b. aedeagus.

632

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 11. Charissa obscurata (Denis & Schiffermüller). a. genitalia, b. aedeagus. 12. Charissa crenulata (Staudinger). a. genitalia, b. aedeagus. 13. Charissa italohelveticus (Rezbanyai-Reser). a. genitalia, b. aedeagus.

633

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 14. Charissa pullata (Denis & Schiffermüller). a. genitalia, b. aedeagus. 15. Charissa difficilis (Alphéraky). a. genitalia, b. aedeagus. 16. Charissa turfosaria (Wehrli). a. genitalia, b. aedeagus.

634

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 17. Charissa mutilata (Staudinger). a. genitalia, b. aedeagus, c. valva. 18. Charissa peloponnesiaria Erlacher & Erlacher. a. genitalia, b. aedeagus, c. valva. 19. Charissa pentheri (Rebel). a. genitalia, b. aedeagus, c. valva.

635

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 20. Charissa assoi (Redondo & Gastón). a. genitalia, b. aedeagus. 21. Charissa corsica (Oberthür). a. genitalia, b. aedeagus. 22. Charissa staudingeri (Wnukowsky). a. genitalia, b. aedeagus.

636

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 23. Charissa subtaurica (Wehrli). a. genitalia, b. aedeagus. 24. Charissa variegata (Duponchel). a. genitalia, b. aedeagus. 25. Charissa mucidaria (Hübner). a. genitalia, b. aedeagus.

637

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 26. Charissa ambiguata (Duponchel). a. genitalia, b. aedeagus. 27. Charissa predotae (Schawerda). a. genitalia, b. aedeagus. 28. Charissa onustaria (Herrich-Schäffer). a. genitalia, b. aedeagus.

638

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 29. Charissa remmi (Viidalepp). a. genitalia, b. aedeagus. 30. Charissa zeitunaria (Staudinger). a. genitalia, b. aedeagus. 31. Charissa glaucinaria (Hübner). a. genitalia, b. aedeagus.

639

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 32. Charissa pfeifferi (Wehrli). a. genitalia, b. aedeagus. 33. Charissa supinaria (Mann). a. genitalia, b. aedeagus. 34. Charissa graecaria (Staudinger). a. genitalia, b. aedeagus.

640

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 35. Psodos alpinata (Scopoli). a. genitalia, b. aedeagus, c. juxta. 36. Psodos canaliculata (Hochenwarth). a. genitalia, b. aedeagus, c. juxta. 37. Psodos werneri Schawerda. a. genitalia, b. aedeagus, c. juxta.

641

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 38. Psodos coracina (Esper). a. genitalia, b. aedeagus, c. juxta. 39. Psodos bentelii Rätzer. a. genitalia, b. aedeagus, c. juxta. 40. Psodos noricana Wagner. a. genitalia, b. aedeagus, c. juxta.

642

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 41. Psodos belzebuth Praviel. a. genitalia, b. aedeagus, c. juxta. 42. Psodos pyrenaica Schawerda. a. genitalia, b. aedeagus, c. juxta. 43. Psodos alpmaritima Wehrli. a. genitalia, b. aedeagus, c. juxta. 44. Psodos alticolaria Mann. a. genitalia, b. aedeagus, c. juxta.

643

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 45. Psodos wehrlii Vorbrodt. a. genitalia, b. aedeagus, c. juxta. 46. Psodos baldensis Wolfsberger. a. genitalia, b. aedeagus, c. juxta. 47. Psodos spitzi Rebel. a. genitalia, b. aedeagus, c. juxta. 48. Psodos quadrifaria (Sulzer). a. genitalia, b. aedeagus, c. juxta.

644

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 49. Sciadia tenebraria (Esper). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 50. Sciadia septaria (Guenée). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 51. Sciadia innuptaria (Herrich-Schäffer). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 52. Sciadia slovenica Leraut. a. genitalia, b. aedeagus, c. vesica, d. juxta arms.

645

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 53. Sciadia dolomitica Huemer & Hausmann. a. genitalia, b. aedeagus, c. vesica (position of large cornutus distorted under cover glass), d. juxta arms. 54. Sciadia zelleraria (Freyer). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 55. Sciadia unicoloraria (Staudinger). a. genitalia, b. aedeagus, c. vesica, d. juxta arms.

646

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 56. Elophos caelibaria (Heydenreich). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 57. Elophos zirbitzensis (Pieszcek). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 58. Elophos operaria (Hübner). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 59. Elophos andereggaria (Harpe). a. genitalia, b. aedeagus, c. vesica, d. juxta arms.

647

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 60. Yezognophos dilucidaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 61. Yezognophos sproengertsi (Püngeler). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 62. Yezognophos vittaria (Thunberg). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm (margin highlighted partially).

648

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1). 63–1. Yezognophos serotinaria serotinaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 63–2. Yezognophos serotinaria dognini (Thierry-Mieg). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm. 63–3. Yezognophos serotinaria serotinoides (Wehrli). a. genitalia, b. aedeagus, c. vesica, d. apex of juxta arm.

649

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 64. Crocota tinctaria (Hübner). a. genitalia, b. aedeagus, c. apex of aedeagus. 65. Crocota peletieraria (Duponchel). a. genitalia, b. aedeagus, c. apex of aedeagus. 66. Crocota pseudotinctaria Leraut. a. genitalia, b. aedeagus, c. vesica, d. apex of aedeagus. 67. Crocota niveata (Scopoli). a. genitalia, b. aedeagus, c. vesica, d. apex of aedeagus (ventral view).

650

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 68. Synopsia sociaria (Hübner). a. genitalia, b. aedeagus, c. vesica, d. valva. 69. Phthonandria strictaria (Lederer). a. genitalia, b. aedeagus, c. vesica, d. uncus (dorsal view). 70. Menophra abruptaria (Thunberg). a. genitalia, b. aedeagus, c. apex of aedeagus, d. valva.

651

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 71. Menophra japygiaria (O. Costa). a. genitalia, b. aedeagus, c. cornuti, d. valva. 72. Menophra berenicidaria (Turati). a. genitalia, b. aedeagus, c. aedeagus, d. valva. 73. Menophra harterti (Rothschild). a. genitalia, b. aedeagus, c. cornuti, d. valva.

652

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 74. Menophra thuriferaria (Zerny). a. genitalia, b. aedeagus, c. cornuti, d. valva. 75. Menophra nycthemeraria (Geyer). a. genitalia, b. aedeagus, c. cornuti. 76. Menophra annegreteae Skou. a. genitalia, b. aedeagus, c. apex of aedeagus. d. valva.

653

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 77. Cleorodes lichenaria (Hufnagel). a. genitalia, b. aedeagus, c. cornutus, d. uncus. 78. Zernyia granataria (Staudinger). a. genitalia, b. aedeagus, c. cornutus, d. uncus. 79. Megalycinia serraria (A. Costa). a. genitalia, b. aedeagus, c. vesica, d. rounded lobe (apparently originating from the tegumen, see arrow in figure a).

654

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 80. Dasypteroma thaumasia Staudinger. a. genitalia, b. aedeagus with juxta, c. aedeagus, d. ­vesica, e. gnathos. 81. Microbiston lanaria (Eversmann). a. genitalia, b. aedeagus, c. apex of aedeagus, d. apical half of valva. 82. Apocheima hispidaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. apex of aedeagus, d. apical half of valva.

655

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 83. Phigalia pilosaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. central part of valva. 84. Hypomecis roboraria (Denis & Schiffermüller. a. genitalia, b. aedeagus, c. vesica, d. central part of valva. 85. Hypomecis punctinalis (Scopoli). a. genitalia, b. aedeagus, c. central part of valva.

656

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 86. Ematurga atomaria (Linnaeus). a. genitalia, b. aedeagus, c. vesica, d. central part of valva. 87. Lycia hirtaria (Clerck). a. genitalia, b. aedeagus, c. vesica, d. cornuti. 88. Lycia necessaria (Zeller). a. genitalia, b. aedeagus.

657

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 89. Lycia lapponaria (Boisduval). a. genitalia, b. aedeagus, c. apex of aedeagus. 90. Lycia isabellae (Harrison). a. genitalia, b. aedeagus, c. apex of aedeagus. 91. Lycia pomonaria (Hübner). a. genitalia, b. aedeagus, c. vesica, d. cornuti.

658

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 92. Lycia zonaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. apex of aedeagus. 93. Lycia alpina (Sulzer). a. genitalia, b. aedeagus, c. apex of aedeagus. 94. Lycia graecarius (Staudinger). a. genitalia, b. aedeagus. 95. Lycia florentina (Stefanelli). a. genitalia, b. aedeagus.

659

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 96. Biston strataria (Hufnagel). a. genitalia, b. aedeagus, c. vesica, d. uncus, gnathos and apex of juxta. 97. Biston rosenbaueri Müller. a. genitalia, b. aedeagus, c. uncus, gnathos and apex of juxta. 98. Biston achyra Wehrli. a. genitalia, b. aedeagus, c. uncus, gnathos and apex of juxta. 99. Biston betularia (Linnaeus). a. genitalia, b. aedeagus, c. vesica, d. uncus and gnathos.

660

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 100. Erannis defoliaria (Clerck). a. genitalia, b. aedeagus, c. vesica. 101. Erannis ankeraria (Staudinger). a. genitalia, b. aedeagus. 102. Erannis declinans (Staudinger). a. genitalia, b. aedeagus.

661

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 103. Kresnaia beschkovi Ganev. a. genitalia, b. aedeagus, c. apex of aedeagus. 104. Agriopis leucophaearia (Denis & Schiffermüller). a. genitalia, b. aedeagus. 105. Agriopis bajaria (Denis & Schiffermüller). a. genitalia, b. aedeagus.

662

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 106. Agriopis aurantiaria (Hübner). a. genitalia, b. aedeagus, c. vesica. 107. Agriopis budashkini Kostjuk. a. genitalia, b. aedeagus. 108. Agriopis marginaria (Fabricius). a. genitalia, b. aedeagus.

663

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 109. Calamodes occitanaria (Duponchel). a. genitalia, b. aedeagus. 110. Calamodes melanaria Oberthür. a. genitalia, b. aedeagus. 111. Paradarisa consonaria (Hübner). a. genitalia, b. aedeagus.

664

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 112. Larerannis orthogrammaria (Wehrli). a. genitalia, b. aedeagus, c. apical half of valva. 113. Chemerina caliginearia (Rambur). a. genitalia, b. aedeagus. 114. Fagivorina arenaria (Hufnagel). a. genitalia, b. aedeagus, c. apical half of valva.

665

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 115. Spartopteryx kindermannaria Staudinger. a. genitalia, b. aedeagus. 116. Athroolopha pennigeraria (Hübner). a. genitalia, b. aedeagus. 117. Athroolopha chrysitaria (Hübner). a. genitalia, b. aedeagus.

666

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 118. Eurranthis plummistaria (Villers). a. genitalia, b. aedeagus, c. apical half of valva. 119. Nychiodes obscuraria (Villers). a. genitalia, b. aedeagus with everted vesica. 120. Nychiodes ragusaria Millière. a. genitalia, b. aedeagus with everted vesica.

667

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 121. Nychiodes andalusiaria Millière. a. genitalia, b. aedeagus with everted vesica. 122. Nychiodes notarioi Expósito. a. genitalia, b. aedeagus with everted vesica. 123. Nychiodes hispanica Wehrli. a. genitalia, b. aedeagus with everted vesica, c-h. variation of valval ornamentation.

668

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 124. Nychiodes waltheri Wagner. a. genitalia, b. aedeagus. 125. Nychiodes amygdalaria (Herrich-Schäffer). a. genitalia, b. aedeagus. 126. Nychiodes dalmatina Wagner. 1a. genitalia, 1b. aedeagus with everted vesica, 2. valva.

669

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 127. Phaselia serrularia (Eversmann). a. genitalia, b. aedeagus. 128. Phaselia algiricaria aragona Wehrli. a. genitalia, b. aedeagus in ventral view, c. aedeagus in lateral view, d. uncus and gnathos.

670

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 129. Ekboarmia atlanticaria (Staudinger). a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 130. Ekboarmia sagnesi Dufay. a. genitalia, b. aedeagus, c. vesica, d. juxta arms. 131. Ekboarmia miniaria Skou, Stüning & Sihvonen. a. genitalia, b. aedeagus, c. vesica, d. ­juxta arms.

671

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 132. Ecleora solieraria (Rambur). a. genitalia, b. aedeagus, c. apical half of valva. 133. Afriberina tenietaria (Staudinger). a. genitalia, b. aedeagus, c. vesica, d. apex of valva. 134. Afriberina salemae Skou & Sihvonen. a. genitalia, b. aedeagus, c. vesica, d. apex of valva.

672

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 135. Sardocyrnia bastelicaria (Bellier). a. genitalia, b. aedeagus, c. apical half of valva. 136. Sardocyrnia fortunaria (Vázquez). a. genitalia, b. aedeagus, c. apical half of valva.

673

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 137. Selidosema brunnearia (Villers). a. genitalia, b. aedeagus, c. apical half of valva. 138. Selidoseman plumaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. apical half of valva. 139. Selidosema pyrenaearia (Boisduval). a. genitalia, b. aedeagus, c. apical half of valva.

674

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 140. Selidosema taeniolaria (Hübner). a. genitalia, b. aedeagus, c. apex of aedeagus. 141. Selidosema ambustaria (Geyer). a. genitalia, b. aedeagus, c. apex of aedeagus. 142. Selidosema erebaria postdentaria (Rothschild). a. genitalia, b. aedeagus, c. apical half of valva.

675

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 143. Peribatodes rhomboidaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view), e. setal comb (male abdominal sternite 3). 144. Peribatodes perversaria (Boisduval). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view). 145. Peribatodes omalensis Rajaei, Skou & Hausmann. a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view).

676

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 146. Peribatodes umbraria (Hübner). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view), d. setal comb (male abdominal sternite 3). 147. Peribatodes ebusaria Vallhonrat. a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view). 148. Peribatodes secundaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. ­uncus (lateral view).

677

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 149. Peribatodes correptaria (Zeller). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view). 150. Peribatodes aragonis (Wehrli). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view). 151. Peribatodes ilicaria (Geyer). a. genitalia, b. aedeagus, c. vesica, d. uncus (lateral view), e. setal comb (male abdominal sternite 3).

678

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 152. Alcis repandata (Linnaeus). a. genitalia, b. aedeagus. 153. Alcis deversata (Staudinger). a. genitalia, b. aedeagus. 154. Alcis jubata (Thunberg). a. genitalia, b. aedeagus.

679

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 155. Deileptenia ribeata (Clerck). a genitalia, b. aedeagus, c. central half of valva. 156. Paraboarmia viertlii (Bohatsch). a. genitalia, b. aedeagus, c. central part of valva. 157. Adactylotis gesticularia (Hübner). a. genitalia, b. aedeagus. 158. Adactylotis contaminaria (Hübner). a. genitalia, b. aedeagus.

680

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 159. Pseudocoremia suavis Butler. a. genitalia, b. aedeagus, c. vesica, d. cornuti at base of vesica, d. apex of juxta arm. 160. Parectropis similaria (Hufnagel). a. genitalia, b. aedeagus, c. central part of valva. 161. Aethalura punctulata (Denis & Schiffermüller). a. genitalia, b. aedeagus.

681

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 162. Ascotis selenaria (Denis & Schiffermüller). a. genitalia, b. aedeagus. 163. Cleora cinctaria (Denis & Schiffermüller). a. genitalia, b. aedeagus. 164. Ectropis crepuscularia (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. everted vesica.

682

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 165. Arichanna melanaria (Denis & Schiffermüller). a. genitalia, b. aedeagus, c. vesica, d. cornuti. 166. Bupalus piniaria (Linnaeus). a. genitalia, b. aedeagus. 167. Megametopon grisolaria (Eversmann). a. genitalia, b. aedeagus.

683

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 168. Tephronia sepiaria (Hufnagel). a. genitalia, b. aedeagus. 169. Tephronia nuragica Fiumi et al. a. genitalia, b. aedeagus. 170. Tephronia codetaria (Oberthür) subsp. sicula (Wehrli). a. genitalia, b. aedeagus. 171. Tephronia lhommaria (Cleu) subsp. melaleucaria (Schwingenschuss). a. genitalia, b. aede­ agus.

684

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 172-1. Tephronia cyrnea (Schawerda). a. genitalia, b. aedeagus (Corsica). 172-2. Tephronia cyrnea (Schawerda). a. genitalia, b. aedeagus (Corsica, variant). 172-3. Tephronia cyrnea (Schawerda). a. genitalia, b. aedeagus (Sardinia). 173. Tephronia lepraria Rebel. a. genitalia, b. aedeagus.

685

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 174. Tephronia psyloritaria (Reisser). a. genitalia, b. aedeagus. 175. Tephronia theophilaria Hausmann. a. genitalia, b. aedeagus. 176. Tephronia espaniola (Schawerda). a. genitalia, b. aedeagus.

686

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 177. Eumannia oppositaria (Mann). a. genitalia, b. aedeagus. 178-1. Eumannia oranaria (Staudinger). a. genitalia, b. aedeagus. 178-2. Eumannia oranaria subsp. castiliaria (Staudinger). a. genitalia, b. aedeagus. 179. Phyllometra culminaria (Eversmann). a. genitalia, b. aedeagus, c. valva. 180. Phyllometra gracilaria Boisduval. a. genitalia, b. aedeagus, c. valva.

687

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 181. Ithysia pravata (Hübner). a. genitalia, b. aedeagus, c. aedeagus (detail), d. juxta processes. 182. Leucobrephos middendorfii (Ménétriés). a. genitalia, b. aedeagus, c. vesica. 183. Myinodes constantina (Hausmann). a. genitalia, b. aedeagus, c. tip of valva.

688

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 184. Kuchleria menadiara (Thierry-Mieg). a. genitalia, b. aedeagus, c. apex of aedeagus 185. Idaea barbuti Tautel & Lévêque. a. genitalia, b. aedeagus, c. apex of aedeagus with ­cornuti (photo: A. Lévêque). 186. Idaea textaria (Lederer). See male genitalia figure in Vol. 2 fig. 49d. 187. Idaea lobaria (Chrétien). a. genitalia, b. aedeagus. 188. Idaea gelbrechti Hausmann. a. genitalia, b. aedeagus.

689

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 189. Oar pratana (Fabricius). See male genitalia figures in Vol. 2 figs 115b, 115c. 190. Scopula drenowskii Sterneck. a. genitalia, b. aedeagus, c. sternum A8. 191. Scopula donovani (Distant). a. genitalia, b. aedeagus, c. sternum A8. 192. Nothocasis rosariae Scalercio et al. a. genitalia, b. aedeagus, c. detail of valva (tip of sacculus magnified) (photo: S. Scalercio, reproduced with permission from copyright holder).

690

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 193. Perizoma juracolaria (Wehrli). a. genitalia, b. aedeagus. 194. Perizoma barrassoi Zahm et al. a. genitalia, b. aedeagus, c. detail of vesica (magnified). 195. Eupithecia carpophilata Staudinger. a. genitalia, b. aedeagus, c. sternum A7 + A8. 196. Eupithecia dubiosa Dietze. a. genitalia, b. aedeagus, c. sternum A8.

691

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 197. Eupithecia furcata Staudinger. a. genitalia, b. aedeagus, c. sternum A8, left anterior lobe distorted. 198. Eupithecia gypsophilata Skou et al.. a. genitalia, b. aedeagus, c. sternum A8. 199. Eupithecia pindosata Weidlich. a. genitalia, b. aedeagus, c. detail of vesica, d. sternum A8.

692

MALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 1) 200. Eupithecia goossensiata Staudinger. a. genitalia, b. aedeagus, c. sternum A8. 201. Ourapteryx niveiscythes Zolotuhin. a. genitalia, b. aedeagus (vesica everted). (photo: V. Zolotuhin, reproduced with permission from copyright holder). 202. Dyscia govii Fiumi & Flamigni. a. genitalia, b. aedeagus. (photo: G. Fiumi, reproduced with permission from copyright holder).

693

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2. ­Circle, if shown, indicates the point of origin of the ductus seminalis.) 1. Odontognophos dumetata (Treitschke). a. genitalia, b. signum. 2. Odontognophos margaritata (Zerny). a. genitalia, b. signum. 3. Odontognophos zacharia (Costa). a. genitalia, b. signum.

694

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 4. Dicrognophos sartata (Treitschke). a. genitalia, b. ostium bursae and adjacent structures. 5. Dicrognophos siciliana Skou & Sihvonen. a. genitalia, b. ostium bursae and adjacent structures. 6. Ortaliella perspersata (Treitschke).

695

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 7. Apochima flabellaria (Hübner). 8. Chondrosoma fiducaria Anker. 9. Gnophos furvata (Denis & Schiffermüller). a. genitalia, b. signum. 10. Gnophos obfuscata (Denis & Schiffermüller).

696

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 11. Charissa obscurata (Denis & Schiffermüller). 12. Charissa crenulata (Staudinger). 13. Charissa italohelveticus (Rezbanyai-Reser).

697

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 14. Charissa pullata (Denis & Schiffermüller). 15. Charissa difficilis (Alphéraky). 16. Charissa turfosaria (Wehrli).

698

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 17. Charissa mutilata (Staudinger). 18. Charissa peloponnesiaria Erlacher & Erlacher. 19. Charissa pentheri (Rebel).

699

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 20. Charissa assoi (Redondo & Gastón). 21. Charissa corsica (Oberthür). 22. Charissa staudingeri (Wnukowsky). a. genitalia, b. genitalia (variant).

700

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 23. Charissa subtaurica (Wehrli). 24. Charissa variegata (Duponchel). 25. Charissa mucidaria (Hübner).

701

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 26. Charissa ambiguata (Duponchel). a. genitalia, b. genitalia (variant). 27. Charissa predotae (Schawerda). 28. Charissa onustaria (Herrich-Schäffer).

702

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 29. Charissa remmi (Viidalepp). 30. Charissa zeitunaria (Staudinger). 31. Charissa glaucinaria (Hübner).

703

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 32. Charissa pfeifferi (Wehrli). 33. Charissa supinaria (Mann). 34. Charissa graecaria (Staudinger).

704

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 35. Psodos alpinata (Scopoli). 36. Psodos canaliculata (Hochenwarth). 37. Psodos werneri Schawerda. Female is unknown to science. 38. Psodos coracina (Esper).

705

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 39. Psodos bentelii Rätzer. 40. Psodos noricana Wagner. 41. Psodos belzebuth Praviel. 42. Psodos pyrenaica Schawerda.

706

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 43. Psodos alpmaritima Wehrli. 44. Psodos alticolaria Mann. 45. Psodos wehrlii Vorbrodt.

707

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 46. Psodos baldensis Wolfsberger. 47. Psodos spitzi Rebel. 48. Psodos quadrifaria (Sulzer).

708

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 49. Sciadia tenebraria (Esper). a. genitalia, b. signum. 50. Sciadia septaria (Guenée). a. genitalia, b. ostium bursae and adjacent structures. 51. Sciadia innuptaria (Herrich-Schäffer). a. genitalia, b. signum.

709

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 52. Sciadia slovenica Leraut. a. genitalia, b. signum. 53. Sciadia dolomitica Huemer & Hausmann. a. genitalia, b. signum. 54. Sciadia zelleraria (Freyer). a. genitalia, b. signum.

710

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 55. Sciadia unicoloraria (Staudinger). a. genitalia, b. ostium bursae and adjacent structures. 56. Elophos caelibaria (Heydenreich). a. genitalia, b. signum. 57. Elophos zirbitzensis (Pieszcek). a. genitalia, b. ostium bursae and adjacent structures.

711

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 58. Elophos operaria (Hübner). a. genitalia, b. signum, c. genitalia (variant), d. signum (variant). 59. Elophos andereggaria (Harpe). a. genitalia, b. signum.

712

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 60. Yezognophos dilucidaria (Denis & Schiffermüller). a. genitalia, b. signum. 61. Yezognophos sproengertsi (Püngeler). a. genitalia, b. ostium bursae and adjacent structures, c. genitalia (variant), d. signum. 62. Yezognophos vittaria (Thunberg). a. genitalia, b. ostium bursae and adjacent structures.

713

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 63–1. Yezognophos serotinaria serotinaria (Denis & Schiffermüller). a. genitalia, b. signum. 63–2. Yezognophos serotinaria dognini (Thierry-Mieg). a. genitalia, b. signum. 63–3. Yezognophos serotinaria serotinoides (Wehrli). a. genitalia, b. signum.

714

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 64. Crocota tinctaria (Hübner). 65. Crocota peletieraria (Duponchel). 66. Crocota pseudotinctaria Leraut. 67. Crocota niveata (Scopoli).

715

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 68. Synopsia sociaria (Hübner). 69. Phthonandria strictaria (Lederer). a. genitalia, b. signum. 70. Menophra abruptaria (Thunberg).

716

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 71. Menophra japygiaria (O. Costa). 72. Menophra berenicidaria (Turati). 73. Menophra harterti (Rothschild).

717

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 74. Menophra thuriferaria (Zerny). 75. Menophra nycthemeraria (Geyer). 76. Menophra annegreteae Skou.

718

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 77. Cleorodes lichenaria (Hufnagel). 78. Zernyia granataria (Staudinger). 79. Megalycinia serraria (A. Costa). a. genitalia, b. ostium bursae and adjacent structures, c. signum with solid centre, d. signum with irregular centre (and detached cornutus).

719

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 80. Dasypteroma thaumasia Staudinger. 81. Microbiston lanaria (Eversmann). 82. Apocheima hispidaria (Denis & Schiffermüller).

720

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 83. Phigalia pilosaria (Denis & Schiffermüller). 84. Hypomecis roboraria (Denis & Schiffermüller. a. genitalia, b. signum. 85. Hypomecis punctinalis (Scopoli). a. genitalia, b. signum. 86. Ematurga atomaria (Linnaeus).

721

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 87. Lycia hirtaria (Clerck). a. genitalia, b. signum. 88. Lycia necessaria (Zeller). a. genitalia, b. signum. 89. Lycia lapponaria (Boisduval). a. genitalia, b. signum.

722

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 90. Lycia isabellae (Harrison). a. genitalia, b. signum. 91. Lycia pomonaria (Hübner). a. genitalia, b. signum. 92. Lycia zonaria (Denis & Schiffermüller).

723

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 93. Lycia alpina (Sulzer). 94. Lycia graecarius (Staudinger). 95. Lycia florentina (Stefanelli).

724

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 96. Biston strataria (Hufnagel). 97. Biston rosenbaueri Müller. Female is unknown to science. 98. Biston achyra Wehrli. Female is unknown to science. 99. Biston betularia (Linnaeus).

725

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 100. Erannis defoliaria (Clerck). 101. Erannis ankeraria (Staudinger). 102. Erannis declinans (Staudinger).

726

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 103. Kresnaia beschkovi Ganev. a. genitalia, b. signum. 104. Agriopis leucophaearia (Denis & Schiffermüller). 105. Agriopis bajaria (Denis & Schiffermüller).

727

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 106. Agriopis aurantiaria (Hübner). a. genitalia, b. signum. 107. Agriopis budashkini Kostjuk. a. genitalia, b. signum. 108. Agriopis marginaria (Fabricius). a. genitalia, b. signum.

728

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 109. Calamodes occitanaria (Duponchel). 110. Calamodes melanaria Oberthür. 111. Paradarisa consonaria (Hübner). a. genitalia, b. signum.

729

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 112. Larerannis orthogrammaria (Wehrli). a. genitalia, b. signum. 113. Chemerina caliginearia (Rambur). a. genitalia, b. signum. 114. Fagivorina arenaria (Hufnagel). a. genitalia, b. signum.

730

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 115. Spartopteryx kindermannaria Staudinger. 116. Athroolopha pennigeraria (Hübner). 117. Athroolopha chrysitaria (Hübner).

731

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 118. Eurranthis plummistaria (Villers). 119. Nychiodes obscuraria (Villers). 120. Nychiodes ragusaria Millière.

732

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 121. Nychiodes andalusiaria Millière. 122. Nychiodes notarioi Expósito. 123. Nychiodes hispanica Wehrli. a. genitalia, b. variation of lamella postvaginalis.

733

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 124. Nychiodes waltheri Wagner. a. genitalia. b. genitalia (holotype of cuencaensis Leraut syn. nov.) 125. Nychiodes amygdalaria (Herrich-Schäffer). 126. Nychiodes dalmatina Wagner.

734

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 127. Phaselia serrularia (Eversmann). a. genitalia, b. signum. 128. Phaselia algiricaria aragona Wehrli. a. genitalia, b. signum.

735

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 129. Ekboarmia atlanticaria (Staudinger). a. genitalia, b. signum. 130. Ekboarmia sagnesi Dufay. a. genitalia, b. signum. 131. Ekboarmia miniaria Skou, Stüning & Sihvonen. a. genitalia, b. ostium bursae and adjacent structures.

736

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 132. Ecleora solieraria (Rambur). a. genitalia, b. ostium bursae and adjacent structures. 133. Afriberina tenietaria (Staudinger). a. genitalia, b. ostium bursae and adjacent structures. 134. Afriberina salemae Skou & Sihvonen. a. genitalia, b. ostium bursae and adjacent structures.

737

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 135. Sardocyrnia bastelicaria (Bellier). 136. Sardocyrnia fortunaria (Vázquez).

738

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 137. Selidosema brunnearia (Villers). a. genitalia, b. signum. 138. Selidoseman plumaria (Denis & Schiffermüller). a. genitalia, b. signum. 139. Selidosema pyrenaearia (Boisduval). a. genitalia, b. signum.

739

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 140. Selidosema taeniolaria (Hübner). a. genitalia, b. signum. 141. Selidosema ambustaria (Geyer). a. genitalia, b. signum. 142. Selidosema erebaria postdentaria (Rothschild).

740

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 143. Peribatodes rhomboidaria (Denis & Schiffermüller). a. genitalia, b. ductus bursae. 144. Peribatodes perversaria (Boisduval). a. genitalia, b. ductus bursae. 145. Peribatodes omalensis Rajaei, Skou & Hausmann. a. genitalia, b. ductus bursae.

741

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 146. Peribatodes umbraria (Hübner). a. genitalia, b. ductus bursae. 147. Peribatodes ebusaria Vallhonrat. a. genitalia, b. ductus bursae. 148. Peribatodes secundaria (Denis & Schiffermüller). a. genitalia, b. ductus bursae.

742

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 149. Peribatodes correptaria (Zeller). a. genitalia, b. ductus bursae. 150-1. Peribatodes aragonis (Wehrli). a. genitalia, b. ductus bursae. 150-2. Peribatodes powelli (Oberthür). Not found in Europe, illustrated for comparison. a. geni­ talia, b. ductus bursae. 151. Peribatodes ilicaria (Geyer). a. genitalia, b. ductus bursae.

743

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 152. Alcis repandata (Linnaeus). 153. Alcis deversata (Staudinger). 154. Alcis jubata (Thunberg).

744

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 155. Deileptenia ribeata (Clerck). a genitalia, b. signum. 156. Paraboarmia viertlii (Bohatsch). a. genitalia, b. signum. 157. Adactylotis gesticularia (Hübner). a. genitalia, signum. 158. Adactylotis contaminaria (Hübner). a. genitalia, b. signum.

745

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 159. Pseudocoremia suavis Butler. a. genitalia, b. signum. 160. Parectropis similaria (Hufnagel). a. genitalia, b. signum. 161. Aethalura punctulata (Denis & Schiffermüller). a. genitalia, b. signum.

746

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 162. Ascotis selenaria (Denis & Schiffermüller). a. genitalia, b. signum. 163. Cleora cinctaria (Denis & Schiffermüller). a. genitalia, b. signum. 164. Ectropis crepuscularia (Denis & Schiffermüller).

747

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 165. Arichanna melanaria (Denis & Schiffermüller). 166. Bupalus piniaria (Linnaeus). 167. Megametopon grisolaria (Eversmann). Female was not available to GME project.

748

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 168. Tephronia sepiaria (Hufnagel). 169. Tephronia nuragica Fiumi et al. 170. Tephronia codetaria (Oberthür) subsp. sicula (Wehrli).

749

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 171. Tephronia lhommaria (Cleu) subsp. cleui Wehrli. a. genitalia (France), b. genitalia (Italy, identification requiring confirmation). 172. Tephronia cyrnea (Schawerda). a. genitalia (Corsica), b. genitalia (Sardinia).

750

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 173. Tephronia lepraria Rebel. 174. Tephronia psyloritaria (Reisser). 175. Tephronia theophilaria Hausmann. 176. Tephronia espaniola (Schawerda).

751

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 177. Eumannia oppositaria (Mann). a. genitalia, b. signum. 178-1. Eumannia oranaria (Staudinger). a. genitalia, b. signum. 178-2. Eumannia oranaria subsp. cebennaria (Chrétien). a. genitalia, b. signum.

752

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 179. Phyllometra culminaria (Eversmann). a. genitalia, b. signum. 180. Phyllometra gracilaria Boisduval. a. genitalia, b. signum. 181. Ithysia pravata (Hübner).

753

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 182. Leucobrephos middendorfii (Ménétriés). 183. Myinodes constantina (Hausmann). 184. Kuchleria menadiara (Thierry-Mieg).

754

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 185. Idaea barbuti Tautel & Lévêque. Tergum A8 removed (photo: A. Lévêque). 186. Idaea textaria (Lederer). See female genitalia figure in Vol. 2 fig. 49b. 187. Idaea lobaria (Chrétien). Tergum A8 moved to the right. 188. Idaea gelbrechti Hausmann.

755

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 189. Oar pratana (Fabricius). See female genitalia figure in Vol. 2 fig. 115b. 190. Scopula drenowskii Sterneck. 191. Scopula donovani (Distant).

756

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 192. Nothocasis rosariae Scalercio et al. a. genitalia, b. antrum (photo: S. Scalercio, reproduced with permission from copyright holder). 193. Perizoma juracolaria (Wehrli). a. genitalia, b. signum. 194. Perizoma barrassoi Zahm et al. a. genitalia, b. signum.

757

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 195. Eupithecia carpophilata Staudinger. 196. Eupithecia dubiosa Dietze. 197. Eupithecia furcata Staudinger.

758

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 198. Eupithecia gypsophilata Skou et al. 199. Eupithecia pindosata Weidlich. 200. Eupithecia goossensiata Staudinger.

759

FEMALE GENITALIA (Scale bar = 1.0 mm, specimen details are shown in Appendix 2) 201. Ourapteryx niveiscythes Zolotuhin. (photo: V. Zolotuhin, reproduced with permission from copyright holder). 202. Dyscia govii Fiumi & Flamigni. a. genitalia, b. genitalia (variant) (both photos: G. Fiumi, reproduced with permission from copyright holder).

760

Appendix 1. Data of the microscopic preparations shown in male genitalia plates 1. Odontognophos dumetata (Treitschke) a-d. genitalia, aedeagus, vesica, valva apex. Croatia: Dubrovnik-Neretva district, 1.7 km SSE Oslje, 215 m, 5.x.2015, slide 2302 Sihvonen (coll. Skou). 2. Odontognophos margaritata (Zerny) a-d. genitalia, aedeagus, vesica, valva apex. Spain: Prov. Teruel, Albarracin, Val de Vecar, 1200 m, 18.ix.2002, slide 2299 Sihvonen (coll. Skou). 3. Odontognophos zacharia (Staudinger) a-d. genitalia, aedeagus, vesica, valva apex. Greece: Prov. Fthiótida, Parnassos Ski Centre, 1750 m, 6.ix.2008, slide 2307 Sihvonen (coll. Skou). 4. Dicrognophos sartata (Treitschke) a. genitalia, Bulgaria: Pazardzhik, Bessa-parski Ridove Hills, 13.ix.1985, slide Sihvonen 2072 (coll. Skou). b-c. aedeagus, vesica. Italy: Puglia, 7 km N Altamura, 475 m, 29.v.2016, slide Sihvonen 2202 (coll. Skou). d. cornutus, Italy: Calabria, Monte Pollino, 800 m, 26.ix.2014, slide Sihvonen 2070 (coll. Skou). 5. Dicrognophos siciliana Skou & Sihvonen a-c. genitalia, aedeagus, vesica. Italy: Sicily, 5 km SE of Castelluzzo, 300 m, 12.v.2003, slide Sihvonen 2092, paratype (coll. Skou). d. cornutus, Italy: Sicily, 5.8 km E Santo Stefano Quisquina, 1000 m, 23–24.ix.2014, slide Sihvonen 2069, paratype (coll. Skou). 6. Ortaliella perspersata (Treitschke) a-e. genitalia, aedeagus (lateral view), aedeagus (ventral view), vesica, uncus. Spain: Tarra­gona, 3 km N of Alcossebre, 300 m, 3−6.vi.2013, slide 2304 Sihvonen (coll. Sihvonen). 7. Apochima flabellaria (Hübner) a-b. genitalia, aedeagus. Greece: Athens, no date, slide Müller 1015 (coll. MNHU). c. vesica, Italy: Sardinia, Soleminis, 22.i.1987, slide Sihvonen 1405 (coll. ZMH) 8. Chondrosoma fiduciaria Anker a-b. genitalia, aedeagus. Austria: Neusiedler See, Zitzmannsdorfer Wiesen, 2.x.1976 e.o., slide Müller 534 (coll. Müller). c. vesica, Hungary: Budapest, 28.x.1945, slide Sihvonen 1390 (coll. ZMH) 9. Gnophos furvata (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germany: Thüringen, Kranichfeld, 14.vii.1979, slide SE–743 (coll. Erlacher). 10. Gnophos obfuscata (Denis & Schiffermüller) a-b. genitalia, aedeagus. Italy: Adamello, Rifugio Mandrone, 2450 m, 08.vii.1994, slide SE–745 (coll. Erlacher).

761

11. Charissa obscurata (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germany: Thüringen, Erfurt-Rhoda, 20.viii.1985, slide Erlacher 35–­1999 (coll. Erlacher). 12. Charissa crenulata (Staudinger) a-b. genitalia, aedeagus. Spain: Granada, Sierra Nevada, Ruta del Veleta, 2000 m, 23.vii.2003, slide SE-758 (coll. Skou). 13. Charissa italohelveticus (Rezbanyai-Reser) a-b. genitalia, aedeagus. Switzerland: Wallis, Raron, 700 m, 14.vii.1987, e.o., slide SE–748 (coll. Erlacher). 14. Charissa pullata (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germany: Thüringen, Rehberg near Blankenberg, 19.vi.1993, slide SE–750 (coll. Erlacher). 15. Charissa difficilis (Alphéraky) a-b. genitalia, aedeagus. Russia: Karachay-Cherkessia, Teberda, 27.iii.1977, slide BM–175 (coll. B. Müller). 16. Charissa turfosaria (Wehrli) a-b. genitalia, aedeagus. Russia: Altai, 26 km south of Seminski pass, ♀: 13–19.vi.2012, e.o., slide JH–253 (coll. Erlacher). 17. Charissa mutilata (Staudinger) a-c. genitalia, aedeagus, sclerotized prickly crest. Turkey, Prov. Trabzon, Doğu Karadeniz Mts., 7 km SE, Oviddağı Geçidi, 2000 m, ♀: 30.v.2000, e.o., slide DNATAX–2322/ Erlacher (coll. Gelbrecht). 18. Charissa peloponnesiaria Erlacher & Erlacher a-b. genitalia, aedeagus. Greece: Peloponnes, Parnon Mts., north east of Kosmas, 850 m, 11.ix.2004, slide SE–721 (coll. Erlacher). c. sclerotized prickly crest, Greece: Peloponnesus, Chelmos Mts., above Kalavrita, 2200 m, 04.ix.1983, slide Sircoulomb–3066 (coll. Sircoulomb). 19. Charissa pentheri (Rebel) a-c. genitalia, aedeagus, sclerotised prickly crest. Macedonia: Ochrid–Petrina, 07.–16. vi.1939, slide SE–724 (coll. ZSM). 20. Charissa assoi (Redondo & Gastón) a-b. genitalia, aedeagus. Spain: Prov. Granada, Sierra Nevada, Road to Veleta, 2000 m, 8.vii.1971, slide SE–103 (coll. ZSM). 21. Charissa corsica (Oberthür) a-b. genitalia, aedeagus. France: Corsica, 21.vi., slide SE–106 (coll. ZFMK). 22. Charissa staudingeri (Wnukowsky) a-b. genitalia, aedeagus. Greece: Tinos, 17.–26.ix.2001, slide SE–52 (coll. ZSM). 23. Charissa subtaurica (Wehrli) a-b. genitalia, aedeagus. Turkey: Cyprus, Mount Troodos, 600 m, v.2000, slide SE–118/ 2004 (coll. Gelbrecht).

762

24. Charissa variegata (Duponchel) a-b. genitalia, aedeagus. Greece: Corfu, Pantokratora, 911 m, 5.–19.vi.1999, slide SE– 119/ 2004 (coll. Gelbrecht). 25. Charissa mucidaria (Hübner) a-b. genitalia, aedeagus. Spain: Prov. Almeria, Cabo de Gata, e.o., ­Gelbrecht).

♀:

ii.1992 (coll.

26. Charissa ambiguata (Duponchel) a-b. genitalia, aedeagus. France: Embrun, 800 m, 27.vi.1988, slide JH–122 (coll. ­Gelbrecht). 27. Charissa predotae (Schawerda) a-b. genitalia, aedeagus. Spain: Miraflores near Madrid, 1200 m, e.o., slide DNATAX–2198/Erlacher (coll. Gelbrecht).

♀:

2.–3.v.1996,

28. Charissa onustaria (Herrich-Schäffer) a-b. genitalia, aedeagus. Turkmenistan: Kopet-Dag Mts., 15 km SE of Nochur, 1300– 1400 m, 28.iv.1991, slide CD–24 (coll. Skou). 29. Charissa remmi (Viidalepp) a-b. genitalia, aedeagus. Russia: Magadan Distr., Kulu, 11.vii.1980, slide SE SE–515, paratype (coll. Skou). 30. Charissa zeitunaria (Staudinger) a-b. genitalia, aedeagus Turkey: Southern Taurus Mts., Zeitun, 1897, slide SE–520, lectotype (coll. MNHB). 31. Charissa glaucinaria (Hübner) a-b. genitalia, aedeagus. Greece: Ioannina, Katarapass, 22.vii.1999, slide JH–97 (coll. E ­ rlacher). 32. Charissa pfeifferi (Wehrli) a-b. genitalia, aedeagus. Turkey, slide SE–08/2005 (coll. ZSM). 33. Charissa supinaria (Mann) a-b. genitalia, aedeagus. Germany: Thüringen, Kobersfels, ♀: 5.v.1993, slide SE–37/1999 (coll. Erlacher). 34. Charissa graecaria (Staudinger) a-b. genitalia, aedeagus. Greece: Hellas, Drama, 1700 m, Mt. Phalakron, above Volas, 17.viii.1985, slide SE– DNATAX–2303/Erlacher (coll. Erlacher). 35. Psodos alpinata (Scopoli) a-c. genitalia, aedeagus, juxta. Austria: North Tyrol, Karwendel Mts., Innsbruck, Nordkette, 2200 m, 5.viii.1970, slide CJ–83 (coll. ZSM). 36. Psodos canaliculata (Hochenwarth) a, c. genitalia, juxta. Switzerland: Canton of Valais, Riffelberg, 26.vii.1935, slide CJ–96 (coll. ZSM). b. aedoeagus, France: Alpes-Maritimes, Saint Martin Vésubie, 1.vii.1941, slide CJ–98 (coll. ZSM).

763

37. Psodos werneri Schawerda a-c. genitalia, aedeagus, juxta. Bosnia-Herzegovina: Balkan, Volujak, 2000 m, slide ­SE–716, lectotype (coll. NHMW). 38. Psodos coracina (Esper) a-c. genitalia, aedeagus, juxta. Austria: Salzburg, Glockner Group, Gleiwitzerhütte, 7.– 13.vii.1941, slide CJ–89 (coll. ZSM). 39. Psodos bentelii (Rätzer) a-c. genitalia, aedeagus, juxta. Switzerland: Canton of Valais, Zermatt, 7.viii.1912, slide CJ–92 (coll. ZSM). 40. Psodos noricana Wagner a-c. genitalia, aedeagus, juxta. Austria: North Tyrol, Ötztal Alps, Obergurgl, 2100 m, vii.1956, slide CJ–106 (coll. ZSM). 41. Psodos belzebuth Praviel a, c. genitalia, juxta. France: Alpes-de-Haute-Provence, Colmars, 29.vi.1924, slide CJ–52 (coll. ZSM). b. aedeagus, France: Alpes-Maritimes, Tete Chaudon, Col Boira, 8.viii.1986, slide ­CJ–65 (coll. TLMF). 42. Psodos pyrenaica Schawerda a-c. genitalia, aedeagus, juxta. France: Pyrénées c., Pic du Midi, [without date], slide Rajaei 2176/2018 (coll. ZSM). 43. Psodos alpmaritima Wehrli a-c. genitalia, aedeagus, juxta. France: Alpes-Maritimes, Cima Argentera, 30.vii.1923, slide G259, lectotype (coll. ZFMK). 44. Psodos alticolaria Mann a-c. genitalia, aedeagus, juxta. Austria: North Tyrol, Ötztal Alps, Rotmoostal, 2400 m, 1.viii.1994, slide CJ–59 (coll. Erlacher). 45. Psodos wehrlii Vorbrodt a-c. genitalia, aedeagus, juxta. Switzerland: Canton of Valais, Mettelhorn, 11.vii.1921, slide CJ–108 (coll. ZFMK). 46. Psodos baldensis Wolfsberger a-c. genitalia, aedeagus, juxta. Italy: Verona, Monte Baldo, 2200 m, vii.1966, slide CJ–99), paratype (coll. TLMF). 47. Psodos spitzi Rebel a-c. genitalia, aedeagus, juxta. Slovenia: Triglav, vii.1922, slide CJ–79 (coll. ZSM). 48. Psodos quadrifaria (Sulzer) a-c. genitalia, aedeagus, juxta. Austria: Styria, Ennstal Alps, Eisenerzer Reichenstein, 1700 m, 15.–31.vii.1925, slide CJ–103 (coll. ZSM). 49. Sciadia tenebraria (Esper) a, c-d. genitalia, vesica, juxta. Austria: Innsbruck, Nordkette, 14.viii.1970, 2200 m, slide S ­ ihvonen 2104 (coll. TLMF). b. aedeagus, Italy: Province Veneto, Below Rif. Di Giussani, 2450 m, 31.vii.2013, slide Sihvonen 2146 (coll. Skou). 764

50. Sciadia septaria (Guenée) a. genitalia. France: Pic du Midi, 2700–3000 m, 24.vii.1950, slide Huemer G1442 (coll. ZSM). b-d. aedeagus, vesica, juxta. Spain: c. 20 km W of Andorra, Vall Ferrera, 30.vii.1957, slide Sihvonen 2139 (coll. Skou). 51. Sciadia innuptaria (Herrich-Schäffer) a-d. genitalia, aedeagus, vesica, juxta. Austria: Dachstein, Simonyhütte, without date, slide Sihvonen 2107 (coll. TLMF). 52. Sciadia slovenica Leraut a-d. genitalia, aedeagus, vesica, juxta. Slovenia: Triglav, 31.vii.1906, slide Sihvonen 2108 (coll. TLMF). 53. Sciadia dolomitica Huemer & Hausmann a-d. genitalia, aedeagus, vesica, juxta. Italy: Pale di San Martino, Rif. Pedrotti alla Rosetta, 2550 m, 6.viii.2013, slide Sihvonen 2106 (coll. TLMF). 54. Sciadia zelleraria (Freyer) a-d. genitalia, aedeagus, vesica, juxta. Italy: Völs am Schlern, 2300–2440 m, 27.vii.2007, slide Sihvonen 2122 (coll. TLMF). 55. Sciadia unicoloraria (Staudinger) a-d. genitalia, aedeagus, vesica, juxta. France: Punta Marguareis, western slope, 2100– 2200 m, 21–23.vii.1990, slide Sihvonen 2137 (coll. TLMF). 56. Elophos caelibaria (Heydenreich) a-d. genitalia, aedeagus, vesica, juxta. Austria: Kärnten, Glockner-Hochtor, 2400–2600 m, 5.vii.1988 (ex pupa), slide Sihvonen 2126 (coll. TLMF). 57. Elophos zirbitzensis (Pieszcek) a-d. genitalia, aedeagus, vesica, juxta. Austria: Zirbitzkogel, Lavanttaler Alpen, 2200 m, 27.vi.1949, slide Sihvonen 2135 (coll. TLMF). 58. Elophos operaria (Hübner) a-d. genitalia, aedeagus, vesica, juxta. Austria: Rax, 1800 m, 13.vi.1967, slide Sihvonen 2116 (coll. TLMF). 59. Elophos andereggaria (de la Harpe) a-d. genitalia, aedeagus, vesica, juxta. Switzerland: LaQuintal, 2000 m, 12.vii.1958, slide Sihvonen 2134 (coll. TLMF). 60. Yezognophos dilucidaria (Denis & Schiffermüller) a-d. genitalia, aedeagus, vesica, juxta. Italy: Dolomites, Wolkenstein in Gröden, 1900 m, 11.vii.2015, slide Sihvonen 2112 (coll. Sihvonen). 61. Yezognophos sproengertsi (Püngeler) a-d. genitalia, aedeagus, vesica, juxta. France: Mercantour, Col de la Cayolle, 1950 m, 7.vii.2012, slide Sihvonen 2132 (coll. TLMF). 62. Yezognophos vittaria (Thunberg) a-d. genitalia, aedeagus, vesica, juxta. Finland: Kaunispää, 4.vii.1994, slide Sihvonen 2080 (coll. Sihvonen).

765

63. Yezognophos serotinaria serotinaria (Denis & Schiffermüller) 63-1a,d. genitalia, apex of juxta arm. Austria: Rangger Köpfl, 1700 m, 4.vii.1937, slide Sihvonen 2174 (coll. TLMF). 63-1b-c. aedeagus, vesica. Italy: Vinschgau, Schnalstal, 1300 m, 2.iv.1986 (ex ovo), slide Sihv­ onen 2114 (coll. Sihvonen) (lateral view). Yezognophos serotinaria dognini (Thierry-Mieg) 63-2a-d. genitalia, aedeagus, vesica, apex of juxta arm. France: Hautes-Pyrénées, Lac d’Orédon, 1900 m, 24.vii.2014, slide Sihvonen 2185 (coll. Skou). Yezognophos serotinaria serotinoides (Wehrli) 63-3a. genitalia, Switzerland: Wallis, Leuk-Guttet, 1000 m, 18.ii.1993 (ex ovo, female 1.viii. 1992), slide Sihvonen 2130 (coll. TLMF). 63-3b-d. aedeagus, vesica, apex of juxta arm. Switzerland: Valais, Saastal, 1900 m, 13–14. vii. 2015, slide Sihvonen 2183 (coll. Skou). 64. Crocota tinctaria (Hübner) a-c. genitalia, aedeagus, apex of aedeagus. Italy: Alpi Cozie, Demonte, Colle Valcavera, 2420 m, 2.viii.2010, slide Sihvonen 2101 (coll. TLMF). 65. Crocota peletieraria (Duponchel) a-c. genitalia, aedeagus, apex of aedeagus. France: L’Hospitalet, Ariege, 13.vii.1946, 1500–1600 m, slide Sihvonen 2103 (coll. TLMF). 66. Crocota pseudotinctaria Leraut a, c. genitalia, vesica. Switzerland: Leukerbad, Torrent, 2300 m, 24.vii.1993 (ex ovo), slide Sihvonen 2097 (coll. Sihvonen). b, d. aedeagus, apex of aedeagus. Switzerland: Wallis, Simplonpass, 2050 m, viii.1985 (ex ovo), slide Sihvonen 2121 (coll. TLMF). 67. Crocota niveata (Scopoli) a-b, d. genitalia, aedeagus, apex of aedeagus. Austria: Saualpe, Hohenwart, 1780 m, 12.vii. 2011, slide Sihvonen 2099 (coll. Sihvonen). c. vesica, Austria: Salzburg, Thomatal, 1950 m, 8.ix.2007 (ex ovo), slide Sihvonen 2158 (coll. TLMF). 68. Synopsia sociaria (Hübner) a-b, d. genitalia, aedeagus, valva. Bulgaria: Varna/Kranevo, 4.viii.1984, slide Müller 351 (coll. Müller). c. vesica, Italy: Calabria, 6 km N Civita, 1030 m, 11.viii.2006, slide Sihvonen 2276 (coll. ZSM). 69. Phthonandria strictaria (Lederer) a-c. genitalia, aedeagus, vesica. Russia: Ural Mountains, Orenburg Region, Kyvandyk Village, 1−11.vi.1995, slide 2229 Sihvonen (coll. Skou). d. uncus (dorsal view, Mongolia: Zavhan Aimak, Tsangaanhayrhan somon env., 1800 m, 30.vi.2004, slide 2227 Sihvonen (coll. Skou). 70. Menophra abruptaria (Thunberg) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Greece: Crete, Drosia, 6.iv.2005, slide Müller 374 (coll. Müller).

766

71. Menophra japygiaria (O. Costa) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Italy: Sardinia, Olbia, San Pantaleo, 13.iv.2007, slide Müller 370 (coll. Müller). 72. Menophra berenicidaria (Turati) a-b. genitalia, aedeagus. Greece: Crete, Gonies/Iraklion, 15.iv.1999, slide Müller 369 (coll. Müller). c-d. aedeagus, apical half of valva. Turkey: Antalya, 16.v.1996 e.o., slide Müller 247 (coll. Müller). 73. Menophra harterti (Rothschild) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Algeria: Guelt es Stel, 15.–23.ix.1933, slide Müller 1041 (coll. ZSM). 74. Menophra thuriferaria (Zerny) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Spain:Teruel, Albarracin 1150 m, 15.vi.2007, slide Müller 361 (coll. Müller). 75. Menophra nycthemeraria (Geyer) a-c. genitalia, aedeagus, apex of aedeagus. France: Provence, Choudon-Noranthe, 800 m, 24.iii.1996 e.o., slide Müller 376 (coll. Müller). 76. Mephra annegreteae Skou a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Spain: Almeria, Tabernas, 25.v.2006, paratype, slide Müller 375 (coll. Müller). 77. Cleorodes lichenaria (Hufnagel) a-d. genitalia, aedeagus, apex of aedeagus, uncus and gnathos. Denmark: Arnager, 13.viii.1979, slide Müller 567 (coll. Müller). 78. Zernyia granataria (Staudinger) a-d. genitalia, aedeagus, apex of aedeagus, uncus and gnathos. Spain:Teruel, Albarracin 1100 m, 4.ix.2008, slide Müller 462 (coll. Müller). 79. Megalycinia serraria (Costa) a-d. genitalia, aedeagus, vesica, lobe between valva base and gnathos (potentially arising from anterior margin of tegumen, see arrow in figure a). Italy: Calabria, Mte. ­Cocuzzo, 1150 m, 4.ix.1997, slide 2274 Sihvonen (coll. ZSM). 80. Dasypteroma thaumasia Staudinger a-b. genitalia, aedeagus with juxta. Spain: Madrid, Aranjuez, 7.x.2008, slide Müller 580 (coll. Müller). c, e. aedeagus, gnathos. Spain: Madrid, Aranjuez, 7.x.2008, slide Müller 568 (coll. Müller). d. vesica, [handwritten label, very difficult to read] Guileda [or Juileda?], 4.x.1933, slide Sihvonen 2278 (coll. ZSM). 81. Microbiston lanaria (Eversmann) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Russia: “Ural-Steppen”, date unknown, slide Müller 1067 (coll. ZFMK). 82. Apocheima hispidaria (Denis & Schiffermüller) a-d. genitalia, aedeagus, apex of aedeagus, apical half of valva. Germany: Brandenburg, Nauen, Paulinenaue, 12.iii.2007, slide Müller 505 (coll. Müller). 767

83. Phigalia pilosaria (Denis & Schiffermüller) a-b, d. genitalia, aedeagus, central part of valva. Germany: Thuringia, Erfurt, Steiger, 26.i.1974, slide Müller 484 (coll. Müller). c. vesica, Finland: Savonlinna, 29–30.iv.1998, slide Sihvonen 1316 (coll. Sihvonen). 84. Hypomecis roboraria (Denis & Schiffermüller) a-b, d. genitalia, aedeagus, central part of valva. Germany: Brandenburg, Grünheide, Löcknitztal, 3.vi.2011, slide Müller 433 (coll. Müller). c. vesica, Finland: Enonkoski, 8–10.vii.1996, slide Sihvonen 2086 (coll. Sihvonen). 85. Hypomecis punctinalis (Scopoli) a-b, d. genitalia, aedeagus, central part of valva. Germany: Brandenburg, Grünheide, Löcknitztal, 3.vi.2011, slide Müller 430 (coll. Müller). c. vesica, Finland: Inkoo, Gölesudden, 19.–26.v.2005, slide Sihvonen 2783 (coll. Sih­ vonen). 86. Ematurga atomaria (Linnaeus) a-b, d. genitalia, aedeagus, central part of valva. Germany: Thuringia, Ilmenau, 500 m, 9.v.1975, slide Müller 560 (coll. Müller). c. vesica, Finland: Rantasalmi, 28.v.1996, slide Sihvonen 1865 (coll. Sihvonen). 87. Lycia hirtaria (Clerck) a-b. genitalia, aedeagus. Germany, Brandenburg, Ahrensfelde, 19.iv.1970, slide Müller 498 (coll. Müller). c-d. vesica, cornuti. Finland: Savonlinna, 15–18.v.1997, slide Sihvonen 1322 (coll. Sih­ vonen). 88. Lycia necessaria (Zeller) a-b. genitalia, aedeagus. Georgia: Tbilisi, Mtatsminda, 26.iii.1989 e.o., slide Müller 574 (coll. Müller). 89. Lycia lapponaria (Boisduval) a-c. genitalia, aedeagus, apex of aedeagus. Latvia: “Livonia”, without date, slide Müller 1010 (coll. MNHU). 90. Lycia isabellae (Harrison) a-c. genitalia, aedeagus, apex of aedeagus. Czech Republic: Moravia, Ždánice, 30.iii.1987, slide Müller 531 (coll. Müller). 91. Lycia pomonaria (Hübner) a-b. genitalia, aedeagus. Germany: Brandenburg, Nauen, Paulinenaue, 12.iii.2007, slide Müller 507 (coll. Müller). c-d. vesica, cornuti. Finland: Savonlinna, 7.v.1997, slide Sihvonen 1318 (coll. Sihvonen). 92. Lycia zonaria (Denis & Schiffermüller) a-c. genitalia, aedeagus, apex of aedeagus. Germany: Brandenburg, Kyritz, Dreetz, 21.iii.1991, slide Müller 494 (coll. Müller). 93. Lycia alpina (Sulzer) a-c. genitalia, aedeagus, apex of aedeagus. Italy: South Tyrol, Paneveggio, 20.iv.1913, slide Müller 530 (coll. Müller).

768

94. Lycia graecarius (Staudinger) a-b. genitalia, aedeagus. Croatia: Krk Island, Punet, 31.iii.2003, slide Müller 529 (coll. Müller). 95. Lycia florentina (Stefanelli) a-b. genitalia, aedeagus. Italy: Roma, 15.iii.1959, slide Müller 528 (coll. Müller). 96. Biston strataria (Hufnagel) a-b, d. genitalia, aedeagus, uncus and gnathos and apex of juxta. Germany: Berlin, Rosenthal, 9.iii.1986 e.l., slide Müller 546 (coll. Müller). c. vesica, Finland: Lemland, 29.iv-6.v.1995, slide Sihvonen 1324 (coll. Sihvonen). 97. Biston rosenbaueri Müller a-c. genitalia, aedeagus, uncus and gnathos and apex of juxta. Croatia: Krk Island, ­Vrhnik, 26.iii.2005, slide Müller 1032 (coll. E. Friedrich). 98. Biston achyra Wehrli a-c. genitalia, aedeagus, uncus and gnathos and apex of juxta. Greece: Samos, Vour­ liotes, 18.iii.2015, slide Müller 1036 (coll. D. Fritsch). 99. Biston betularia (Linnaeus) a-b, d. genitalia, aedeagus, uncus and gnathos. Germany: Saxony, Erzgebirge, Schlettau, 3.vii.2014, slide Müller 489 (coll. Müller). c. vesica, Finland: Lemland, 20–27.vii.1996, slide Sihvonen 1325 (coll. Sihvonen) 100. Erannis defoliaria (Clerck) a-b. genitalia, aedeagus. Germany: Brandenburg, Tiefensee, 10.x.1980, slide Müller 582 (coll. Müller). c. vesica, Finland: Savonlinna, 7–13.x.1997, slide Sihvonen 1331 (coll. Sihvonen) 101. Erannis ankeraria (Staudinger) a-b. genitalia, aedeagus. Slovenia: Podgorski Kras, Podgorje, 31.iii.2004, slide Müller 562 (coll. Müller). 102. Erannis declinans (Staudinger) a-b. genitalia, aedeagus. “Amasia”, no date, slide Müller 1026 (coll. MNHU). 103. Kresnaia beschkovi Ganev a-c. genitalia, aedeagus, apex of aedeagus. Bulgaria: Pirin mts., Kresna gorge, 17.xi.2014, slide Müller 1051 (coll. St. Beshkov). 104. Agriopis leucophaearia (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germany: Brandenburg, Nauen, Brieselang, 4.iv.1987, slide Müller 475 (coll. Müller). 105. Agriopis bajaria (Denis & Schiffermüller) a-b. genitalia, aedeagus. Italy: Sardinia, Olbia, Palau, 22.xii.2007 e.l., slide Müller 478 (coll. Müller). 106. Agriopis aurantiaria (Hübner) a-b. genitalia, aedeagus. Germany: Brandenburg, Tiefensee, 18.x.1980, slide Müller 474 (coll. Müller). c. vesica, Finland: Helsinki, 5.x.1995, slide Sihvonen 1330 (coll. Sihvonen).

769

107. Agriopis budashkini Kostjuk a-b. genitalia, aedeagus. Greece: Crete, Asfendos, 8.xi.2015 e.l., slide Müller 563 (coll. Müller). 108. Agriopis marginaria (Fabricius) a-b. genitalia, aedeagus. Germany: Thuringia, Arnstadt, Neusiss, 21.iii.1971, slide Müller 472 (coll. Müller). 109. Calamodes occitanaria (Duponchel) a-b. genitalia, aedeagus. France: Alpes-de-Haute-Provence, Digne, 6.x.1993, slide Müller 427 (coll. Müller). 110. Calamodes melanaria Oberthür a-b. genitalia, aedeagus. Italy: Calabria, Fiumara Trionto (CS), 21.x.1999, slide Müller 1049 (coll. ZSM). 111. Paradarisa consonaria (Hübner) a-b. genitalia, aedeagus. Germany: Thuringia, Thuringian Forest, Martinroda, 9.v.1976, slide Müller 398 (coll. Müller). 112. Larerannis orthogrammaria (Wehrli) a-c. genitalia, aedeagus, apical half of valva. Russia: Far East (Primorje), Suchanskij ­Rudnik, 20.x.1933, slide Müller 1057 (coll. ZFMK). 113. Chemerina caliginearia (Rambur) a-b. genitalia, aedeagus. France: Gironde, Villenave-d’Ornon, 10.iii.1955 e.l., slide Müller 487 (coll. Müller). 114. Fagivorina arenaria (Hufnagel) a-c. genitalia, aedeagus, apical half of valva. Croatia: Istria, Ćićarija mts., Buzet, 800– 1200 m, 16.vii.1986, slide Müller 439 (coll. Müller). 115. Spartopteryx kindermannaria Staudinger a-b. genitalia, aedeagus. Russia: Republic Altai, Altai mts., Aktash, 1480 m, 16.vi.2012, slide Müller 446 (coll. Müller). 116. Athroolopha pennigeraria (Hübner) a-b. genitalia, aedeagus. Spain: Almeria, Gérgal, 19.ix.2004 e.o., slide Müller 557 (coll. Müller). 117. Athroolopha chrysitaria (Hübner) a-b. genitalia, aedeagus. Italy: Sicilia, Prov. Palermo, S. Martino, 20.v.1905, slide Müller 1048 (coll. MNHU). 118. Eurranthis plummistaria (Villers) a-c. genitalia, aedeagus, apical half of valva. France: Alpes-de-Haute-Provence, Digne, 19.iv.1992, slide Müller 509 (coll. Müller). 119. Nychiodes obscuraria (Villers) a-b. genitalia, aedeagus with everted vesica. Italy: Südtirol, Vinschgau, 16.vii.1976, slide Rajaei 2007/2016 (coll. Müller). 120. Nychiodes ragusaria Millière a-b. genitalia, aedeagus with everted vesica. Italy: Sardinia, 2008, vii.2009 ex ovo, slide Rajaei 2100/2017 (coll. Löbel). 770

121. Nychiodes andalusiaria Millière a-b. genitalia, aedeagus. Portugal: Manteigas, slide Fazekas 2591 (coll. ZFMK). 122. Nychiodes notarioi Expósito a-b. genitalia, aedeagus. Spain: Teruel, SE Sierra de Javalambre, 1700–1800 m; 24.vii.78, slide Rajaei 2150/2017 (coll. SMNS). 123. Nychiodes hispanica Wehrli a-b. genitalia, aedeagus. Spain: Granada, N 37°04’28” W 003°01’23”, Sierra Nevada, 1840 m, 12.vii.2009, slide Rajaei 2009/2017 (coll. SMNS). c. Paralectotype. Variation of valval ornamentation, Spain: Sierra Nevada, 10.vii.1926, slide Wehrli 4052 (coll. ZFMK). d. Variation of valval ornamentation, Spain: Malaga, 25 km SW Ronda Algatocin, 865 m, 31.vii.1999, slide Rajaei 2013/2016 (coll. Müller). e. Variation of valval ornamentation, Spain: Malaga, 25 km SW Ronda Algatocin, 865 m, 31.vii.1999, slide Rajaei 2164/2017 (coll. Müller). f. Variation of valval ornamentation, Spain: Malaga, 25 km SW Ronda Algatocin, 865 m, 31.vii.1999, slide Rajaei 2165/2017 (coll. Müller). g. Variation of valval ornamentation, Moroco: Hoher Atlas, Oukaimeden Umg. Zedernwald, N31°13’58’’ E07°49’13’’, vii.2011 e. o., slide Rajaei 2142/2017 (coll. Stadie). h. Variation of valval ornamentation, Spain: Andalusia, Granada, N37°04’28’’, W3°01’23’’, Sierra Nevada, 1840 m, 15.vii.2009, slide Rajaei 2163/2017 (coll. Müller). 124. Nychiodes waltheri Wagner a-b. genitalia, aedeagus. Bulgaria: Rhodopi Mts. Studen Kladenets, 250 m, 23.v.1990, slide Rajaei 2118/2017 (coll. Skou). 125. Nychiodes amygdalaria (Herrich-Schäffer) a-b. genitalia, aedeagus. Turkey: Aksehir, 1000 m, 20.vii.–23.viii.1968, slide Rajaei 2089/ 2017 (coll. Smns). 126. Nychiodes dalmatina Wagner a-b. genitalia, aedeagus. Greece: Thessaly, 5 km S Aghiokambos, 5 m, 31.v.–1.vi.2013, slide Rajaei 2115/2017 (coll. Skou). c. valva (arrow shows the tiny wart on sacculus part of valva), Bulgaria: Pirin Mt., Lilja­ novo, 550 m, 14.vii.1981 ex ovo, slide Rajaei 2000/2016 (coll. Müller). 127. Phaselia serrularia (Eversmann) a-b. genitalia, aedeagus. Russia: Orenburg district, Prokorska, 21.vi.1999, slide Müller 1060 (coll. P. Skou). 128. Phaselia algiricaria aragona Wehrli a-b. genitalia, aedeagus. Spain: Teruel, Albarracin, 1200 m, 25.–26.vi.1991, slide Müller 527 (coll. Müller). c-d. aedeagus, uncus and gnathos. Spain: Teruel, Albarracin, 1200 m, 1.vii.1991, slide Müller 570 (coll. Müller). 129. Ekboarmia atlanticaria (Staudinger) a-d. genitalia, aedeagus, vesica, juxta. Portugal: Salema, 40 m, 19.iv.2007, slide Sihvonen 2053 (coll. Skou).

771

130. Ekboarmia sagnesi Dufay a-b. genitalia, aedeagus. Spain: Leon, Valle de Luna, 1200 m, 2.vii.2011, slide Sihvonen 2157 (coll. Skou). c-d. vesica, juxta. France: Valle de la Romanche, 1200 m, 22–23.vi.1999 slide Sihvonen 2095 (coll. Skou). 131. Ekboarmia miniaria Skou, Stüning & Sihvonen a-d. genitalia, aedeagus, vesica, juxta. Portugal: Grandola, Ameiras de Baixo, 20.vi.2011, holotype, slide Sihvonen 2050 (coll. Skou). 132. Ecleora solieraria (Rambur) a-c. genitalia, aedeagus, apical half of valva. Spain: Teruel, Albarracin, 1200 m, 12.vi. 2007, slide Müller 526 (coll. Müller). 133. Afriberina tenietaria (Staudinger) a, d. genitalia, apex of valva. Spain: Prov. Huelva, Matalascanas, 20 m, 16.iv.2007, slide Sihvonen 2243 (coll. Skou). b-c. aedeagus, vesica. Spain: Prov. Teruel, 4 km W Albarracin, 1185 m, 7.viii.2007, slide Sihvonen 2147 (coll. Skou). 134. Afriberina salemae Skou & Sihvonen a-b. genitalia, aedeagus. Portugal: Prov. Algarve, Salema, 40 m, 19.iv.2007, slide 2247 Sihvonen (coll. Skou), paratype. c-d. vesica, apex of valva. Portugal: Algarve, Salema, 55 m, 22.iv.2017, slide Sihvonen 2240 (coll. Skou), paratype. 135. Sardocyrnia bastelicaria (Bellier) a-c. genitalia, aedeagus, apical half of valva. France: Corsica, 1906, slide Müller 1002 (coll. MNHU). 136. Sardocyrnia fortunaria (Vázquez) a-c. genitalia, aedeagus, apical half of valva. Spain: Teruel, Albarracin, 1200 m, 5.ix.2006, slide Müller 448 (coll. Müller). 137. Selidosema brunnearia (Villers) a-c. genitalia, aedeagus, apical half of valva. Danmark: Bornholm, 30.vi.1990 e.o., slide Müller 407 (coll. Müller). 138. Selidosema plumaria (Denis & Schiffermüller) a-c. genitalia, aedeagus, apical half of valva. Hungary: Budapest, Budaörs, 30.viii.1980, slide Müller 404 (coll. Müller). 139. Selidosema pyrenaearia (Boisduval) a-c. genitalia, aedeagus, apical half of valva. Spain: Teruel, Albarracin, 1350 m, 9.ix.2008, slide Müller 406 (coll. Müller). 140. Selidosema taeniolaria (Hübner) a-c. genitalia, aedeagus, apex of aedeagus. “Southern France”, 28.vi.2006 e.o., slide Müller 411 (coll. Müller). 141. Selidosema ambustaria (Geyer) a-c. genitalia, aedeagus, apex of aedeagus. Italy: Sicilia, Girgenti (now Agrigento), 3.viii.1910 e.o., slide Müller 1020 (coll. MNHU).

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142. Selidosema erebaria postdentaria (Rothschild) a-c. genitalia, aedeagus, apical half of valva. Algeria: Guelt es Stel, x.1929, slide Müller 1063 (coll. MNHU). 143. Peribatodes rhomboidaria (Denis & Schiffermüller) a-c, e. genitalia, aedeagus, vesica, setal comb (male abdominal sternite 3). Germany: Berlin-­ Rosenthal, 27.v.2002, slide Rajaei 1996/2016 (coll. Müller). d. uncus (lateral view). Germany: Berlin-Rosenthal, 12.vi.2014, slide Müller 466 (coll. Müller). 144. Peribatodes perversaria (Boisduval) a-d. genitalia, aedeagus, vesica, uncus (lateral view). Switzerland: Helvetia/Valesia, Visperterminen, 28.vi.1995 e. o., slide Rajaei 1978/2016 (coll. Müller). 145. Peribatodes omalensis Rajaei, Skou & Hausmann a-d. genitalia, aedeagus, vesica, uncus (lateral view). Greece: Crete, 10.5 km SW Omalos Plateau, 18.vi.2014, 760 m, slide Rajaei 2164/2017 Paratype (coll. Skou). 146. Peribatodes umbraria (Hübner) a-e. genitalia, aedeagus, vesica, uncus (lateral view), setal comb (male abdominal sternite 3). France: Corsica 42°05’25,8” N 009°12’09,2” E, 2 km SW Ghisoni, 730 m, 02.vi.2008, slide Rajaei 1983/2016 (coll. Müller). 147. Peribatodes ebusaria Vallhonrat a-c. genitalia, aedeagus, vesica. Spain: Ibiza, 4.2 km SW Sant Rafel, Puig des Fornas Coll des Rossellons, 220 m, 30.iii.–3.iv.2016, slide Rajaei 2178/2018 (coll. Skou). d. uncus (lateral view), photo modified from Vallhonrat, 2012, Fig. 4, Holotype. 148. Peribatodes secundaria (Denis & Schiffermüller) a-d. genitalia, aedeagus, vesica, uncus (lateral view). Austria: Kärnten 46°50’23” N, 014°43’09” E, Lavanttaler Alpen, Saualpe, 1335 m, 06.vii.2011, slide Rajaei 1992/2016 (coll. Müller). 149. Peribatodes correptaria (Zeller) a-d. genitalia, aedeagus, vesica, uncus (lateral view). Greece: Creta, N 35.11’, E 25.40’, Agios Nikolaos, Bucht von Istron, 01.vii.1994 e. o., slide Rajaei 1987/2016 (coll. Müller). 150. Peribatodes aragonis (Wehrli) a-d. genitalia, aedeagus, vesica, uncus (lateral view). Spain: Teruel, N 40°25’57”, W 001°24’03”, Serrania de Albarracin, 4 km NE Albarracin, 1100 m, 04.ix.2009 e. o., slide Rajaei 1980/2016 (coll. Müller). 151. Peribatodes ilicaria (Geyer) a-c, e. genitalia, aedeagus, vesica, setal comb (male abdominal sternite 3). Spain: Murcia, 640 m, Aledo W of Totana, 04.vi.2003, slide Rajaei 1990/2016 (coll. Müller). d. uncus (lateral view), Spain: Murcia, 640 m, Aledo W of Totana, 04.vi.2003, slide Rajaei 1989/2016 (coll. Müller). 152. Alcis repandata (Linnaeus) a-b. genitalia, aedeagus. Germany: Thuringia, Thuringian Forest, Ilmenau, 4.vi.1973 e.l., slide Müller 423 (coll. Müller).

773

153. Alcis deversata (Staudinger, 1892) a-b. genitalia, aedeagus. Slovakia: Trenčin, Mníchova Lehota, 20.iv.1986 e.o., slide Müller 425 (coll. Müller). 154. Alcis jubata (Thunberg) a-b. genitalia, aedeagus. France: Haute-Savoie, Chamonix, 20.vii.1890, slide Müller 1013 (coll. MNHU). 155. Deileptenia ribeata (Clerck) a-c. genitalia, aedeagus, apical half of valva. Germany: Thuringia, Thuringian Forest, Ilmenau, 16,vii.1969, slide Müller 400 (coll. Müller). 156. Paraboarmia viertlii (Bohatsch) a-c. genitalia, aedeagus, central part of valva. Czech Republic: Moravia, Valtice, 15.vii.1981, slide Müller 445 (coll. Müller). 157. Adactylotis gesticularia (Hübner) a-b. genitalia, aedeagus. Spain: Catalonia, Girona, Santa Coloma, 8.ix.1993 e.o., slide Müller 443 (coll. Müller). 158. Adactylotis contaminaria (Hübner) a-b. genitalia, aedeagus. Spain: Catalonia, Olot, Pruit, 8.vii.1993, slide Müller 444 (coll. Müller). 159. Pseudocoremia suavis Butler a-b. genitalia, aedeagus. United Kingdom: Cornwall, Golophin Cross, 27.ix.2016, slide Sihvonen 2199 (coll. Clarke). c-e. vesica, cornuti at base of vesica, apex of juxta arm. United Kingdom: Cornwall, ­Golophin Cross, 27.ix.2016, slide Sihvonen 2198 (coll. Clarke). 160. Parectropis similaria (Hufnagel) a-c. genitalia, aedeagus, central part of valva. Germania: Berlin, Wilhelmshagen, 14.vi.1975, slide Müller 402 (coll. Müller). 161. Aethalura punctulata (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germania, Mecklenburg, Güstrow, Kluess, 14.v.1980, slide Müller 576 (coll. Müller). 162. Ascotis selenaria (Denis & Schiffermüller) a-b. genitalia, aedeagus. Italy: Toscana, Florence, 14.vii.1938 e.o., slide Müller 441 (coll. Müller). 163. Cleora cinctaria (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germania: Brandenburg, Guben, 14.ii.1987 e.o., slide Müller 421 (coll. Müller). 164. Ectropis crepuscularia (Denis & Schiffermüller) a-b. genitalia, aedeagus. Hungary: Budapest, János-Hegy, 9.iv.1986, slide Müller 396 (coll. Müller). c. vesica. Finland: Savonlinna, Laitaatsilta, 3.–9.vi.1997, slide Sihvonen 2784 (coll. Sihvonen).

774

165. Arichanna melanaria (Denis & Schiffermüller) a-b. genitalia, aedeagus. Germania: Brandenburg, Melchow, 8.vii.1982, slide Müller 416 (coll. Müller). c-d. vesica, cornuti. Finland: Riihimäki, 1–10.vii.1999, slide Sihvonen 2151 (coll. Sihvonen). 166. Bupalus piniaria (Linnaeus) a-b. genitalia, aedeagus. Germania: Brandenburg, Wünsdorf, 3.vi.2014, slide Müller 461 (coll. Müller). 167. Megametopon grisolaria (Eversmann) a-b. genitalia, aedeagus. Kazakhstan: Urda, 15.ix.2000, slide Müller 1053 (coll. P. Skou). 168. Tephronia sepiaria (Hufnagel) a-b. genitalia, aedeagus. Spain: Burgos, 14.viii.1900, slide ZSM G 6015 (ZSM). 169. Tephronia nuragica Fiumi, Flamigni, Zilli & Hausmann a-b. genitalia, aedeagus. France: Corsica, Pinarello, late vi.1973, slide ZSM G 13293 (ZSM). 170. Tephronia codetaria (Oberthür) subsp. sicula (Wehrli) a-b. genitalia, aedeagus. Italy: Sicily Madonie, 1200 m, 26.viii.[1927], slide ZSM G 6538 (ZSM). 171. Tephronia lhommaria (Cleu) subsp. melaleucaria (Schwingenschuss) a-b. genitalia, aedeagus. Spain: Central, Aranjuez, El Regajal, 23.ix.1989, slide ZSM G 9977 (ZSM). 172. Tephronia cyrnea (Schawerda) 172–1a-b. genitalia, aedeagus. Italy: Corsica, Francardo, 4.viii.1988, slide ZSM G 13255 Corsica (ZSM). 172–2a-b. genitalia, aedeagus. Italy: Corsica Corte, late vi.1973, slide ZSM G 13281 Corsica (ZSM). 172 3a-b. genitalia, aedeagus. Italy: Sardinia Stagno de Platamona, 6 km E Porto Torres, 5 m, 23.v.2004, slide ZSM G 20476 Sardinia (ZSM). 173. Tephronia lepraria Rebel a-b. genitalia, aedeagus. Hungary: Csongrad, Szeged-Királyhalom, 19.–29.vii.1933, slide ZSM G 13269 (ZSM). 174. Tephronia psyloritaria (Reisser) a-b. genitalia, aedeagus. Greece: Crete Idneon, 17.viii.1982, slide ZSM G 13751 (ZSM). 175. Tephronia theophilaria Hausmann a-b. genitalia, aedeagus. Italy: Alto Adige, Vinschgau, Schnalstal, 800 m, early vii.1969, slide ZSM G 20499 (ZSM, paratype). 176. Tephronia espaniola (Schawerda) a-b. genitalia, aedeagus. Spain: Teruel, Albarracin, Val de Vecar, 1100 m, 29.vii.2014, slide ZSM G 20550 (ZSM). 177. Eumannia oppositaria (Mann) a-b. genitalia, aedeagus. Greece: Rhodope Mts., Granitis, 800 m, 31.vii.1966, slide ZSM G 20480 (ZSM).

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178. Eumannia oranaria (Staudinger) 1a-b. genitalia, aedeagus. Morocco: Middle Atlas, Bin-el-Quidane (Ben Mellal), 4.vii.1972, slide ZSM G 6020 (ZSM). 2a-b. genitalia, aedeagus. Spain: Juneda, 12.vii.1933, slide ZSM G 13259 (ZSM) (subsp. castiliaria (Staudinger)). 179. Phyllometra culminaria (Eversmann) a-c. genitalia, aedeagus, valva. Hungary: Pilis-hegysék, Pilisvörösvár, 17.v.1956, slide ZSM G 20503 (ZSM). 180. Phyllometra gracilaria Boisduval a-c. genitalia, aedeagus, valva. Spain: Aragon, Albarracin, 21.vi.1963, slide ZSM G 13278 (ZSM). 181. Ithysia pravata (Hübner) a-d. genitalia, aedeagus, aedeagus detail, juxta processes. Ukraine: Crimea, Lenino, 7 km E Kamenskoje, 5.xi.2010, slide ZSM G 20505 (ZSM). 182. Leucobrephos middendorfii (Ménétriés) a-c. genitalia, aedeagus, vesica. Russia: Polar Ural, Shuryskary village W, forest tundra, 5.v.2001, slide Sihvonen 2212 (coll. Nupponen). 183. Myinodes constantina Hausmann a-c. genitalia, aedeagus, tip of valva. Tunisia: Thala, vi.1911, slide ZSM G 20536 (ZSM). 184. Kuchleria menadiara (Thierry-Mieg) a-c. genitalia, aedeagus, aedeagus detail. Italy: Sicily, 6.5 km E Pozallo, Santa Maria del Focallo, 5 m, 20.–21.ix.2014, slide ZSM G 20535 (ZSM). 185. Idaea barbuti Tautel & Lévêque a-c. genitalia, aedeagus, aedeagus detail. Portugal: Minho, Braga, Tebosa, 160 m, slide CT 517 (MNHN, holotype). 186. Idaea textaria (Lederer) See male genitalia figure in Vol. 2 fig. 49d: Turkey: Taurus, Marasch, 800 m, 1.v.1928, slide ZSM G 2158 (ZSM). 187. Idaea lobaria (Chrétien) a-b. genitalia, aedeagus. Algeria: Biskra, 23.v.1907, slide ZSM G 14377 (ZSM, lectotype). 188. Idaea gelbrechti Hausmann a-b. genitalia, aedeagus. Morocco: Middle Atlas, Ifrane, 1700 m, 12.–15.ix.1998 (e.o.), slide ZSM G 10577 (ZSM, paratype). 189. Oar pratana (Fabricius) See male genitalia figure in Vol. 2 Vol. 2 fig. 115b: Israel, Neot HaKikkar, –390 m, 14.–17.xii.1989 (ZSM, subsp. mortuaria Staudinger); fig. 115c: Spain, Gran Canaria, Maspalomas, iii.1967 (ZSM, paratype subsp. baezi Hausmann). 190. Scopula drenowskii Sterneck a-c. genitalia, aedeagus, sternum A8. Turkey: Amasia [without date], Hausm. 20601 (MNHU, type ‘aequata’ Staudinger).

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191. Scopula donovani (Distant) a-c. genitalia, aedeagus, sternum A8. Morocco: Taroudant, iv.1928, slide ZSM G 20513 (ZSM). 192. Nothocasis rosariae Scalercio, Infusino & Hausmann a-c. genitalia, aedeagus, detail of valva. Italy: Calabria, Vivaio Sbanditi, Longobucco, 13.x.2014, slide CREASAM 50 (CREA-SAM, paratype). 193. Perizoma juracolaria (Wehrli) a-c. genitalia, aedeagus. detail of vesica. Italy: Lazio, Apennino centr., Rieti, Leonessa, 3.5 km S Mte. Terminillo, 900 m, 20.–28.vii.2009, slide ZSM G 20541 (ZSM). 194. Perizoma barrassoi Zahm, Cieslak & Hausmann a-b. genitalia, aedeagus. Italy, Abruzzo, Majella, Valle Cannella, Manzini hut, 2350 m, 23.vii.1988, slide ZSM G 668Z (ZSM, holotype). 195. Eupithecia carpophilata (Püngeler) a-c. genitalia, aedeagus, sternum A8. Kyrgyzstan: south-east, Inilchek S env. Kalndy m.r., 3500 m, [without date], slide ZSM G 20543 (ZSM). 196. Eupithecia dubiosa Dietze a-c. genitalia, aedeagus, sternum A8. Israel: northern coast, border to Lebanon, Nahal Betset, 23.iii, 1987, slide ZSM G 1087 (ZSM). 197. Eupithecia furcata Staudinger a-c. genitalia, aedeagus, sternum A8. Turkey: Anatolia, Erzincan, 12.vi.1913, slide ZSM G 20546 (ZSM). 198. Eupithecia gypsophilata Skou, Mironov & Rietz a-c. genitalia, aedeagus, sternum A8. Spain: Zaragoza, Burajaloz, 30.vii.1962, Bujaraloz, 8.viii.1962, slide ZSM G 20552 (ZSM, paratype). 199. Eupithecia pindosata Weidlich a-d. genitalia, aedeagus, detail of vesica, sternum A8. Greece: Thessalia, 1 km NNW Krania, 1040 m, 19.iv.2018, slide Rajaei 2171/2018 (coll. Skou). 200. Eupithecia goossensiata Mabille a-c. genitalia, aedeagus, sternum A8. Germany: Kassel, Wilh’hausen, 23.vii.1950, slide ZSM G 3646 (ZSM). 201. Ourapteryx niveiscythes Zolotuhin a-b. genitalia, aedeagus. Russia: Lower Volga region, Astrakhan reserve, Obzhorovo, 2.vi.1970, slide V. Zolotuhin (ZISP, holotype). 202. Dyscia govii Fiumi & Flamigni a-b. genitalia, aedeagus. Sardinia, Olbia (Olbia-Tempio), Mte. Pinu, 450 m, 9.ix.2011, slide G. Fiumi (coll. P. Barberis, paratype).

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Appendix 2. Data of the microscopic preparations shown in female genitalia plates 1. Odontognophos dumetata (Treitschke) a-b. genitalia, signum. Croatia: Split-Dalmatia district, 2.5 km N Brela, 360 m, 4.x.2015, slide 2303 Sihvonen (coll. Skou). 2. Odontognophos margaritata (Zerny) a-b. genitalia, signum. Spain: Albarracin, ix.1926/1930 [label has two years], slide 2300 Sihvonen (coll. Skou). 3. Odontognophos zacharia (Staudinger) a-b. genitalia, signum. Turkey: Antalya, Akseki 5 km W by Ismit, 1000 m, 08.x.1999 ex ovo, slide 2313 Sihvonen (coll. Stadie). 4. Dicrognophos sartata (Treitschke) a. genitalia, Greece: Central Greece, 1 km NW Arachova, 1070 m, 9.vi.2013, slide Sihvonen 2074 (coll. Skou). b. posterior part of corpus bursae, Bulgaria: Rhodopi Mts, Assenovgrad, 14.ix.1985, slide ­Sihvonen 2073 (coll. Skou). 5. Dicrognophos siciliana Skou & Sihvonen a. genitalia, Italy: Sicily, 5.8 km ESE Santo Stefano Quisquina, 1000 m, 2.vi.2016, paratype, slide Sihvonen 2209 (coll. Skou). b. posterior part of corpus bursae, Italy: Sicily, 5.5 km ESE Collesano, Rifugio Orestano, 1100 m, 17.ix.2014, paratype, slide Sihvonen 2068 (coll. Skou). 6. Ortaliella perspersata (Treitschke) Genitalia, Spain: Tarragona, 3 km N of Alcossebre, 3−6.vi.2013, slide 2305 Sihvonen (coll. Sihvonen). 7. Apochima flabellaria (Hübner) Genitalia, Italy: Sicily, no date, slide Müller 1014 (coll. MNHU). 8. Chondrosoma fiduciaria Anker Genitalia, Austria: Neusiedler See, Zitzmannsdorfer Wiesen, x.1976 e.o., slide Müller 573 (coll. Müller). 9. Gnophos furvata (Denis & Schiffermüller) a-b. genitalia, Germany: Thüringen, Buchfarter Höhlen, 15.viii.1993, slide SE–744 (coll. Erlacher). 10. Gnophos obfuscata (Denis & Schiffermüller) Genitalia, Turkey: Erzurum, Rize Dağları, 2700 m, 5.viii.2001, slide SE–747 (coll. Stadie). 11. Charissa obscurata (Denis & Schiffermüller) Genitalia, Greece: Crete, Imbros, 1020 m, 2.x.2011, slide SE–738 (coll. Friedrich). 12. Charissa crenulata (Staudinger) Genitalia, Spain: Teruel, Albaracin, Moscardon, 1450 m, e.o., ♀: 13.viii.1998, slide DNATAX–2193/Erlacher (coll. Gelbrecht).

779

13. Charissa italohelveticus (Rezbanyai-Reser) Genitalia, Switzerland: Wallis, Raron, 700 m, e.o., 4.iv.1987, slide SE–755 (coll. B. Müller). 14. Charissa pullata (Denis & Schiffermüller) Genitalia, Germany: Thüringen, Asbach, Dietzenröder Stein, 496 m, 10.vii.2006, slide ­SE–751 (coll. Erlacher). 15. Charissa difficilis (Alphéraky) Genitalia, Russia: South Ural, Orenburg region, Donskoje village, 1.–4.vi.1998, slide ­SE–752 (coll. Skou). 16. Charissa turfosaria (Wehrli) Genitalia, Russia: Altai Mts., 26 km south of Seminski pass, 1000 m, e.o., ♀: 13.–19. vi.2012, slide JH–254 (coll. Stadie). 17. Charissa mutilata (Staudinger) Genitalia, Turkey: Ispir, Doğu Karadeniz Mts., Mount Korga, Köprüköy, 28.vii.2001, slide DNATAX–2320/Erlacher (coll. Gelbrecht). 18. Charissa peloponnesiaria Erlacher & Erlacher Genitalia, Greece: Peloponnesus, Chelmos Mountains, Aroania Ori, 1500 m, 20.ix. 1991, slide Viidalepp–54 (coll. ZMUC). 19. Charissa pentheri (Rebel) Genitalia, Macedonia: Ochrid-Petrina, 06.vi.1935, slide SE–725 (coll. ZSM). 20. Charissa assoi (Redondo & Gastón) Genitalia, Spain: Murcia, Aledo, 640 m, e.o., E ­ rlacher).

♀:

4.vi.2003, slide SE–106 (coll.

21. Charissa corsica (Oberthür) Genitalia, Italy: Ovette Malu, 7 km s Orgosolo, 960 m, 2.–16.vi.2002, slide DNATAX– 2371/Erlacher (coll. Gelbrecht). 22. Charissa staudingeri (Wnukowsky) a. genitalia, Greece: Rhodos, Lindos, 23.iv.1985, slide SE–56 (coll. ZSM). b. genitalia, Greece: Tinos, 17.–26.ix.2001, slide SE–54 (coll. ZSM). 23. Charissa subtaurica (Wehrli) Genitalia, Turkey: Samos, Kokkari, 05.v.2008, slide SE–328 (coll. Erlacher). 24. Charissa variegata (Duponchel) Genitalia, Greece: Kardaras, 13.ix.2004, slide SE–63 (coll. Friedrich). 25. Charissa mucidaria (Hübner) Genitalia, Morocco: Mount Atlas, Tizi-n-Test, 1700–1800 m, ♀: 11.–12.iv.1997, slide DNATAX– 2187/Erlacher (coll. Gelbrecht). 26. Charissa ambiguata (Duponchel) a. genitalia, Russia: Ural Mts., Cheljabinsk, Chebarkul village, 10.–19.vii.1993, slide CD–038 (coll. Skou). b. genitalia, France: Escouloubre-les-Bains, e.o., ♀: 6.viii.1996, slide CD–70 (coll. E ­ rlacher).

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27. Charissa predotae (Schawerda) Genitalia, Spain: Andalusia, Almeria/Granada, Sierra Nevada, Posada de los Arrieros, 1700 m, 20.v.2007, slide SE–267 (coll. Gelbrecht). 28. Charissa onustaria (Herrich-Schäffer) Genitalia, Bulgaria: Pirin Mts., Liljanovo, 550 m, e.o., 25.viii.1982, slide CD–054 (coll. Erlacher). 29. Charissa remmi (Viidalepp) Genitalia, Russia: Magadan, Upper Kolyma, 800 m, 11.vii.1980, slide SE–516 (coll. Skou). 30. Charissa zeitunaria (Staudinger) Genitalia, Turkey: Southern Taurus Mts., Zeitun, 1897, slide SE–521, paralectotype (coll. MNHB). 31. Charissa glaucinaria (Hübner) Genitalia, Austria: North Tyrol, Fließ, 1000 m, 15.vi.1994, slide SE–754 (coll. E ­ rlacher). 32. Charissa pfeifferi (Wehrli) Genitalia, Greece, Peloponnes, Mega Spileo bei Kalavrita, 800 m, 4.v.1998, slide DNATAX–2191/Erlacher. 33. Charissa supinaria (Mann) Genitalia, Germany: Thüringen, Kobersfels, ­Erlacher).

♀:

5.v.1993, slide SE–38/1999 (coll.

34. Charissa graecaria (Staudinger) Genitalia, Turkey: Akşhehir, Sultan-Dag, 1700–2200 m, 15.–30.viii.1934, slide SE– 559 (coll. ZFMK). 35. Psodos alpinata (Scopoli) Genitalia, Austria: Totes Gebirge, Warscheneck, 1500 m, 23.vii.1928, slide CJ−73 (coll. ZSM). 36. Psodos canaliculata (Hochenwarth) Genitalia, Austria: North Tyrol, Ötztal Alps, Breslauer Hütte, 2800 m, 28.vii.1953, slide CJ−95 (coll. ZSM). 37. Psodos werneri Schawerda Female is unknown to science. 38. Psodos coracina (Esper) Genitalia, Switzerland: Canton of Grisons, Lower Engadine, Flüela-Pass, 2300 m, 2.vii.1925, slide CJ−84 (coll. ZSM). 39. Psodos bentelii (Rätzer) Genitalia, Italy: Adamello, Rifugio Mandrone, 2500 m, 7.−10.vii.1964, slide CJ−64 (coll. TLMF). 40. Psodos noricana Wagner Genitalia, Austria: North Tyrol, Karwendel Mts., Innsbruck, Nordkette, 2300 m, 24.vii.1994, slide CJ−46 (coll. Erlacher).

781

41. Psodos belzebuth Praviel Genitalia, France: Alpes-Maritimes, Col de la Cayolle, 2400−2500 m, 1.viii.1973, slide CJ−100 (coll. ZSM). 42. Psodos pyrenaica Schawerda Genitalia, France: Pyrénées c., Pic du Midi, [without date], slide Rajaei 2177/2018 (coll. ZSM). 43. Psodos alpmaritima Wehrli Genitalia, France: St. Martin Vesubie, 14.vii.1913, slide ZSM G 14257 (coll. ZSM). 44. Psodos alticolaria Mann Genitalia, Austria: North Tyrol, Ötztal Alps, Rotmoostal, 2300 m, 15.vii.1973, slide CJ−104 (coll. ZSM). 45. Psodos wehrlii Vorbrodt Genitalia, Switzerland: Canton of Valais, Mettelhorn, 18.vii.1919, slide CJ−110 (coll. ZFMK). 46. Psodos baldensis Wolfsberger Genitalia, Italy: Verona, Monte Baldo, 2200 m, vii.1969, slide CJ−37 (coll. ZSM). 47. Psodos spitzi Rebel Genitalia, Slovenia: Triglav, 2500 m, vii.1934, slide CJ−78 (coll. TLMF). 48. Psodos quadrifaria (Sulzer) Genitalia, Switzerland: Canton of Grisons, Albula, Weissenstein, vii.1912, slide CJ−80 (coll. ZSM). 49. Sciadia tenebraria (Esper) a. genitalia, Switzerland: Valesia, Gomer Grat [unclear, handwritten], 18.viii.[19?]89, slide ­Sihvonen 2105 (coll. TLMF). b. signum, Italy: Veneto, Below Rif. Di Giussani, 2450 m, 31.vii.2013, slide Sihvonen 2138 (coll. Skou). 50. Sciadia septaria (Guenée) a-b. genitalia, ostium bursae and adjacent structures. France: Hautes-Pyrénées, Gèdre, without date, slide Sihvonen 2154 (coll. ZSM). 51. Sciadia innuptaria (Herrich-Schäffer) a-b. genitalia, signum. Austria: Dachstein, Simony Hütte, 2200 m, 15.vii.1928, slide Sihvonen 2153 (coll. ZSM). 52. Sciadia slovenica Leraut a-b. genitalia, signum. Slovenia: Soccatal, Bovec Mt. Canin, 05.viii.2002, 2280 m, slide Sihvonen 2109 (coll. TLMF). 53. Sciadia dolomitica Huemer & Hausmann a-b. genitalia, signum. Italy: Südtirol, Antermojasee, 15.vii.1924, slide Huemer G11547 (coll. ZSM). 54. Sciadia zelleraria (Freyer) a. genitalia, Italy: Völs af Schlern, Schlern, Petz, 2450–2560 m, 27.vii.2007, slide Sihvonen 2123 (coll. TLMF).

782

b.

signum, Italy: Monte Baldo, 2200 m, ex pupa 7.[19]66, slide Sihvonen 2160 (coll. TLMF).

55. Sciadia unicoloraria (Staudinger) a-b. genitalia, ostium bursae and adjacent structures. Italy: Piedmont, Bardonecchia, near Lago Sommeiller, 2950 m, 7.vii.1998, slide Flamigni (coll. Flamigni). 56. Elophos caelibaria (Heydenreich) a-b. genitalia, signum. Austria: Teriol. sept., Timmelsjoch, 2600 m, vii.1974 (ex pupa), slide Sihvonen 2129 (coll. TLMF). 57. Elophos zirbitzensis (Pieszcek) a-b. genitalia, ostium bursae and adjacent structures. Austria: Steiermark, Zirbitzkogel, ex larva 24.vi.1943, slide Sihvonen 2167 (coll. TLMF). 58. Elophos operaria (Hübner) a-b. genitalia, signum. Austria: Eisenerzer, Reichen, 14.vii.1938, slide Sihvonen 2117 (coll. TLMF). c-d. genitalia, signum. Austria: Steiermark, Zirbitzkogel, 1500–1800 m, 23.6.1991, slide Sihvonen 2119 (coll. TLMF). 59. Elophos andereggaria (de la Harpe) a. genitalia, Switzerland: Wallis [without date, but taken before 1948 according to other label data], slide Sihvonen 2166 (coll. ZSM). b. signum, France: Savoie, Cormet d’Arêches, 29.vii.1987, 2100 m, slide Sihvonen 2170 (coll. Sihvonen, donation from R. Mazel in 2016). 60. Yezognophos dilucidaria (Denis & Schiffermüller) a-b. genitalia, signum. Austria: Kärnten, St. Jakob im Lesachtal, Mussen, 3–4.viii.1999, 1800 m, slide Sihvonen 2113 (coll. TLMF). 61. Yezognophos sproengertsi (Püngeler) a-b. genitalia, ostium bursae and adjacent structures. France: Alpes-Maritimes, Park Natio­nal Mercantour, 1950 m, 7.vii.2012, slide Sihvonen 2173 (coll. TLMF). c-d. genitalia, signum. Italy: Alpi Cozie, Demonte NW, Colle Valcavera NE, 4.viii.2010, 2420 m, slide Sihvonen 2133 (coll. TLMF). 62. Yezognophos vittaria (Thunberg) a-b. genitalia, ostium bursae and adjacent structures. Norway: Vestlige Finnmark, 10 km S Burfjord, 200–300 m, 20.vii.1998, slide Sihvonen 2081 (coll. Sihvonen). 63. Yezognophos serotinaria serotinaria (Denis & Schiffermüller) 1a-b. genitalia, signum. Austria: Tirolis borealis, Ötztal Ötz. Umgeb. [without date], slide Sihvonen 2190 (coll. TLMF). Yezognophos serotinaria dognini (Thierry-Mieg) 2a-b. genitalia, signum. France: Dept. Hautes-Pyrénées, Lac d’Orédon, 1900 m, 24.vii. 2014, slide Sihvonen 2186 (coll. Skou). Yezognophos serotinaria serotinoides (Wehrli) 3a-b. genitalia, signum. Switzerland: Wallis, Leuk-Guttet, 1000 m, ex pupa 14.11.[19]91, slide Sihvonen 2131 (coll. TLMF). 64. Crocota tinctaria (Hübner)

783

b.

Genitalia, Italy: Alpi Cozie, Demonte NW, Colle Valcavera NE, 2420 m, 2.viii.2010, slide Sihvonen 2102 (coll. TLMF).

65. Crocota peletieraria (Duponchel) Genitalia, Spain: c. 20 km W of Andorra, Vall Ferrera, 25.vii.1957, slide Sihvonen 2140 (coll. Skou). 66. Crocota pseudotinctaria Leraut Genitalia, Switzerland: Leukerbad, Torrent 2300 m, 16.ix.1993 (ex ovo from a female 24.07.1993), slide Sihvonen 2098 (coll. Sihvonen). 67. Crocota niveata (Scopoli) Genitalia, Austria: Lavanttaler Alpen, Saualpe 1770 m, 14.vii.2011, slide Sihvonen 2100 (coll. Sihvonen). 68. Synopsia sociaria (Hübner) Genitalia, Bulgaria: Varna/Kranevo, 5.viii.1984 e.o., slide Müller 556 (coll. Müller). 69. Phthonandria strictaria (Lederer) a-b. genitalia, signum. Russia: Ural Mountains, Orenburg Region, Kyvandyk Village, 1−11.vi.1995, slide 2230 Sihvonen (coll. Skou). 70. Menophra abruptaria (Thunberg) Genitalia, Spain: Catalonia, Girona, Sa. De Montseny, 25.ix.1991 e.o., slide Müller 378 (coll. Müller). 71. Menophra japygiaria (O. Costa) Genitalia, Tunesia: Tabarka, 23.vi.1993, slide Müller 553 (coll. Müller). 72. Menophra berenicidaria (Turati) Genitalia, Turkey: Antalya, 29.v.1996 e.o., slide Müller 379 (coll. Müller). 73. Mephra harterti (Rothschild) Genitalia, Algeria: Guelt es Stel, 15.–23.ix.1933, slide Müller 1042 (coll. ZSM). 74. Mephra thuriferaria (Zerny) Genitalia, Spain: Teruel, Albarracin, 8.ix.2006, slide Müller 365 (coll. Müller). 75. Mephra nycthemeraria (Geyer) Genitalia, France: Provence, Choudon-Noranthe, 800 m, 30.iii.1996 e.o., slide Müller 381 (coll. Müller). 76. Mephra annegreteae Skou Genitalia, Spain: Almeria, Tabernas, 29.v.2006, paratype, slide Müller 380 (coll. Müller). 77. Cleorodes lichenaria (Hufnagel) Genitalia, France: Midi-Pyrénées, Lacerede, 1.ix.2002 e.o., slide Müller 566–1 (coll. Müller). 78. Zernyia granataria (Staudinger) Genitalia, Spain: Huesca, Candasnos, 28.viii.2001, slide Müller 1044 (coll. Skou).

784

79. Megalycinia serraria (Costa) a-b, d. genitalia, ostium bursae and adjacent structures, signum with irregular centre (and detached cornutus). Italy: Basilicata, Monte Caperino, 1400 m, 25.ix.1973, slide S ­ ihvonen 2259 (coll. Skou). c. signum with solid centre, Italy: Calabria, 3 km E Orsomarso, 10.viii.2007, slide Müller 1047 (coll. ZSM). 80. Dasypteroma thaumasia Staudinger Genitalia, Spain: Madrid, Ciempopzuelos, Cerros de Palomera, 17.ix.2000, slide Müller 569 (coll. Müller). 81. Microbiston lanaria (Eversmann) Genitalia, Kazakhstan: Uralsk (Oral), date unknown, slide Müller 1069 (coll. Skou). 82. Apocheima hispidaria (Denis & Schiffermüller) Genitalia, Germany: Brandenburg, Nauen, Brieselang, 23.iii.1981, slide Müller 504 (coll. Müller). 83. Phigalia pilosaria (Denis & Schiffermüller) Genitalia, Germany: Brandenburg, Berkenbrück, 16.ii.1981 e.o., slide Müller 483 (coll. Müller). 84. Hypomecis roboraria (Denis & Schiffermüller) a-b. genitalia, signum. Germany: Brandenburg, Guben, Grano, 8.v.1989 e.o., slide Müller 432 (coll. Müller). 85. Hypomecis punctinalis (Scopoli) a-b. genitalia, signum. Germany: Brandenburg, Mühlenbeck, 6.iv.1987 e.o., slide Müller 429 (coll. Müller). 86. Ematurga atomaria (Linnaeus) Genitalia, Germany: Mecklenburg, Müritz National Park, 26.iv.2014, slide Müller 415 (coll. Müller). 87. Lycia hirtaria (Clerck) a-b. genitalia, signum.Germany: Brandenburg, Ahrensfelde, 19.iv.1970, slide Müller 496 (coll. Müller). 88. Lycia necessaria (Zeller) a-b. genitalia, signum. Caucasus mer. occ., date unknown, slide Müller 1024 (coll. MNHU). 89. Lycia lapponaria (Boisduval) a-b. genitalia, signum. “Livonia” (Latvia and Estonia), date unknown, slide Müller 1004 (coll. MNHU). 90. Lycia isabellae (Harrison) a-b. genitalia, signum. Germany: Saxonia, Zittau montains, 25.ii.1986 e.o., slide Müller 535 (coll. Müller). 91. Lycia pomonaria (Hübner) a-b. genitalia, signum. Germany: Surroundings of Berlin, 9.iii.1937 e.l., slide Müller 506 (coll. Müller).

785

92. Lycia zonaria (Denis & Schiffermüller) Genitalia, Germany: Brandenburg, Kyritz, Dreetz, 15.iii.1992 e.o., slide Müller 493 (coll. Müller). 93. Lycia alpina (Sulzer) Genitalia, Alpes, date unknown, slide Müller 1006 (coll. MNHU). 94. Lycia graecarius (Staudinger) Genitalia, Croatia: Mostar, 14.iii.1944, slide Müller 1005 (coll. MNHU). 95. Lycia florentina (Stefanelli) Genitalia, Italy: Emilia-Romagna, date unknown, slide Müller 1008 (coll. MNHU). 96. Biston strataria (Hufnagel) Genitalia, Germany: Brandenburg, Königs Wusterhausen, Kablow, 4.iv.1980, slide Müller 491 (coll. Müller). 97. Biston rosenbaueri Müller Female is unknown. 98. Biston achyra Wehrli Female is unknown. 99. Biston betularia (Linnaeus) Genitalia, Germany: Brandenburg, Zerpenschleuse, 5.vii.1980, slide Müller 1056 (coll. Busse). 100. Erannis defoliaria (Clerck) Genitalia, Germany: Thuringia, Ilmenau, 500 m, 4.xii.1979 e.o., slide Müller 480 (coll. Müller). 101. Erannis ankeraria (Staudinger) Genitalia, Slovenia: Podgorje, 24.iii.2012 e.o., slide Müller 1066 (coll. B. May). 102. Erannis declinans (Staudinger) Genitalia, Turkey: Erzurum, Mescit Dağları, 27.x.2010 e.l., slide Müller 1070 (coll. ­Gelbrecht). 103. Kresnaia beschkovi Ganev a-b. genitalia, signum. Bulgaria: Pirin mts., Kresna gorge, 26.xi.2009, slide Müller 1052 (coll. Beshkov). 104. Agriopis leucophaearia (Denis & Schiffermüller) Genitalia, Czech Republic: Bohemia, Prague, 10.iii.1972, slide Müller 476 (coll. Müller). 105. Agriopis bajaria (Denis & Schiffermüller) Genitalia, Italy: Sardinia, Olbia, San Pantaleo, 30.xii.2007, e.l., slide Müller 477 (coll. Müller). 106. Agriopis aurantiaria (Hübner) a-b. genitalia, signum. Germany: Brandenburg, Berkenbrück, 20.xi.1980, e.l., slide Müller 473 (coll. Müller).

786

107. Agriopis budashkini Kostjuk a-b. genitalia, signum. Greece: Crete, Asfendos, 8.xi.2015 e.l., slide Müller 564 (coll. Müller). 108. Agriopis marginaria (Fabricius) a-b. genitalia, signum. Germany: Brandenburg, Glienicke Nordbahn, 27.iii.1980, slide Müller 471 (coll. Müller). 109. Calamodes occitanaria (Duponchel) Genitalia, Spain: Teruel, Albarracin, 1200 m, 11.ix.2008, slide Müller 426 (coll. Müller). 110. Calamodes melanaria Oberthür Genitalia, Italy: Calabria, Fiumara Trionto (CS), 21.x.1999, slide Müller 1050 (coll. ZSM). 111. Paradarisa consonaria (Hübner) a-b. genitalia, signum. Germany: Thuringia, Thuringian Forest, Martinroda, 30.iv.1968, slide Müller 399 (coll. Müller). 112. Larerannis orthogrammaria (Wehrli) a. genitalia. South Korea: [5 km SE of Pocheon] Mt Jugeum-san, 15.xi.1997, slide Sihvonen 2778 (coll. Kim). b. signum. Japan: Tochigi Nikko, 15.xi.1986, slide Nakajima 3214 (photo and coll. ­Nakajima). 113. Chemerina caliginearia (Rambur) a-b. genitalia, signum. Spain: Catalonia, Girona, Vidreras, 12.iii.1991, slide Müller 486 (coll. Müller). 114. Fagivorina arenaria (Hufnagel) a-b. genitalia, signum. Croatia: Istria, Ćićarija mts., Buzet, 800–1200 m, 16.vii.1986 e.o., slide Müller 438 (coll. Müller). 115. Spartopteryx kindermannaria Staudinger Genitalia, “Tibet, Kuku-Noor”, [China: Qinghai, Qinghai Hu], vi.1898, slide Müller 1009 (coll. MNHU). 116. Athroolopha pennigeraria (Hübner) Genitalia, Spain: Almería, Gérgal, 16.ix.2004 e.o., slide Müller 517 (coll. Müller). 117. Athroolopha chrysitaria (Hübner) Genitalia, pink colour mark, no further data, slide Müller 1017 (coll. Staudinger in MNHU). 118. Eurranthis plummistaria (Villers) Genitalia, France: Alpes-de-Haute-Provence, Digne, 19.iv.1992, slide Müller 508 (coll. Müller). 119. Nychiodes obscuraria (Villers) Genitalia. Italy: South Tyrol, Kompatsch, 23.vii.1979, slide Müller 390 (coll. Müller). 120. Nychiodes ragusaria Millière Genitalia, Italy: Kalabria, Tropea Coccorina, 200 m, 15.vi.2008, slide Rajaei 2154/2016 (coll. Fiebig). 787

121. Nychiodes andalusiaria Millière Genitalia, Spain: Sierra Estrella, 20.x.1931, slide Rajaei 2035/2016 (coll. Zsm). 122. Nychiodes notarioi Expósito Genitalia, Spain: Gerona, 1400 m, Sierra del Montseny, 08.vii.1993, slide Rajaei 2006/2016 (coll. Müller). 123. Nychiodes hispanica Wehrli a. genitalia, Spain: Sierra Nevada, 06.vii.1926, slide Fazekas 2595 Lectotype (coll. ZFMK). b. variation of lamella postvaginalis. Spain: Malaga, 25 km SW Ronda, Algatocin, 865 m, 28.vii.98, slide Rajaei 2144/2017 (coll. Stadie). 124. Nychiodes waltheri Wagner a. genitalia, Bulgaria: Sozopol, 16.xi.1987 ex. ovo., slide Müller 386 (coll. Müller). b. genitalia, Spain: Serrania de Cuenca, Huelamo, 1400 m, slide Leraut 10821 (Holotype of cuencaensis Leraut syn. nov.) (coll. MNHN) 125. Nychiodes amygdalaria (Herrich-Schäffer) Genitalia, Turkey: Artvin, SE-Side, Barhal-valley, 4 km NE Altıparmak, 100 m, 31.7.–3.8.1983, slide Rajaei 2088/2017 (coll. SMNS). 126. Nychiodes dalmatina Wagner Genitalia, Bulgaria: Pirin Mts., Liljanovo, 550 m, 21.viii.1981 ex. ovo., slide Rajaei 2001/2017 (coll. Müller). 127. Phaselia serrularia (Eversmann) a-b. genitalia, signum. Turkey: Amasya Province, Amasya, 13.–27.vi.1978, slide Müller 1061 (coll. SMNS). 128. Phaselia algiricaria aragona Wehrli a-b. genitalia, signum. Spain: Teruel, Albarracín, 2.vi.2007, slide Müller 571 (coll. Müller). 129. Ekboarmia atlanticaria (Staudinger) a-b. genitalia, signum. Spain: 10 km NW of Tarifa, Punta Paloma, 30 m, 22–23.v.2006, slide Sihvonen 2049 (coll. Skou). 130. Ekboarmia sagnesi Dufay a-b. genitalia, signum. France: Valle de la Romanche, 1200 m, 22–23.vi.1999, slide Sihvonen 2096 (coll. Skou). 131. Ekboarmia miniaria Skou, Stüning & Sihvonen a-b. genitalia, ostium bursae and adjacent structures. Portugal: Grandola, 15.vi.2009, slide Sihvonen 2051, paratype (coll. Skou). 132. Ecleora solieraria (Rambur) a-b. genitalia, ostium bursae and adjacent structures. Spain: Teruel, Albarracín, 15.vi.2007, slide Müller 554 (coll. Müller). 133. Afriberina tenietaria (Staudinger) a-b. genitalia, ostium bursae and adjacent structures. Spain: Prov. Huelva, Matalascanas, 35 m, 18.iv.2017, slide Sihvonen 2242 (coll. Skou).

788

134. Afriberina salemae Skou & Sihvonen a. genitalia, Portugal: Algarve, Salema Camping, 55 m, 22.iv.2017, slide Sihvonen 2256 (coll. Skou), paratype. b. ostium bursae and adjacent structures, Portugal: Algarve, Salema Camping, 55 m, 22.iv. 2017, slide Sihvonen 2241 (coll. Skou), paratype. 135. Sardocyrnia bastelicaria (Bellier) Genitalia, France: Corsica, 1906, slide Müller 1001 (coll. MNHU). 136. Sardocyrnia fortunaria (Vázquez) Genitalia, Spain: Teruel, Albarracín, 8.vi.2007, slide Müller 447 (coll. Müller). 137. Selidosema brunnearia (Villers) a-b. genitalia, signum. Denmark: Bornholm, 17.vii.1990, slide Müller 412 (coll. Müller). 138. Selidosema plumaria (Denis & Schiffermüller) a-b. genitalia, signum. Hungary: Budapest, Budaörs, 18.v.1984 e.o., slide Müller 409 (coll. Müller). 139. Selidosema pyrenaearia (Boisduval) a-b. genitalia, signum. Spain: Amauida, Avila, 1431 m, 14.ix.2004, slide Müller 1055 (coll. J. Ylla). 140. Selidosema taeniolaria (Hübner) a-b. genitalia, signum. “Southern France”, 29.vi.2006 e.o., slide Müller 413 (coll. Müller). 141. Selidosema ambustaria (Geyer) a-b. genitalia, signum. Italy: Sicilia, Girgenti (now Agrigent), 3.viii.1910 e.o., slide Müller 1019 (coll. MNHU). 142. Selidosema erebaria postdentaria (Rothschild) Genitalia, Italy: Island of Pantelleria, 8.–13.x.2007, slide Müller 579 (coll. Müller). 143. Peribatodes rhomboidaria (Denis & Schiffermüller) a-b. genitalia, ductus bursae. Germany: Stuttgart, Kräherwald, 20.viii.1955, slide Rajaei 2168/2018 (coll. SMNS). 144. Peribatodes perversaria (Boisduval) a-b. genitalia, ductus bursae. Switzerland: Wallis, slide Rajaei 2167/2018 (coll. SMNS). 145. Peribatodes omalensis Rajaei, Skou & Hausmann a-b. genitalia, ductus bursae. Greece: Western Crete, Omalos Plateau S of Chania, 53°20’39’’N, 23°54’18’’E, 27.vi.2012, 1050 m, slide Rajaei 2147/2017 (coll. SMNS). 146. Peribatodes umbraria (Hübner) a-b. genitalia, ductus bursae. Italy: Sardinia, N 41.02’, E 09.29’, Costa Smeralda San Pantaleo 200 m, 02.–09.vi.2005, slide Rajaei 1982/2016 (coll. Müller). 147. Peribatodes ebusaria Vallhonrat a. genitalia, Spain: Ibiza, 4.2 km SW Sant Rafel, Puig des Fornas Coll des Rossellons, 220 m, 30.iii.–3.iv.2016, slide Rajaei 2175/2018 (coll. Skou). b. ductus bursae, photo modified from Vallhonrat, 2012, Fig. 5, Paratype.

789

148. Peribatodes secundaria (Denis & Schiffermüller) a-b. genitalia, ductus bursae. Austria: Kärnten 46°50’23” N, 014°43’09” E, Lavanttaler Alpen Saualpe, 1335 m, 09.vii.2011, slide Rajaei 1993/2016 (coll. Müller). 149. Peribatodes correptaria (Zeller) a-b. genitalia, ductus bursae. Greece: Creta, N 35.08’, E 25.44’, Istro, south of Agios Nikolaos 15.v.1997, slide Rajaei 1988/2016 (coll. Müller). 150. Peribatodes aragonis (Wehrli) 1a-b. genitalia, ductus bursae. Spain: Teruel, N 40°24’45”, W 001°25’43”, Serrania de Albarracin, 1140 m, 05.ix.2008, slide Rajaei 1981/2016 (coll. Müller). Peribatodes powelli (Oberthür). Not found in Europe, illustrated for comparison. 2a-b. genitalia, ductus bursae. Morocco: Ifrane, v.1950, slide Leraut 11457 (coll. MNHN). 151. Peribatodes ilicaria (Geyer) a-b. genitalia, ductus bursae. Spain: Murcia, 640 m, Aledo westlich Totana, 04.vii.2003, slide Rajaei 1990/2016 (coll. Müller). 152. Alcis repandata (Linnaeus) Genitalia, Germany: Thuringia, Ilmenau, 500 m, 10.vi.1973 e.o., slide Müller 422 (coll. Müller). 153. Alcis deversata (Staudinger) Genitalia, Slovakia: Trenčin, Mníchova Lehota, 22.v.1986 e.o., slide Müller 424 (coll. Müller). 154. Alcis jubata (Thunberg) Genitalia, Germany: Brandenburg, Märkisch Buchholz, vii.1902, e.l., slide Müller 1012 (coll. MNHU). 155. Deileptenia ribeata (Clerck) a-b. genitalia, signum. Germany: Mecklenburg, Serrahn, 28.ix.1989 e.o., slide Müller 401 (coll. Müller). 156. Paraboarmia viertlii (Bohatsch) a-b. genitalia, signum. Hungary: “Fünfkirchen”, (Southern Transdanubia, Pécs), no date, slide Müller 1003 (coll. MNHU). 157. Adactylotis gesticularia (Hübner) a-b. genitalia, signum. Spain: Teruel, Albarracín, 11.vi.2007, slide Müller 442 (coll. Müller). 158. Adactylotis contaminaria (Hübner) a-b. genitalia, signum. Spain: no other data, slide Müller 1011 (coll. MNHU). 159. Pseudocoremia suavis Butler a. genitalia, United Kingdom: Cornwall, Godolphin Cross, 27.ix.2016, slide Sihvonen 2197 (coll. Clarke). b. signum, United Kingdom: Cornwall, Godolphin Cross, 27.ix.2016, slide Sihvonen 2196 (coll. Clarke). 160. Parectropis similaria (Hufnagel) a-b. genitalia, signum. Germany: Thuringia, Thuringian Forest, Arnstadt, 5.vi.1971, slide Müller 403 (coll. Müller). 790

161. Aethalura punctulata (Denis & Schiffermüller) a-b. genitalia, signum. Germany: Brandenburg, Nauen, Finkenkrug, 10.v.1970, slide Müller 463 (coll. Müller). 162. Ascotis selenaria (Denis & Schiffermüller) a-b. genitalia, signum. Cyprus: Agia Napa, 5.viii.2001 e.o., slide Müller 440 (coll. Müller). 163. Cleora cinctaria (Denis & Schiffermüller) a-b. genitalia, signum. Germany: Thuringia, Ilmenau, 500 m, 17.v.1969, slide Müller 420 (coll. Müller). 164. Ectropis crepuscularia (Denis & Schiffermüller) Genitalia, Hungary: Budapest, Kesztölz, 21.ii.1986 e.o., slide Müller 431 (coll. Müller). 165. Arichanna melanaria (Denis & Schiffermüller) Genitalia, Germany: Baden-Württemberg, Ringschnait, vi.1927 e.o., slide Müller 417 (coll. Müller). 166. Bupalus piniaria (Linnaeus) Genitalia, Germany: Brandenburg, Alt Schadow, 8.vi.2014, slide Müller 465 (coll. Müller). 167. Megametopon grisolaria (Eversmann) Female genitalia not available. 168. Tephronia sepiaria (Hufnagel) Genitalia. Austria: Wien, Mödling, 20.vi.1889, slide ZSM G 20474 (ZSM). 169. Tephronia nuragica Fiumi, Flamigni, Zilli & Hausmann Genitalia. France: Corsica, Pinarello, late vi.1973, slide ZSM G 13294 (ZSM). 170. Tephronia codetaria (Oberthür) subsp. sicula (Wehrli) Genitalia. Italy: Sicily, Madonie, 1200 m, 24.viii.[1927], slide ZSM G 20475 (ZSM). 171. Tephronia lhommaria (Cleu) subsp. cleui Wehrli a. genitalia. France: Hautes Alpes, St.-Crépin, 28.vi.1933, slide ZSM G 13254 (ZSM) b. genitalia. Italy: Abruzzo, San Donato, mid-vii.1964, slide ZSM G 13254 (ZSM) (identification requiring confirmation). 172. Tephronia cyrnea (Schawerda) a. genitalia. France: Corsica, Bonifacio, Pont de Ventilegne, 29.v.2014, slide ZSM G 20478 (ZSM, paratype ‘tonnara’ Tautel). b. genitalia. Italy: Sardinia, Fonni, 14.viii.2003, slide ZSM G 20835 (ZSM). 173. Tephronia lepraria Rebel Genitalia. Hungary: KNP Bugac nyires nada s, 4.viii.1980, slide ZSM G 20496 (ZSM). 174. Tephronia psyloritaria (Reisser) Genitalia. Greece: Crete, Chora Sfakion, Askifou, Kares, 740 m, 7.viii.2012, slide ZSM G 20485 (ZSM).

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175. Tephronia theophilaria Hausmann Genitalia. Italy: Basilicata, upper Noce valley, near Lagonegro, 700 m, 2.ix.1998, slide ZSM G 13296 (ZSM, paratype). 176. Tephronia espaniola (Schawerda) Genitalia. Spain: Teruel, Moscardón, 1400 m, 28.vii.2014, slide ZSM G 20551 (ZSM). 177. Eumannia oppositaria (Mann) a-b. genitalia, signum. Turkey: 55 km W Konya, 1300 m, 9.viii.1990, slide ZSM G 20492 (ZSM). 178. Eumannia oranaria (Staudinger) 1a-b. genitalia, signum. Morocco: Middle Atlas, Ifrane, 1700 m, 25.vii.1975, slide ZSM G 20500 (ZSM). 2a-b. genitalia, signum. France: Var, St. Tropez, 2.vii.1948, slide ZSM G 13285 (ZSM) (subsp. cebennaria (Chrétien)). 179. Phyllometra culminaria (Eversmann) a-b. genitalia, signum. Hungary: Pilisvörösvar, 4.v.1934, slide ZSM G 20504 (ZSM). 180. Phyllometra gracilaria Boisduval a-b. genitalia, signum. Spain: Soria, Sierra de Moncayo, Beraton, 1200 m, 20.vi.1968, slide ZSM G 13291 (ZSM). 181. Ithysia pravata (Hübner) Genitalia. Russia: Sarepta [without date], slide ZSM G 20506 (ZSM). 182. Leucobrephos middendorfii (Ménétriés) Genitalia. Russia: Polar Urals, Jablonevoj Mts., Jukutsk [without date], slide 2174/2018 H. Rajaei (ZMKU). 183. Myinodes constantina (Hausmann) Genitalia. Tunisia: Djebel Penalba, late iii.1933, slide ZSM G 20537 (ZSM). 184. Kuchleria menadiara (Thierry-Mieg) Genitalia. Italy: Sicily, 6.5 km E Pozallo, Santa Maria del Focallo, 5 m, 20.–21.ix.2014, slide ZSM G 20539 (ZSM). 185. Idaea barbuti Tautel & Lévêque Genitalia. Portugal: Minho, Braga, Tebosa, 160 m, slide AL 31 (MNHN, paratype). 186. Idaea textaria (Lederer) See female genitalia figure in Vol. 2 fig. 49b: Turkey: Taurus, Marasch, 800 m, 8.xi.1924 (e.o.), slide ZSM G 2159 (ZSM). 187. Idaea lobaria (Chrétien) Genitalia. Algeria: Biskra, 9.vi.1907, slide ZSM G 20510 (ZSM, paratype). 188. Idaea gelbrechti Hausmann Genitalia. Morocco: Middle Atlas, Ifrane, 1700 m, 12.ix.1998 (e.o.), slide ZSM G 11284 (ZSM, paratype).

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189. Oar pratana (Fabricius) See female genitalia figure in Vol. 2 fig. 115b: Tunisia: Tunis, 7.v.1912, slide ZSM G 1627 (ZSM). 190. Scopula drenowskii Sterneck Genitalia. Turkey: Kızılcahamam, 6.vi.1995, slide ZSM G 20512 (ZSM). 191. Scopula donovani (Distant) Genitalia. Morocco: Ait Melloul, 13.iv.1969, slide ZSM G 20538 (ZSM). 192. Nothocasis rosariae Scalercio, Infusino & Hausmann a-b. genitalia, antrum. Italy: Calabria, Timpone Magara, Saracena, 18.xi.2015, slide CREASAM 60 (CREA-SAM) 193. Perizoma juracolaria (Wehrli) a-b. genitalia, signum. Italy: Lazio, Apennino centr., Rieti, Leonessa, 3.5 km S Mte. Terminillo, 900 m, 20.–28.vii.2009, slide ZSM G 20542 (ZSM). 194. Perizoma barrassoi Zahm, Cieslak & Hausmann a-b. genitalia, signum. Italy: Lazio, Mte. Terminillo, 1700–1900 m, 13.vii.2004, slide ZSM G 15840 (ZSM). 195. Eupithecia carpophilata (Püngeler) Genitalia. Russia: Altai, south-west, Katun valley, 10 km W Katanda, 1200 m, 15.– 19.vii.1983, slide ZSM G 20544 (ZSM). 196. Eupithecia dubiosa Dietze Genitalia. Israel: northern coast, border to Lebanon, Nahal Ksiv, 24.iii, 1987, slide ZSM G 1093 (ZSM). 197. Eupithecia furcata Staudinger Genitalia. Turkey: north-west, Baysehir gölü, 4.v.1964, slide ZSM G 20547 (ZSM). 198. Eupithecia gypsophilata Skou, Mironov & Rietz Genitalia. Spain: Zaragoza, Bujaraloz, 8.viii.1962, slide ZSM G 14315 (ZSM, paratype). 199. Eupithecia pindosata Weidlich Genitalia. Greece: Thessalia, Notia Pindos, Tringia, 6 km N Kranea/Ahelóos, 1050 m, 5.v. 2003, slide ZSM G 20549 (ZSM, paratype). 200. Eupithecia goossensiata Mabille Genitalia. Germany: Lower Saxonia, Vennermoor near Osnabrück, 30.vii.1985, slide ZSM G 20545 (ZSM). 201. Ourapteryx niveiscythes Zolotuhin Genitalia. Russia: Lower Volga region, Astrakhan reserve, cordon nr. 3, 1.–10. viii.2004, slide V. Zolotuhin (ZISP, paratype). 202. Dyscia govii Fiumi & Flamigni a. genitalia. Italy: Sardinia, Olbia (Olbia-Tempio), Mte. Pinu, 450 m, 17.v.2008, slide G. Fiumi (coll. P. Barberis, allotype). b. genitalia. Italy: Sardinia, Cagliari, Sinnai, 200 m, 7.x.2007, slide G. Fiumi (coll. M. Guidi, paratype).

793

Revised, annotated systematic checklist of the ­Geometridae of Europe and adjacent areas, Vols 1–6 by Axel Hausmann and Pasi Sihvonen (eds) 1 The checklist shows, in systematic order, the taxa (from subfamily level to subspecies level) of the European Geometridae treated in Vols 1–6, updated and including most newly described species and taxonomic changes since the publication date (GME1: 2001; GME2: 2004; GME3: 2012; GME4: 2003; GME5: 2015). All changes are annotated with footnotes. Just a few synonyms and incorrect subsequent spellings (with ‘=’ ahead) are added, when in use until recently. 810 taxa occurring in areas immediately adjacent to Europe (as defined here) are inserted (marked with an asterisk) in the list with an indication of their distribution in these adjacent countries. It will be interesting to see whether, and when, some of these species will be found in Europe. Altogether, the list of European Geometridae now comprises 999 species and 289 validated subspecies, with additional 492 species and 320 subspecies mentioned in this checklist (most from adjacent countries). Current European taxon numbers can be broken down to subfamily level as follows: Archiearinae: 5 species, 1 subspecies (cf. Hausmann 2001) ‘Orthostixinae’ [putative tribe of Ennominae?]: 1 species, 0 subspecies (cf. Hausmann 2001) (in addition two extralimital species; one subspecies). Desmobathrinae: 4 species, 1 subspecies (cf. Hausmann 2001) (in addition three extralimital species). Geometrinae: 32 species, 7 subspecies (cf. Hausmann 2001) (+27sp/13subsp.) (in addition 27 extralimital species; 13 subspecies). Ennominae: 327 species, 78 subspecies (cf. Skou & Sihvonen 2015 and present volume; three species are added from Vol. 1: genera Epirranthis/Alsophila) (in addition 187 extralimital species; 191 subspecies). Sterrhinae: 206 species, 55 subspecies; four species are added from Vol. 3: genus Lythria) (cf. Hausmann 2004) (in addition 121 extralimital species; 58 subspecies). Larentiinae: 424 species, 147 subspecies (cf. Hausmann & Viidalepp 2012; Mironov 2003) (in addition 152 extralimital species; 57 subspecies). Our understanding of the higher classification of the Geometridae phylogeny has improved significantly during recent years. This is mainly due to the progress in molecular phylo­ genetic studies and the checklist has benefited greatly from such input (for instance ­Sihvonen et al. 2011; Ounap et al. 2016; Jiang et al. 2017; Ban et al. 2018; Murillo-Ramos et al. 2019). Despite these advances, relationships at the genus level, and particularly at the tribus level, are still partly immature due to limited taxon and gene sampling. The majority of Euro­pean Geometridae genera have not been included in such multi-gene phylogenetic analysis yet. Therefore the results of the modern molecular phylogenies are incorporated in the checklist critically, acknowledging that our understanding on the relationships of taxa is likely to change. Furthermore, we would like to remind our readers that because there is no 1

The checklists for the genera treated in GME6 have been prepared by the authors as indicated and specified in the introduction to this volume. The checklists for Perizomini and Eupitheciini are based on that published in GME4 (Mironov 2003), with numerous modifications.

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­ nambiguous way to transfer information from the phylogenetic hypotheses into a linear u checklist, the readers are encouraged to study the original hypotheses in parallel. The checklist does not include species imported to Europe which are occasionally found in greenhouses, plant nurseries or are otherwise considered not being part of the European fauna. In the following list the European species are followed by a combination of numbers, referring to the GME volume and the number of the species in this volume, e.g. 1/1 meaning GME Vol. 1 (Hausmann 2001), species number 1.

Subfamily Archiearinae Fletcher, 1953 Archiearis Hübner, 1823 A. parthenias (Linnaeus, 1761). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/1 Boudinotiana Leraut, 2002 2 B. notha (Hübner, 1803). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/2 B. puella (Esper, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/3 B. p. mediterranea Ganev, 1984 B. touranginii Berce, 1870. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/3a Leucobrephos Grote, 1874 3 L. middendorfii (Ménétriés, 1858) 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/182 (= mongolicum Vojnits, 1977) 4

Subfamily Orthostixinae Meyrick, 1892 5 Orthostixis Hübner, 1823 O. cribraria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/4 * O. c. amanensis Wehrli, 1932 [Turkey] * O. cinerea (Rebel, 1916) [Cyprus] * O. calcularia (Lederer, 1853) [Turkey, Georgia, ­Armenia, Azerbaijan]

Subfamily Desmobathrinae Meyrick, 1886 6 Gypsochroa Hübner, 1825 G. renitidata (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/5 Myinodes Meyrick, 1892 M. interpunctaria (Herrich-Schäffer, 1839). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/6 M. i. atlantica Hausmann, 1994 M. constantina Hausmann, 1994 7. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/183 2 Genus described after GME1 (Leraut 2002). 3 Siberian genus and species recorded as new for Europe (see information in this volume). 4 See Beljaev (2016). 5 Validated at subfamily rank in GME1, but validity of subfamily status tentatively questioned by Sihvonen et al. (2011), not yet having analysed the type genus of Orthostixinae (Orthostixis), but having represented the tribe by another genus (Naxa). The work on this question is in progress. Orthostixinae are synonymized with Desmobathrinae and tribus Desmobathrini in Beljaev (2016). 6 Genus Epirrantis and “subfamilia” Alsophilinae (treated in Vol. 1) transferred to Ennominae (see below). 7 North African species, recorded as new for Europe (see information in this volume).

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M. shohami Hausmann, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/7 * Eumegethes Staudinger, 1898 * E. tenuis Staudinger, 1898 [Morocco, Algeria, Tunisia, Libya] * Drepanopterula Hedicke, 1923 * D. zanoni (Turati, 1919) [Libya] * D. limaria (Christoph, 1885) [Caucasus (Russia), Azerbaijan?]

Subfamily Geometrinae Stephens, 1829 8, 9 Tribus Heliotheini Expósito, 1978 Heliothea Boisduval, 1840 H. discoidaria Boisduval, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/11 Tribus Pseudoterpnini Warren, 1893 Aplasta Hübner, 1823 A. ononaria (Fuessly, 1783). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/12 Holoterpna Püngeler, 1900 H. pruinosata (Staudinger, 1898). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/13 Pingasa Moore, 1887 P. lahayei (Oberthür, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/14 Pseudoterpna Hübner, 1823 P. pruinata (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/15 P. coronillaria (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/16 P. c. flamignii Hausmann, 1997 P. c. cinerascens (Zeller, 1847) * P. c. algirica Wehrli, 1929 [Morocco, Algeria, Tunisia] * P. c. axillaria Guenée, 1858 [Turkey, Levant] * P. c. halperini Hausmann, 1996 [Levant] * P. rectistrigaria Wiltshire, 1948 [Cyprus] P. corsicaria (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/17 * P. lesuraria Lucas, 1933 [Morocco] 10 Tribus Geometrini Stephens, 1829 Geometra Linnaeus, 1758 G. papilionaria Linnaeus, 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/18

8 9

Subfamily monophyletic. As a sister group of Ennominae it can remain in the herewith proposed position. Systematics at tribus level controversial. We largely follow here the subdivision of GME1, but further research is necessary. 10 Leraut (2009) raises Pseudoterpna lesuraria f. stiparia to subspecies rank, without justification, diagnosis or geographical background, thus not acceptable.

797

Tribus Comibaenini Inoue, 1961 Comibaena Hübner, 1823 C. bajularia (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/19 (= pustulata Hufnagel, 1767) * C. b. tikhonovi Hausmann, 1999 [Ciscaucasus] C. pseudoneriaria Wehrli, 1926. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/20 * C. serrulata Fletcher, 1963 [Azerbaijan?, Iran] * C. levequei Leraut, 2009 [North Africa] 11 * Microbaena Hausmann, 1996 * M. pulchra (Staudinger, 1898) [Israel] * M. p. minor Hausmann, 1996 [Sudan] Proteuchloris Hausmann, 1995 P. neriaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/21 Thetidia Boisduval, 1840 subgenus Thetidia Boisduval, 1840 T. plusiaria Boisduval, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/22 subgenus Aglossochloris Prout, 1912 * T. fulminaria (Lederer, 1870) [Iran, Kazakhstan] T. correspondens (Alpheraky, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/23 * T. crucigerata (Christoph, 1887) [Armenia, Azerbaijan] * T. euryrithra (Prout, 1935) [Jordan] subgenus Antonechloris Raineri, 1994 T. smaragdaria (Fabricius, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/24 T. s. gigantea (Millière, 1874) T. s. volgaria (Guenée, 1858) * T. persica Hausmann, 1996 [Turkey, Armenia, Azerbaijan] * T. silvia Hausmann, 1991 [Jordan] * T. bilineata Hausmann, 1991 [Jordan] T. sardinica (Schawerda, 1934) 121�������������������������������������������������������������������������������� 1/25 (Tribus Hemistolini Inoue, 1961) 13 Hemistola Warren, 1893 H. chrysoprasaria (Esper, 1795). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/26 (= immaculata: sensu auct. nec Thunberg, 1784) (= biliosata: sensu auct. nec Villers, 1789) H. c. occidentalis Wehrli, 1929 H. siciliana Prout, 1935 14 1������������������������������������������������������������������������������������������ 1/27 11 The original description presents ‘differential features’ (of just four males from the same locality) from C. pseudo­ neriaria in habitus and genitalia which, however, appear weak and require confirmation by morphology of other material and DNA barcoding. Doubtful but preliminarily to be accepted. 12 In Leraut (2009) suspected to be subspecies of T. smaragdaria without examining material (not figured) and ignoring the differential analysis in GME1. To be maintained at species rank. 13 Hemistolini subordinated under Hemitheini in Beljaev (2016) and Ban et al. (2018). Subtribe status may be considered. 14 In Leraut (2009) suspected to be subspecies of H. chrysoprasaria without justifying or presenting arguments

798

Xenochlorodes Warren, 1897 X. olympiaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/28 (= beryllaria Mann, 1853) * X. o. cremonaria (Staudinger, 1897) [Turkey] * X. nubigena (Wollaston, 1858) [Madeira] * X. magna Wolff, 1977 [Madeira] * Victoria Warren, 1897 * V. plantei Herbulot, 1976 [C. Israel] * V. eremita Hausmann, 1993 [Sinai, S. Israel] * Hierochthonia Prout, 1912 * H. pulverata (Warren, 1901) [Turkey, Lebanon] * H. semitata (Püngeler, 1902) [Israel] (Tribus Comostolini Inoue, 1961) 15 Eucrostes Hübner, 1823 E. indigenata (Villers, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/29 (= lanjeronica Hausmann, 1996) 16 (Tribus Jodini Inoue, 1961) 17 Jodis Hübner, 1823 J. lactearia (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/30 J. putata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/31 (Tribus Thalerini Herbulot, 1963) 18 Thalera Hübner, 1823 T. fimbrialis (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/32 Dyschloropsis Warren, 1895 D. impararia (Guenée, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/33 * Culpinia Prout, 1912 * C. prouti (Thierry-Mieg, 1913) [Turkey, Cyprus, Levant] Bustilloxia Expósito, 1979 * B. saturata (Bang-Haas, 1906) [Morocco, Algeria] B. s. iberica Hausmann, 1995 19 1���������������������������������������������������������������������������� 1/34

for this view. To be maintained at species rank. 15 Comostolini subordinated under Hemitheini in Beljaev (2016) and Ban et al. (2018). Subtribe status may be considered. 16 In Leraut (2009) downgraded to synonymy, justified by the statement that such forms “are found elsewhere as isolated individuals”. That view may be preliminarily accepted until detailed examination about gene-flow is available. 17 Jodini subordinated under Hemitheini in Beljaev (2016) and Ban et al. (2018). Subtribe status may be considered. 18 Thalerini subordinated under Hemitheini in Beljaev (2016) and Ban et al. (2018). Subtribe status may be considered. 19 In Leraut (2009) downgraded to synonymy, justified only by the statement that such habitus forms are found also in Morocco, ignoring differential features in male pectination and female genitalia (Hausmann 1995c; 2001). To be raised again to subspecies rank (stat.rev.).

799

Kuchleria Hausmann, 1995 K. insignata Hausmann, 1995 201������������������������������������������������������������������������������������ 1/35 (= menadiara: (partim) sensu Prout, 1912) K. menadiara (Thierry-Mieg, 1893) 216���������������������������������������������������������� 6/184 (1/35gh) * K. m. atlagenes (Prout, 1935) [Morocco] * K. m. ephedrae (Prout, 1935) [Morocco, Algeria] * K. therapaena (Prout, 1924) [Algeria, Tunisia] Tribus Hemitheini Bruand, 1846 Hemithea Duponchel, 1829 H. aestivaria (Hübner, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/36 Chlorissa Stephens, 1831 C. viridata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/37 C. cloraria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/38 * C. pretiosaria (Staudinger, 1877) [Turkey, Armenia, Azerbaijan] 22 * C. asphaleia Wiltshire, 1966 [Turkey] Phaiogramma Gumppenberg, 1887 P. etruscaria (Zeller, 1849). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/39 (= pulmentaria Guenée, 1858) P. faustinata (Millière, 1868). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/40 (Tribus Microloxiini Hausmann, 1996) 23 Microloxia Warren, 1893 M. herbaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/41 M. h. virideciliata (Bubacek, 1926) 24 M. h. advolata (Eversmann, 1837) * M. ruficornis (Warren, 1897) [North Africa, Levant] * M. simonyi (Rebel, 1894) [Canary Islands, Morocco] * M. schmitzi Hausmann, 1995 [Canary Islands, Morocco] * Acidaliastis Hampson, 1896 * A. micra Hampson, 1896 [North Africa, Levant] * A. m. galactea Rungs, 1943 [Morocco] * Acidromodes Hausmann, 1996 * A. saharae (Wiltshire, 1985) [Algeria]

20 In Leraut (2009) downgraded to “synonym or subspecies” of K. menadiara despite citation of differential characters but without bringing arguments for synonymy (“to me does not appear an authentic species”). To be raised to species rank again (stat.rev.). 21 New records for Europe (Sicily), see information in this volume. 22 Ad interim this species was recorded in the GME website as recorded from the Astrachan desert, but further investigations revealed that this record was misidentified (C. viridata). 23 Microloxiini subordinated under Hemitheini in Beljaev (2016) and Ban et al. (2018). Subtribe status may be considered. 24 In Leraut (2009) downgraded to synonymy. Genetically there are strong arguments to keep virideciliata as separate taxon (1.5% genetical distance from populations of nominate subspecies in Italy). Moreover there are differences in habitus (near to constant), and slight differences in male and female genitalia (Hausmann 2001). Therefore better to be raised again to subspecies rank (stat.rev.).

800

* Hemidromodes Prout, 1916 * H. robusta (Prout, 1913) [Algeria?] * H. r. affinis (Rothschild, 1915) [Morocco, Algeria] * H. sabulifera (Prout, 1922) [NW India] * H. s. hessa (Prout, 1935) [Levant, W Arabia, Iran]

Subfamily Ennominae Duponchel, 1845 25 Tribus Abraxini Warren, 1893 26 Abraxas Leach, 1815 27 A. grossulariata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/1 A. g. notabilis Wehrli, 1939 * A. g. conspurcata Butler, 1878 [Japan] * A. g. dsungarica Wehrli, 1939 [Central Asia] * A. g. minor Herz, 1905 [East Asia] * A. g. ribesata Staudinger, 1892 [Central Asia] A. sylvata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/2 * A. s. ijimai Inoue, 1955 [Japan] * A. s. intensa Cockerell, 1906 [Japan] * A. s. ischna Wehrli, 1939 [Russia: Sajan Mountains] * A. s. microtate Wehrli, 1931 [East Asia] A. pantaria (Linnaeus, 1867). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/3 Ligdia Guenée, 1858 L. adustata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/4 * L. lassulata (Rogenhofer, 1873) [Turkey] 28 Odontognophos Wehrli, 1951 29 O. dumetata (Treitschke, 1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/1 O. d. hibernica Forder, 1993 * O. faroulti (Rothschild, 1914) [Algeria] 30 O. margaritata (Zerny, 1927) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/2 O. zacharia (Staudinger, 1879). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/3 * O. z. dagestanensis Wehrli, 1953 [Caucasus: Dage­stan]

25 Covered by Geometrid Moths of Europe volumes 5 (Skou & Sihvonen 2015) & 6 (current volume). Arrangement of tribes considers the latest molecular results (for instance Jiang et al. (2017) and Murillo-Ramos et al. (2019), COI data, (genitalia) morphology and external features. 26 Abraxini + Cassymini + Eutoeini + Macariini form a well-supported assemblage of taxa in the multi-gene analysis, but the current taxon coverage at the genus level is too scarce to draw decisive conclusions on the classification (Murillo-Ramos et al. 2019). 27 Abraxas is listed under Macariini in Beljaev (2016). 28 Type locality is Gülek, Bulgar Dağları (Bolkar Mountains) on Taurus mountains in Turkey (Rogenhofer 1873: 572), not Bulgaria as stated in Scoble (1999). 29 Odontognophos position in the Abraxini + Cassymini + Eutoeini group supported by multi-gene analysis (Murillo-Ramos et al. in prep.) and by morphology. Odontognophos was listed under the broad concept of Macariini in Beljaev (2016). 30 Validated at species level, see this volume, Remarks under O. margaritata.

801

Tribus Cassymini Holloway, 1994 31 Lomaspilis Hübner, 1825 32 L. marginata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/5 * L. opis Butler, 1878 [Japan] * L. o. amurensis (Hedemann, 1881) [Russia: Amur] L. o. nigrita Heydemann, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/6 L. bithynica Wehrli, 1954. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/7 Stegania Guenée, 1845 33 S. cararia (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/8 * S. c. lungtanensis (Wehrli, 1939) [South-East China] S. trimaculata (Villers, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/9 * S. t. ochrearia (Bang-Haas, 1910) [North Africa] S. dilectaria (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/10 * S. d. trimaculoides (Wehrli, 1931) [Turkey] S. dalmataria Guenée, 1858. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/11 * S. d. altaica (Wehrli, 1939) [Russia: Altai] * Zamarada Moore, 1887 * Z. torrida Fletcher, 1974 [Levant, Egypt, Algeria, Africa] * Z. latilimbata Rebel, 1948 [Egypt] Dicrognophos Wehrli, 1951 34 D. sartata (Treitschke, 1827) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/4 D. siciliana Skou & Sihvonen, 2019. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/5 * D. amanensis (Wehrli, 1934) [Turkey] * D. anophaea (Wehrli, 1939) [Turkey, Iraq] 35 * D. chorista (Wehrli, 1939) [Turkey, Iraq] 36 * D. orthogonius (Wehrli, 1939) [Turkey, Iran] 37 * D. pseudosnelleni (Rjabov, 1964) [Turkey, Armenia to Turkmenistan] 38 Tribus Macariini Guenée, 1858 39 Acanthovalva Krüger, 2001 A. inconspicuaria (Hübner, 1819) 405���������������������������������������������������������������������������� 5/12 Narraga Walker, 1861 N. fasciolaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/13 31 32 33 34 35 36 37 38 39 40

802

See comments under Abraxini. Lomaspilis is listed under Macariini in Beljaev (2016). See comments under Abraxini. Stegania is listed under Macariini in Beljaev (2016). See comments under Abraxini. Position in the Abraxini + Cassymini + Eutoeini group supported by morphology (see information in this volume). Recorded for Turkey in Koçak & Kemal (2009). Recorded for Turkey in Koçak & Kemal (2009). Recorded for Turkey in Koçak & Kemal (2009). Recorded for Turkey in Koçak & Kemal (2009). See comments under Abraxini. The taxon tsekubia (Wehrli, 1940) from China: Sichuan was subordinated as subspecies under A. inconspic­ uaria in the checklist of GME5, but the text correctly states that it was recognized as bona species in Krüger (2001).

* N. f. fumipennis Prout, 1915 [China] * N. f. turkestanica Moucha & Povolny, 1957 [Turke­stan] * N. nelvae (Rothschild, 1912) [North Africa] * N. n. cappadocica Herbulot, 1943 [Turkey, Armenia] N. n. catalaunica Herbulot, 1943. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/14 N. tessularia (Metzner, 1845). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/15 * N. t. ilia Wehrli, 1940 [Central Asia] Heliomata Grote & Robinson, 1866 H. glarearia (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/16 Isturgia Hübner, 1823 I. famula (Esper, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. f. brunnea (Le Cerf, 1923) [Morocco] I. limbaria (Fabricius, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. l. anzascaria (Staudinger, 1892) I. l. delimbaria (Staudinger, 1892) I. l. nigrostriata (Heydemann, 1938) I. l. pedemontaria (Staudinger, 1892) I. l. rablensis (Zeller, 1868) I. l. styriaca (Schwingenschuss, 1911) I. l. graeca Krajcik, 2015 41 I. roraria (Fabricius, 1776) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= Palaeocrocota ostrogovichi (Caradja, 1930)) 42 I. homochromata (Mabille, 1869). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. murinaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. catalaunaria (Guenée, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. deerraria (Walker, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. disputaria (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. spodiaria (Lefèbvre, 1832). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. assimilaria (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. miniosaria (Duponchel, 1829). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. m. rungsi Leraut, 2009 [Morocco] I. berytaria (Staudinger, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sparsaria (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. messapiaria (Sohn-Rethel, 1929). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. arenacearia (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. a. mongolica (Vojnits, 1974) [Mongolia] I. punctistrigaria (Christoph, 1897). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. aspersaria (Krüger, 1939) [Libya] * I. exustaria (Staudinger, 1897) [Israel, Jordan, Sinai] * I. hausmanni Krüger, 2001 [Tunisia] * I. hedemanni (Christoph, 1885) [Iran, Near East, Turkmenistan, Uzbekistan] * I. hopffearia (Staudinger, 1878) [Turkey, Trans­caucasus] * I. perviaria (Lederer, 1855) [Middle East, southern India] * I. rubrior (Hausmann, 1990) [Morocco]

5/17 5/18

5/19 5/20 5/21 5/22 5/23 5/24 5/25 5/26 5/27 5/28 5/29 5/30 5/31 5/32

41 Probably a mountain form of Isturgia roraria (D. Stüning, pers. comm.). Work in progress. 42 Junior synonym according to Rakosy et al. (2016), previously discussed in GME5. The monotypic genus Pal­ aeocrocota Stanescu, 2002 effectively becomes a junior synonym of Isturgia.

803

* I. tennoa (Pinker, 1978) [Canary Islands] * I. terminipuncta Krüger, 2001 [Tunisia] * I. tozeurensis Leraut, 2009 [Tunisia] Itame Hübner, 1823 I. vincularia (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/33 * I. v. latefasciata Rothschild, 1914 [Algeria] * I. v. lycioidaria Herbulot, 1957 [Tunisia] * I. v. mrassinaria (Oberthür, 1923) [Morocco] * I. teknaria Powell & Rungs, 1942 [North Africa] 43 Neognopharmia Staudinger, 1892 N. stevenaria (Boisduval, 1840) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/34 * N. horhammeri (Brandt, 1941) [Iran] 44 * N. cataleucaria (Staudinger, 1901) [Turkey] 45 Gnopharmia Staudinger, 1892 46 G. irakensis Wehrli, 1938 (Turkey to Afghanistan and Pakistan) * G. rubraria Staudinger, 1892 [Turkey to Levant] * G. colchidaria (Lederer, 1870) [Turkey to Pakistan] G. c. objectaria Staudinger, 1892) 47 * G. erema Wehrli, 1930 [Turkey] 48 Macaria Curtis, 1826 49 M. notata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * M. n. kirina (Wehrli, 1940) [East Asia] * M. n. appalachiata Ferguson, 2008 [North America] M. alternata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. signaria (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * M. s. dispuncta Walker, 1860 [North America] M. liturata (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= deceptans Staudinger, 1901; = hispanica H ­ erbulot, 1957) 50 * M. l. pressaria Christoph, 1893 [Russia: Amur] * M. l. depressa (Inoue, 1955) [Japan] M. wauaria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * M. w. africana (Zerny, 1934) [Morocco, Algeria] * M. w. chinensis (Sterneck, 1928) [China]

5/35 5/36 5/37 5/38

5/39

43 See additional information in Skou & Sihvonen (2015). 44 See additional information in Skou & Sihvonen (2015). 45 This species, described from southern Turkey, cited as “Gnopharmia cataleucaria” by Koçak & Kemal (2009), but belonging to genus Neognopharmia (Rajaei et al. 2012). 46 Genus recently revised and close association to Neognopharmia diagnosed by Rajaei et al. (2015): Zootaxa 3360: 1–52. 47 Listed at species rank for Turkey in Kocak & Kemal (2009), downgraded to subspecies level in Rajaei et al. (2012). 48 Recorded for Turkey in Koçak & Kemal (2009), only confirmed for Iraq by Rajaei et al. (2012). 49 In North America, the genus name Speranza (based on the Holarctic species brunneata) was used for many years, confirmed by Ferguson (2008), followed by some North American checklists, e.g. Pohl et al. (2015). Nevertheless Speranza is a synonym of Macaria (Scoble & Krüger 2002; Skou & Sihvonen 2015; Pohl et al. 2016). Ferguson finished his manuscript in the year of 2001 (although printed much later), thus before the publication of Scoble & Krüger (2002). 50 Regarded as synonym in the text part of GME5 (though listed at subspecies rank in GME5 checklist).

804

* M. w. koreaebia (Bryk, 1949) [Korea] M. artesiaria (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/40 M. ichnusae Govi & Fiumi, 2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/41 M. carbonaria (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/42 M. brunneata (Thunberg, 1784). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/43 * M. b. imperatoria (Kardakoff, 1928) [Russia: Ussuri] * M. b. sordida (Butler, 1881) [Japan] M. loricaria (Eversmann, 1837). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/44 M. fusca (Thunberg, 1792) 51 5�������������������������������������������������������������������������������������� 5/45 * Boarmioides Lucas, 1932 52 * B. colpias (Prout, 1928) [Morocco, Algeria] Chiasmia Hübner, 1823 C. clathrata (Linnaeus, 1758) 535������������������������������������������������������������������������������������ 5/46 C. c. aurata (Turati, 1905) C. c. hugginsi Baynes, 1959 C. c. retata (Haworth, 1809) * C. c. albifenestra (Inoue, 1942) [Japan] * C. c. almacola Wehrli, 1934 [Turkey] * C. c. azrouensis (Wehrli, 1937) [North Africa] * C. c. centralasiae (Krulikowsky, 1911) [Central Asia, south Kazakhstan] * C. c. djakonovi Kardakoff, 1928 [Russia: Ussuri] * C. c. hoenei Schultze, 1954 [China] * C. c. kurilata (Bryk, 1942) [Russia: Kuril Islands] * C. c. punctata (Bang-Haas, 1910) [China] C. aestimaria (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/47 C. a. sareptanaria (Staudinger, 1871) 54 * C. a. kuldschana (Wehrli, 1940) [China] * C. a. tunesiella (Lucas, 1949) [Tunisia, North Africa?] * C. syriacaria (Staudinger, 1871) [Cyprus] 55 * C. latimarginaria (Rebel, 1907) [North Africa] * C. saburraria (Eversmann, 1851) [Russia: southern Siberia] * C. tenuiata (Staudinger, 1901) [Lebanon, potentially Syria] 56

51 See text in Skou & Sihvonen (2015) for additional details as concerns Macaria fusca (Krüger, 1939) from Libya. 52 Possibly a junior synonym of Macaria Curtis, 1826, see also Scoble & Krüger (2002). 53 See remarks under this species in GME5: validity of the putative subspecies has not been examined. 54 László et al. (2018) raise taxon sareptanaria to species status (stat. rev.), based on quantitative differences in the male and female genitalia. The mentioned male genitalia structures have been illustrated earlier by Skou & Sihvonen (2015). László et al. (2018) did not study taxon sareptanaria from its type locality (Russia: Sarepta), but instead from Iran. The very limited morphological data available cannot exclude the possiblility that the dentate plate in the male vesica varies in a clinal manner, being larger in Europe and smaller in ­Asian popu­lations. COI barcodes from 55 specimens of C. aestimaria from 15 countries do not support species status for sareptanaria. C. aestimaria splits up to five different BINs in a geographical mosaic pattern (with overlaps), none of them unequivocally matching the distribution area claimed for sareptanaria. 55 Eliminated from the European fauna list in Skou & Sihvonen (2015). Cyprus is not regarded as part of the European fauna in the GME series, whilst László et al. (2018) report this species “from Europe” based on records from Cyprus. 56 Potentially a junior synonym of C. syriacaria (Staudinger, 1871), see Skou & Sihvonen 2015).

805

* ‘Semiothisa’ Hübner, 1818 * S. erevanica (Wardikjan, 1985) [Armenia] 57 Digrammia Gumppenberg, 1887 D. rippertaria (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/48 * D. r. flavularia (Püngeler, 1902) [Uzbekistan] Rhoptria Guenée, 1858 R. asperaria (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/49 R. dolosaria (Herrich-Schäffer, 1848). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/50 Ortaliella Hausmann, 1993 O. gruneraria (Staudinger, 1862) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/51 * O. palaestinensis (Calberla, 1891) [Israel, Jordan, Lebanon] * O. stonei (Hausmann, 1991) [Jordan] O. perspersata (Treitschke, 1827) 586������������������������������������������������������������������������������ 6/6 * Stueningia Hausmann, 1993 * S. poggearia (Lederer, 1855) [Lebanon, NW Israel] * S. p. meyi Hausmann, 1993 [Syria, C Israel] * S. wolfi Hausmann, 1993 [Turkey] * S. w. taurica Hausmann, 1993 [S Turkey] Tribus Lithinini Forbes, 1948 Petrophora Hübner, 1811 P. chlorosata (Scopoli, 1763) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/53 Pachycnemia Stephens, 1829 59 P. hippocastanaria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/62 P. tibiaria (Rambur, 1829). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/63 * P. t. aindrahamaria (Oberthür, 1923) [North Africa] P. t. benesignata (Bellier, 1861) Lithinini of uncertain association Perigune Gumppenberg, 1887 P. binaevata (Mabille, 1869). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/54 * P. b. austautaria (Oberthür, 1881) [Algeria, Libya] * P. b. fraternaria (Rothschild, 1914) [North Africa] 60 P. convergata (Villers, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/55 P. c. exoglypha (Praviel, 1936) P. narbonea (Linnaeus, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/56 P. n. obliterata (Staudinger, 1901) * P. jordanaria (Staudinger, 1901) [Israel, Jordan] 57 Systematic combination uncertain. Originally combined to Semiothisa Hübner, 1818, but Semiothisa sensu stricto is restricted to the Neotropics (Scoble & Krüger 2002). 58 Transferred from Odontognophos Wehrli, 1951 (Gnophini) to Ortaliella Hausmann, 1993 (Macariini) comb. n. in the current volume. 59 Systematic position in Lithinini confirmed by the multi-gene analysis of Murillo-Ramos et al. (2019). 60 Skou & Sihvonen (2015) noted that Perigune binaevata, P. convergata and their subspecies form a species complex that needs further study. Lászlo et al. (2018, p. 39) raised taxon fraternaria to species rank, justified only by the statement: “Due to rather different external habitus, fraternaria is raised to species rank.”

806

Tribus Epirranthini McGuffin, 1981 61 Epirranthis Hübner, 1823 62 E. diversata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/8 Tribus Hypochrosini Guenée, 1858 Heterolocha Lederer, 1853 63 H. laminaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/59 * H. l. euxantha Wehrli, 1937 [East China] * H. l. lungtana Wehrli, 1940 [South East China] * H. l. sutschanska Wehrli, 1937 [Russia: Amur] Hypochrosini of uncertain association Plagodis Hübner, 1823 64 P. pulveraria (Linnaeus), 1758. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/60 * P. p. japonica (Butler, 1881) [Japan: Honshu] * P. p. jezoensis (Inoue, 1954) [Japan: Hokkaido] * P. p. montana (Inoue, 1954) [Japan: Honshu] * P. p. occiduaria (Walker, 1861) [USA] * P. p. singularis Vojnits, 1975 [Mongolia] * P. p. violacearia Prout, 1915 [Russia: Amur] 65 P. dolabraria (Linnaeus, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/61 * P. d. costisignata Wehrli, 1939 [China] Tribus Epionini Bruand, 1846 Epione Duponchel, 1829 E. repandaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/65 E. vespertaria (Linnaeus, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/66 * E. v. hespera Bryk, 1949 [Korea] Therapis Hübner, 1823 T. flavicaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/67 Epionini of uncertain association Cepphis Hübner, 1823 C. advenaria (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/52 61 Transferred from Desmobathrinae (GME1) to Ennominae. Combination with Ennominae unambiguously supported by recent molecular studies. The Ennominae position was discussed in GME5. However, the idea of Ennominae position was earlier used in McGuffin (1981) for the North America Epirranthis species, followed by Viidalepp (1996) and Kullberg et al. (2001, following Viidalepp), the last discussing larval chaetotaxy. Epirranthini is subordinated under Ennomini in Viidalepp (1996) and under Lithinini in Beljaev (2016) although E. diversata is not fern-feeding as most Lithinini. 62 Transferred from Desmobathrinae (GME1) to Ennominae, see previous footnote. 63 Heterolocha is listed under Epionini in Beljaev (2016). In multi-gene analysis Hypochrosini + Anagogini + Epionini group together, but the data is too scarce to draw decisive conclusions on the classification (Murillo-­ Ramos et al. 2019). 64 Plagodis is listed under Epionini in Beljaev (2016). See comments under Heterolocha and GME5 for additional information. 65 Compare extra information in GME5. See comments under Heterolocha.

807

Pseudopanthera Hübner, 1823 P. macularia (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/68 (= aureoadflava Stauder, 1915; = meridionalis ­Galvagni, 1908) 66 * P. m. cryptica Beljaev, 1997 [Russia: Irkutsk] * P. syriacata (Guenée, 1858) [Turkey to Levant] * Pynthanosis Turati, 1922 67 * P. henricaria (Oberthür, 1881) [Morocco, Algeria] Eilicrinia Hübner, 1823 68 E. cordiaria (Hübner, 1790) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/69 * E. c. signigera Butler, 1889 [Caucasus, Armenia, Turkey, India] E. subcordaria Herrich-Schäffer, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/70 * E. s. acardia Stichel, 1911 [Iran] E. trinotata (Metzner, 1845). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/71 Tribus Apeirini Stekolnikov & Kuznetzov, 1982 69 Apeira Gistl, 1848 A. syringaria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/75 Tribus Ennomini Duponchel, 1845 70 Ennomos Treitschke, 1825 E. autumnaria (Werneburg, 1859) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. a. koreennomos Bryk, 1949 [Korea] * E. a. pyrrosticta Wehrli, 1940 [China] * E. a. sinica Yang, 1978 [North China] E. quercinaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. alniaria (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fuscantaria (Haworth, 1809) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. f. algeriensis Prout, 1929 [North Africa] * E. f. effractaria Freyer, 1841 [southern European part of Russia] 71 E. erosaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. e. ifranaria Rungs, 1941 [Morocco] * E. e. pseudoquercaria Wehrli, 1940 [Algeria] E. quercaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. q. duercki Reisser, 1958 72 * E. q. freidbergi Hausmann, 1997 [Eastern Mediterra­nean] 66 67 68 69

5/76

5/77 5/78 5/79 5/80 5/81

Regarded synonym in the text part of GME5 (though listed at subspecies rank in GME5 checklist). Systematic position unclear, here placed near Pseudopanthera Hübner, 1823, cf. Wehrli (1939–1954) Eilicrinia is listed under Caberini in Beljaev (2016). In multi-gene analysis Apeirini (Apeira) grouped next to Epione, but the data on the potentially related Hypo­ chrosini + Anagogini + Epionini lineages are too scarce to draw decisive conclusions on the classification (Murillo-Ramos et al. 2019). 70 Beljaev (2016) listed Ourapteryx under Ennomini, thus synonymizing Ourapterygini with Ennomini. The close relationship is confirmed by the multi-gene analysis of Murillo-Ramos et al. (2019). 71 Possibly junior synonym of E. fuscantaria (Haworth), see Skou & Sihvonen (2015). 72 László et al. (2018) raised taxon duercki to species rank, justifiying it only by the statement: “In our opinion, the genitalia difference found between E. quercaria and the Cretan taxon [=duercki] justifies their separation at species rank.” For genetic patterns see GME5.

808

* E. q. lissochila (Prout, 1929) [Cyprus] * E. q. olivaria Brandt, 1938 [Iran] * E. vartianae László, 2018 [Iran] 73 * E. fraxineti Wiltshire, 1947 [South Turkmenistan, Iran] * E. kasyi László, 2018 [Armenia] 74 Ourapteryx Leach, 1814 75 O. sambucaria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/96 O. niveiscythes Zolotuhin, 2017 76 6���������������������������������������������������������������������������� 6/201 * O. malatyensis Wehrli, 1936 [Turkey] 77 * O. persica Ménétriés, 1858 [Caucasus to N. Iran] 78 (= falciformis Inoue, 1993) 79 * Coenina Walker, 1860 80 * C. dentataria Swinhoe, 1904 [Israel, Arabia, Africa] * C. islamitica Amsel, 1935 [Palestine] 81 * C. paulusi (Rebel, 1906) [Israel, Arabia] Ennomini of uncertain association Selenia Hübner, 1823 82 S. dentaria (Fabricius, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/83 S. lunularia (Hübner, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/84 * S. l. gamra Rungs, 1950 [Morocco] * S. l. kuldjana Wehrli, 1940 [China] S. tetralunaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/85 Artiora Meyrick, 1892 A. evonymaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/86 Crocallis Treitschke, 1825 83 C. tusciaria (Borkhausen, 1793). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/89 * C. t. cypriaca Fischer, 2003 [Cyprus] * C. t. taurica Wehrli, 1934 [Turkey, Eastern Mediterranean] * C. t. transcaucasica Wehrli, 1940 [Turkey, Caucasus] C. albarracina Wehrli, 1940. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/90 73 E. vartianae is very similar to E. quercaria. Species status needs proper study, the description is based on small quantitative differences only. Year of description is 2018, not 2017 as mentioned on p. 41 (László et al. 2018). 74 E. kasyi is very similar to E. fuscantaria and E. fraxineti. Species status needs proper study, because the description is based on small quantitative morphological differences only, and females are unknown. Year of description is 2018, not 2017 as mentioned on p. 41 (László et al. 2018). 75 Ourapteryx is listed under Ennomini in Beljaev (2016), thus synonymizing Ourapterygini with Ennomini. The close relationship is confirmed by the multi-gene analysis of Murillo-Ramos et al. (2019). 76 Recently described from European Caucasus and Russia, see information in this volume. 77 Not mentioned in GME5, see Zolotuhin (2017). 78 For additional information see Skou & Sihvonen (2015). 79 Synonymy according to Zolotuhin (2017). 80 Systematic position unclear, here placed in the Ennomini following Wiltshire (1990), Hacker & Hausmann (1999), and Hausmann et al 2016. The strange behaviour of rolled forewing position is reminiscent of a similar behaviour in Apochima flabellaria. 81 Possibly just a synonym of C. dentataria. 82 Selenia is listed under Epionini in Beljaev (2016). 83 Crocallis is listed under Odontoperini in Beljaev (2016).

809

C. a. centraliberica Gastón & Redondo, 2009 C. elinguaria (Linnaeus, 1758) 845���������������������������������������������������������������������������������� 5/91 C. dardoinaria Donzel, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/92 C. auberti Oberthür, 1883. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/93 C. boisduvalaria (Lucas, 1849). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/95 C. helenaria Ruckdeschel, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/94 * C. inexpectata Warnecke, 1940 [Turkey] 85 * C. i. caucasi Wehrli, 1954 [Caucasus] * C. i. cappadoica Stadie & Fiebig, 2014 [Turkey] * C. elingomorpha Stadie & Fiebig, 2014 [Caucasus] * C. merzbacheri Stadie & Fiebig, 2014 [Kazakhstan] * C. loebeli Stadie & Fiebig, 2014 [Turkey] * C. rothei Stadie & Fiebig, 2014 [Turkey] * C. sylvana Fiebig, 2014 [Turkey] * C. bacalladoi Pinker, 1978 [Canary Islands] * C. jordanaria Staudinger, 1898 [Jordan, Israel] * C. matillae Pinker, 1974 [Canary Islands] * C. rjabovi Wehrli, 1936 [Transcaucasus, Armenia] Tribus Campaeini Forbes, 1948 86 Campaea Lamarck, 1816 C. margaritaria (Linnaeus, 1761). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/110 Gerinia Leraut, 2009 G. honoraria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/111 Hylaea Hübner, 1822 H. fasciaria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/112 * H. f. cedricola (Wehrli, 1929) [Turkey, potentially Near East] * H. f. flavella (Wehrli, 1940) [Transcaucasus] H. mediterranea Sihvonen et al., 2014 87 5������������������������������������������������������������������ 5/113 H. pinicolaria (Bellier, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/114 * H. compararia (Staudinger, 1895) [North Africa] 88 * H. poeymiraui (Oberthür, 1922) [Morocco] 89 Adalbertia Wehrli, 1931 A. castiliaria (Staudinger, 1900) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/115 Pungeleria Rougemont, 1903 P. capreolaria (Denis & Schiffermüller 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/116 84 See following footnote. 85 For potential occurrence of a cryptic taxon (summarized under elinguaria) close to inexpectata and elinguaria in Europe see Skou & Sihvonen (2015). Crocallis inexpectata group was revised by Stadie & Fiebig (2014). 86 In multi-gene analysis the Campaeini is sister to the Prosopolophini + Alsophilini + Wilemanini lineage (­Murillo-Ramos et al. 2019). 87 Full authorship: Sihvonen, Skou, Flamigni, Fiumi & Hausmann. 88 See Sihvonen et al. (2014) for additional information. For discussion of the taxon ‘Hemithea squalidaria’ see op. cit and GME5 p. 391. 89 Lászlo et al. (2018) combined taxon poeymiraui with genus Pungeleria Rougemont, 1903, contradicting the proposed Hylaea combination (Leraut 2009, Skou & Sihvonen 2015).

810

Tribus Alsophilini Herbulot, 1962 90 Alsophila Hübner, 1825 A. aescularia (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1/9 A. aceraria (Denis & Schiffermüller, 1775) 911�������������������������������������������������������������� 1/10 (= quadripunctaria Esper, 1801) Tribus Prosopolophini Warren, 1894 (= Colotoini Wehrli, 1940) 92 Compsoptera Blanchard, 1845 C. opacaria Hübner, (1819) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/139 (= caesaraugustanus Redondo, 1995) 93 C. jourdanaria (Serres, 1826). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/140 C. j. anargyra (Turati, 1913) C. argentaria (Herrich-Schäffer, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/141 Colotois Hübner, 1823 94 C. pennaria (Linnaeus, 1761). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/98 C. p. carbonii Hartig, 1976 * C. p. mauretanaria (Stättermayer, 1930) [Morocco, Algeria] * C. p. paupera Hausmann, 1995 [Cyprus] * C. p. ussuriensis Bang-Haas, 1927 [Russia: Ussuri, Japan] * C. furcata Hausmann, 1995 [Israel] Dasycorsa Prout, 1915 D. modesta (Staudinger, 1879). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/88 * D. m. marerufa Wiltshire, 1940 [East-Mediterra­nean] 95 Apochima Agassiz, 1847 96 (= Bellachima Kemal et al. 2018) 97 90 Alsophilini clearly nested in Ennominae between the tribes Colotoini, Campaeini and Prosopolophini in the molecular phylogeny of Sihvonen et al. (2011), and as a sister-taxon to the Wilemanini + Prosopolophini in multi-gene analysis (Murillo-Ramos et al., in prep), therefore not supporting the separate tribe Alsophilini under Desmobathrinae (!) as suggested in Beljaev (2016). 91 In Leraut (2009) as Inurois aceraria. The combination with Inurois was proposed subjectively (“it is clear”) without indication of characters or synapomorphies. However, the shape of uncus and the much shorter terminal process of valva distinguish A. aceraria from Inurois. The presence of a cornutus is not a valuable differential feature between both genera (cf. Inoue 1943). Herewith transferred back to Alsophila (A. aceraria comb. rev.). 92 Colotoini are subordinated under Prosopolophini in Beljaev (2016). 93 See GME5, p. 462. 94 Colotois is listed under Prosopolophini in Beljaev (2016). 95 Probably synonym of D. modesta. 96 Apochima is listed under Prosopolophini in Beljaev (2016) who regards Colotoini as synonym with Prosopo­ lophini. For possible relationship to genus Coenina, see above (Ennomini of uncertain position). Relationship between Colotois, Apochima and Chondrosoma supported by shape of uncus and suggested by DNA barcode similarity. 97 A. diaphanaria combined with the newly introduced genus name Bellachima in Kemal et al. (2018). Since the molecular analysis shows several flaws (just based on the mtDNA COI gene for genus diagnosis; just based on bootstrap support and not considering the genetic distances from A. flabellaria (8.5%) and A. (d.) rjabovi; no comparison with genetic distances from other closely allied genera like Chondrosoma etc.), the validity of the genus is not accepted here.

811

A. flabellaria (Heeger, 1838). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/7 (= marmararia Buresch, 1915) * A. diaphanaria (Püngeler, 1904) [Turkey] * A. d. rjabovi (Wehrli, 1936) [Armenia to Iran] Chondrosoma Anker, 1854 98 C. fiduciaria Anker, 1854. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/8 * Dorsispina Nupponen & Sihvonen, 2013 99 * D. furcicornaria Nupponen & Sihvonen, 2013 [Kazakh­stan] Tribus Caberini Duponchel, 1845 Cabera Treitschke, 1825 C. pusaria (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/100 C. leptographa Wehrli, 1939. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/101 * C. l. birographa Viidalepp & Remm, 1982 (Russia: Khabarovsk) 100 C. exanthemata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/102 * C. e. bryantaria (Taylor, 1906) [Canada] * C. e. hamica Wehrli, 1939 [Mongolia] Tribus Baptini Forbes, 1948 Lomographa Hübner, 1825 101 L. bimaculata (Fabricius, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/104 * L. b. subnotata (Warren, 1895) [Japan] * L. b. pallidiola (Matsumura, 1931) [Japan] 102 L. temerata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/105 Tribus Theriini Herbulot, 1963 Aleucis Guenée, 1845 A. distinctata (Herrich-Schäffer, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/106 A. orientalis (Staudinger, 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/107 * A. kuznetzovi (Viidalepp, 1992) [Turkey, Levant] 103 * A. mimetes Wehrli, 1932 [Turkey, Levant] 104

98 Tentatively classified to Prosopolophini (this volume), see text for details. Relationship between Colotois, Apochima and Chondrosoma supported by shape of uncus and suggested by DNA barcode similarity. In Viidalepp (1996) placed in Bistonini between Apocheima and Agriopis. Here tentatively classified as member of Prosopolophini (new classification). 99 Potential relationships of Apochima, Chondrosoma, Dorsispina and Wilemania discussed in Nupponen & Sih­ vonen (2013), however, without assigning Dorsispina to any Ennominae tribe. Here tentatively classified as member of Prosopolophini (new classifcation). 100 Misspelled to biographa in the checklist of GME5. 101 Lomographa is listed under Caberini in Beljaev (2016). In multi-gene analysis (Murillo-Ramos et al. 2019), Lomographa is part of the Baptini + Theriini lineage, in agreement with results of Sihvonen et al. (2011). 102 Validated at subspecies rank by Scoble (1999), not examined by us. 103 Transferred from genus Epitherina, and newly recorded for the Near East in Dey et al. (2019). 104 Omitted in the checklist of GME5.

812

* Ramitia Viidalepp, 1989 105 * R. kufrana Seven, 2015 [E. Turkey] 106 Theria Hübner, 1825 T. rupicapraria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/108 * T. r. crypta Wehrli, 1940 [Russia: Caucasus] T. primaria (Haworth, 1809) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/109 Tribus Gnophini Duponchel, 1845 Gnophos Treitschke, 1825 G. furvata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/9 G. obfuscata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/10 (=androgynus Reisser, 1936; = lutescentaria Staudinger, 1871; = marsicaria Dannehl, 1933; = myrtillata Thunberg, 1788; = nivea Schawerda, 1913) Charissa Curtis, 1826 nomen protectum 107 (= Hyposcotis Hübner, 1825 nomen oblitum) Subgenus Charissa Curtis, 1826 C. obscurata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/11 (= argillacearia Staudinger, 1871; = calceata Staudinger, 1861; = lafauryata Oberthür, 1913; = pullaria Hübner, 1799) C. o. bellieri (Oberthür, 1913) * C. canariensis (Rebel, 1911) [Canary Islands: Tenerife, below 1200 m] * C. c. teidensis Pinker, 1969 [Canary Islands: Tenerife, Teide: above 2000 m] * C. c. bartolomensis Pinker, 1969 [Canary Islands: Gran Canaria] * Subgenus Sacrognophos Wehrli, 1951 * C. sacraria (Staudinger, 1894), comb. n. [Israel, Turkey, Algeria, Libya, Egypt] 108 * C. crosi Thierry-Mieg, 1910, comb. n. [Algeria, Morocco] 109 * C. similaria (Rothschild, 1914), comb. n. [Algeria, Libya] 110 Subgenus Costignophos Wehrli, 1951 C. crenulata (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/12 C. c. aragonensis (Zerny, 1927) C. c. avilarius (Reisser, 1936) C. c. pyrenaica (Oberthür, 1913) C. italohelveticus (Rezbanyai-Reser, 1986). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/13 105 Assigned to tribe Theriini in Viidalepp (1996). 106 Probably subspecies of R. ghirshmani (or even synonym), cf. Dey et al. (2019). 107 Koçak & Kemal (2009) suggest to validate the genus name Hyposcotis, Erlacher & Erlacher (2017a) propose to suppress that name in relation to Charissa, whenever the two are considered to be synonyms, currently pending a decision from the Commission (ICZN). Checklist provided by S. Erlacher, supplemented by A. Hausmann. New and revised combinations with subgenera of Charissa suggested by morphology (­Erlacher, publication(s) in preparation). 108 Combination with subgenus Sacrognophos and subordination under Charissa suggested by morphology (Erlacher in prep.). 109 Combination with subgenus Sacrognophos and subordination under Charissa suggested by morphology (Erlacher in prep.). 110 Combination with subgenus Sacrognophos and subordination under Charissa suggested by morphology (Erlacher in prep.).

813

C. pullata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/14 (= albarinata Millière, 1859; = confertata Staudinger, 1871; = impectinata Guenée, 1858; = kovacsi Vojnits, 1967; = nubilata Fuchs, 1872) Subgenus Dysgnophos Wehrli, 1951 C. difficilis (Alphéraky, 1883). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/15 C. d. teberdensis (Müller, 1978) [western Caucasus] * C. d. urmensis (Wehrli, 1953) [eastern Caucasus] C. turfosaria (Wehrli, 1922). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/16 (= macguffini Smiles, 1979) Subgenus Cnestrognophos Wehrli, 1951 C. mutilata (Staudinger, 1878). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/17 C. peloponnesiaria Erlacher & Erlacher, 2017. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/18 C. pentheri (Rebel, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/19 (= petrina Thurner, 1936) * C. luticiliata (Christoph, 1889) comb. n. [Turkey, Transcaucasus, Turkmenistan, Iran] 111 * C. l. subvariegata (Staudinger, 1898) [Israel, Jordan, Egypt] (= claytoni Wiltshire, 1948) * C. annubilata (Christoph, 1885) comb. n. [Turkey, Georgia, Azerbaijan, Iran] 112 * C. a. darashama (Wehrli, 1953), comb. n. [Trans­caucasus] 113 * C. anthina (Wehrli, 1953), comb. n. [Turkey, Trans­caucasus] 114 * C. effendii (Viidalepp & Pirijev, 1993), comb. n. [Transcaucasus: Azerbaijan] 115 * C. nannodes (Wehrli, 1936), comb. n. [Turkey] 116 * C. adjectaria (Staudinger, 1897), comb. n. [Turkey, Iran, Israel, Jordan] 117 * C. a. armenia (Wehrli, 1953) [Armenia] * C. a. daritshevae Viidalepp, 1992) [Turkmenistan] * C. rjabovi (Wehrli, 1939), comb. n. [Turkey, Trans­caucasus] 118 * C. libanotica (Wehrli, 1931), comb. n. [Turkey to Israel] 119 * C. quadrimaculata (Chrétien, 1913), comb. n. [­Algeria, Libya] 120 * C. liliputaria Turati, 1934, comb. n. [Libya] 121 Subgenus Euchrognophos Wehrli, 1951 C. assoi (Redondo & Gastón, 1996). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. corsica (Oberthür, 1913). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. staudingeri (Wnukowsky, 1929). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. subtaurica (Wehrli, 1934) (nec Wehrli, 1932). . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. variegata (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. dubitaria (Staudinger, 1892) [Turkey, Transcaucasus, Iran] 122 111 112 113 114 115 116 117 118 119 120 121 122

814

Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Combination with subgenus Cnestrognophos suggested by morphology (Erlacher in prep.). Described from Turkey.

6/20 6/21 6/22 6/23 6/24

* C. symmicta (Wehrli, 1953), comb. n. [Turkey, Caucasus, Transcaucasus] 123 * C. lequatrei (Herbulot, 1988) [Jordan, Yemen] * C. recticostaria (Turati, 1926) nomen dubium [­Libya] 124 C. mucidaria (Hübner, 1799) 125/���������������������������������������������������������������������������������� 6/25 * C. rothschildi (Prout, 1929) [Morocco] 126 * C. porphyratus (Zerny, 1936) [Morocco] Subgenus Kemtrognophos Wehrli, 1951 C. ambiguata (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. a. herbuloti Leraut, 2009 127 C. a. sheljuzhkoi (Schawerda, 1924) C. a. pullularia (Herrich-Schäffer, 1852) C. predotae (Schawerda, 1929) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. onustaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. o. serraria (Guenée, 1858) * C. ciscaucasica (Rjabov, 1964), comb. n. [Turkey, Caucasus, Transcaucasus] 128 C. remmi (Viidalepp, 1988). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. zeitunaria (Staudinger, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. mardinaria (Staudinger, 1901) [Turkey to Levant] * C. argillata (Brandt, 1938), comb. n. [Turkey, Iran] 129 * C. idrisarius (Zerny, 1934) [Morocco] 130

6/26

6/27 6/28 6/29 6/30

Subgenus Neognophina Wehrli, 1946 (= Rhopalognophos Wehrli, 1951) C. glaucinaria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/31 C. pfeifferi (Wehrli, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/32 C. supinaria (Mann, 1854) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/33 C. s. intermedia Wehrli, 1917 (= gulsensis Wolfsberger, 1959; = f­rancothuringiaca Bergmann, 1955; = dioszeghyi Kovács, 1954; = budensis Kovács, 1954; = benestriata Kovács, 1954) Subgenus Trilobignophos Wehrli, 1951 C. graecaria (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/34 (= certhiatus Rebel & Zerny, 1931; = minorasiaticus Wehrli, 1936) * C. pallescens (Rjabov, 1964) [Turkey, Caucasus: Dagestan] 131 * C. talyshensis (Wehrli, 1936), comb. n. [Caucasus, Transcaucasus, Iran] 132 * C. zejae (Wehrli, 1953) [Caucasus, Transcaucasus] 133 123 Combination with subgenus Euchrognophos suggested by morphology (Erlacher in prep.). 124 Detailed discussion will be published by Erlacher (in prep.) 125 The name Hyposcotis, based on the type species C. mucidaria, is proposed by Erlacher & Erlacher (2017) to be suppressed in relation to Charissa, whenever the two are considered to be synonyms, currently pending a decision from the Commission (ICZN). Further research needs to clarify whether C. mucidaria and the following two species can remain in subgenus Euchrognophos or not. 126 Raised from subspecies rank (under C. variegata) to species rank (see text in this volume). 127 Downgraded from species rank (see text in this volume). 128 Combination with subgenus Kemtrognophos suggested by morphology (Erlacher in prep.). 129 Combination with subgenus Kemtrognophos suggested by morphology (Erlacher in prep.). 130 Systematic position of C. idrisarius within the genus Charissa is still unclear (Erlacher in prep.). 131 Combination with subgenus Trilobignophos suggested by morphology (Erlacher in prep.). 132 Combination with subgenus Trilobignophos suggested by morphology (Erlacher in prep.). 133 Combination with subgenus Trilobignophos suggested by morphology (Erlacher in prep.).

815

* Subgenus Organognophos Wehrli, 1951 * C. wanensis (Wehrli, 1939), comb. n. [Turkey] 134 * C. asymmetra (Wehrli, 1939), comb. n. [Trans­caucasus to Iran] 135 Psodos Treitschke, 1825 136 (= Glacies Millière, 1874) P. alpinata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/35 P. canaliculata (Hochenwarth, 1785). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/36 P. c. schwingenschussi Wehrli, 1919 P. werneri Schawerda, 1916. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/37 P. coracina (Esper, 1805) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/38 (= transiens Wehrli, 1921; = tundranoides Malicky, 1966; = pseudonoricana Wehrli, 1921) P. c. bureschi Varga, 1975 P. c. dioszeghyi Schmidt, 1930 P. c. lappona Wehrli, 1921 * P. c. tundrana Wehrli, 1919 [central Asia] P. bentelii (Rätzer, 1890) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/39 P. b. perlinii Turati, 1914 P. b. retyezatensis Bartha, 1933 P. noricana Wagner, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/40 (= kusdasi Wehrli, 1945; = burmanni Tarmann, 1984) P. n. carpathica Schwingenschuss, 1915 P. belzebuth Praviel, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/41 P. pyrenaica Schawerda, 1919. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/42 P. alpmaritima Wehrli, 1924. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/43 P. alticolaria Mann, 1853. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/44 (= chalybaeus Zerny, 1916) P. a. faucium Favre, 1905 P. a. gedrensis Rondou, 1903 P. wehrlii Vorbrodt, 1918. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/45 P. baldensis Wolfsberger, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/46 P. spitzi Rebel, 1907. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/47 P. quadrifaria (Sulzer, 1776). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/48 P. q. pyrenaea Oberthür, 1884 P. q. sudetica Sterneck, 1918 Sciadia Hübner, 1822 S. tenebraria (Esper, 1806). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. t. wockearia (Staudinger, 1871) S. t. taurusica Huemer & Hausmann, 2009 S. septaria (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. innuptaria (Herrich-Schäffer, 1852) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. slovenica Leraut, 2008. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dolomitica Huemer & Hausmann, 2009. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134 Combination with subgenus Organognophos suggested by morphology (Erlacher in prep.). 135 Combination with subgenus Organognophos suggested by morphology (Erlacher in prep.). 136 Checklist provided by S. Erlacher & H. Rajaei.

816

6/49 6/50 6/51 6/52 6/53

S. zelleraria (Freyer, 1836) 137/������������������������������������������������������������������������������������ 6/54 S. unicoloraria (Staudinger, 1871) 138/�������������������������������������������������������������������������� 6/55 Elophos Boisduval, 1840 E. caelibaria (Heydenreich, 1851). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. c. spurcaria (de la Harpe, 1853) E. zirbitzensis (Pieszcek, 1902). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. operaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. o. hoefneri (Rebel, 1903) E. o. necopinatus (Rezbanyai-Reser, 1992) E. andereggaria (de la Harpe, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Yezognophos Matsumura, 1927 (=Parietaria Leraut, 1981) Y. dilucidaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Y. sproengertsi (Püngeler, 1914). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Y. vittaria (Thunberg, 1788) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Y. v. mendicaria (Herrich-Schäffer, 1852) * Y. v. kononis Matsumura, 1927 [Japan] Y. serotinaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Y. s. dognini (Thierry-Mieg, 1910) Y. s. serotinoides (Wehrli, 1922) Crocota Hübner, 1823 C. tinctaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. t. estachyi Leraut, 1999 C. t. vintejouxi Leraut, 1999 C. peletieraria (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pseudotinctaria Leraut, 1999. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. niveata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

6/56 6/57 6/58 6/59

6/60 6/61 6/62 6/63

6/64 6/65 6/66 6/67

* Scodionista Joannis, 1912 * S. mischii (Turati & Krüger, 1936) [Libya] * S. amoritaria (Püngeler, 1902) [Egypt, Levant, Transcaucasus to Afghanistan] * S. a. irrorata Wiltshire, 1990 [Saudi Arabia] * Synopsidia Djakonov, 1935 * S. phasidaria (Rogenhofer, 1873) [Transcaucasus to Afghanistan] Synopsia Hübner, 1825 139 S. sociaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/68 (=almasa Schawerda, 1912; = propinquaria ­Boisduval, 1840; = unitaria Prout, 1915) Siona Duponchel, 1829 S. lineata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/117 Hypoxystis Prout, 1915 H. pluviaria (Fabricius, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/73 137 Transferred from Elophos to Sciadia, see text (this volume) for details. 138 Transferred from Elophos to Sciadia, see text (this volume) for details. 139 Gnophini position is supported by multi-gene analysis (Murillo-Ramos et al. 2019). Synopsia is listed under Gnophini in Beljaev (2016).

817

Angerona Duponchel, 1829 A. prunaria (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/99 * A. p. aquafortis Bryk, 1949 [Korea] * A. p. coreola Wehrli, 1940 [Korea] * A. p. dominans Bryk, 1942 [Russia: Kuril Islands] * A. p. kentearia Staudinger, 1892 [Mongolia] * A. p. mongoligena Bryk, 1949 [northern Mongolia] * A. p. prouterona Bryk, 1949 [Korea] * A. p. turbata Prout, 1929 [Japan] * A. p. valens Wehrli, 1940 [Russia: Ussuri, Korea] Phthonandria Warren, 1894 140 P. strictaria (Lederer, 1839) 141/������������������������������������������������������������������������������������ 6/69 Chariaspilates Wehrli, 1953 C. formosaria (Eversmann, 1837). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/118 Aspitates Treitschke, 1825 A. mundataria (Stoll, 1782) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * A. m. tonghata (Felder & Rogenhofer, 1875) [Japan] * A. m. uncinataria (Vojnits, 1975) [Mongolia] A. gilvaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. g. burrenensis Cockayne, 1951 * A. g. kukunorensis (Wehrli, 1953) [China] * A. g. minimus (Vojnits, 1975) [Mongolia] * A. g. orientaria (Alphéraky, 1883) [Central Asia] * A. taylorae (Butler, 1893) [Canada] A. t. sibirica (Djakonov, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. orciferaria (Walker, 1862). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * A. o. baffinensis (Munroe, 1963) [Canada] * A. o. churchillensis (Munroe, 1963) [Canada] * A. o. occidentalis (Munroe, 1963) [Canada] A. forbesi (Munroe, 1963) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. albaria (Bartel, 1903) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ochrearia (Rossi, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * A. acuminaria (Eversmann, 1851) [Caucasus, Georgia, Dagestan] * A. collinaria (Holt-White, 1894) [Canary Islands]

5/119 5/120

5/121 5/122

5/123 5/124 5/125

Dyscia Hübner, 1825 D. raunaria (Freyer, 1851). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/126 D. royaria Tautel & Billi, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/127 D. govii Fiumi & Flamigni, 2013 142/���������������������������������������������������������������6/202 (5/128f) * D. innocentaria (Christoph, 1885) [Turkmenistan] * D. i. osmanica (Wagner, 1931) [Turkey, Cyprus to Israel] D. i. sicanaria (Oberthür, 1923). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/128

140 Phthonandria position in the Gnophini is supported by multi-gene analysis (Murillo-Ramos et al. 2019). 141 Transferred from Megalycinia to Phthonandria, see text (this volume) for details. 142 In GME5 regarded as subspecies of D. innocentaria, upgraded to species rank again in Flamigni et al. (2016), here confirmed (see information in this volume).

818

* D. malatyana Wehrli, 1934 [Turkey to Jordan and Israel] D. m. albersaria Warnecke, 1940 [Turkmenistan] D. m. clara Pugaev & Zolotuhin, 2006. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * D. m. senecai Wiltshire, 1990 [Libya] * D. m. dagestana Wehrli, 1934 [Transcaucasus] 143 D. crassipunctaria (Rebel, 1916). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. conspersaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . * D. c. sultanica Wehrli, 1936 [Turkey] D. c. turturaria (Boisduval, 1840) D. fagaria (Thunberg, 1784). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . D. f. albidaria (Staudinger, 1871) D. f. emucidaria (Hübner, 1813) D. f. favillacearia (Hübner, 1799) D. penulataria (Hübner, 1819). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * D. p. atlasica Leraut, 2009 [Morocco, Algeria)] * D. p. combustaria (Oberthür, 1923) [Algeria] D. lentiscaria (Donzel, 1837) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * D. l. agacles (Oberthür, 1923) [Algeria] D. distinctaria (Bang-Haas, 1910). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * D. atlantica (Reisser, 1933) [Morocco] * D. galactaria Turati, 1934 [North Africa] * D. holli (Oberthür, 1910) [North-West Africa] * D. negrama Wehrli, 1953 [Turkey, Caucasus] * D. nobiliaria (Bang-Haas, 1906) [North Africa] * D. plebejaria (Oberthür, 1910) [North-West Africa] * D. rungsi Herbulot, 1981 [Morocco] * D. simplicaria (Rebel, 1939) [Cyprus]

5/129 5/130 5/131 5/132

5/133 5/134 5/135

Perconia Hübner, 1823 P. strigillaria (Hübner, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/136 P. baeticaria (Staudinger, 1871) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/137 Menophra Moore, 1887 144 M. abruptaria (Thunberg, 1792). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= dalmata Galvagni, 1907) * M. a. maura (Oberthür, 1913) [Algeria, Tunisia] * M. a. canariensis (Rebel, 1917) [Canary Islands] M. japygiaria (O. Costa, 1849). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= fidelensis Mendes, 1909) M. berenicidaria (Turati, 1924). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M. b. cretacaria (Reisser, 1958) M. harterti (Rothschild, 1912). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * M. h. dresnayi (D. Lucas, 1932) [Morocco: Moyen Atlas] 145 * M. h. dubiosa (Albers & Warnecke, 1941) [­Morocco: Moyen Atlas; Algeria] 146

6/70

6/71 6/72 6/73

143 See Trusch & Erlacher (2002). 144 Gnophini according to Jiang et al. (2017) and Beljaev (2016), and confirmed by multi-gene analysis (Murillo-­ Ramos et al. 2019). 145 Subspecies rank according to Rungs (1981). 146 Subspecies rank according to Rungs (1981), synonym in Scoble (1999).

819

* M. h. undulosa (Albers & Warnecke, 1941) [Morocco: Western plains] 147 * M. tameliltensis Leraut, 2009 [Morocco] M. thuriferaria (Zerny, 1927) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/74 * M. maderae (Bethune-Baker, 1891) [Madeira] M. nycthemeraria (Geyer, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/75 (= dinicola Wehrli, 1941) M. annegreteae Skou, 2007 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/76 Cleorodes Warren, 1894 148 C. lichenaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/77 C. l. distinctaria (Hanson, 1962) C. l. jaspidaria (Wagner, 1919) Gnophini of uncertain association Zernyia Prout, 1929 Z. granataria (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/78 * Z. maroccana Reisser, 1933 [Morocco] * Z. algiricaria Lucas, 1940 [Algeria] * Z. annularis Prout, 1929 [Algeria] * Z. gnophoides Prout, 1929 [Algeria] * Z. nelvai Prout, 1929 [Algeria] * Z. selidosema Prout, 1929 [Algeria] Megalycinia Wehrli, 1939 149 M. serraria (A. Costa, 1882). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/79 * M. scalaria (Christoph, 1893) [Turkey, Transcaucasus: Georgia] * M. engys (Wehrli, 1939) [Turkey, Iraq] 150 Dasypteroma Staudinger, 1892 D. thaumasia Staudinger, 1892. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/80 * D. t. powelli (Oberthür, 1914) [Morocco, Algeria] * Wehrliola Strand, 1932 151 * W. revocaria (Staudinger, 1892) [Turkey to Levant] * W. inexpectata Kemal & Uçak, 2018 [eastern Turkey] 152

147 Subspecies rank according to Rungs (1981), synonym in Scoble (1999). 148 Gnophini according to Viidalepp et al. (2007), confirmed by multi-gene analysis (Murillo-Ramos et al. 2019), associating closely with Siona and Menophra. 149 Megalycinia has earlier been classified as Boarmiini. 150 Recorded for Turkey in Koçak & Kemal (2009). 151 Wehrliola is structurally very similar to Megalycinia (P. Sihvonen, unpublished). 152 Recently described from Hakkari prov., see Kemal & Uçak (2018).

820

Tribus Odontoperini Duponchel, 1845 153 Odontopera Stephens, 1831 O. bidentata (Clerck, 1759) 154/������������������������������������������������������������������������������������ 5/87 * O. b. exsul (Tshetverikov, 1905) [Russia: Sajan] * O. b. asiatica (Bang-Haas, 1907) [Russia: Sajan] 155 * O. b. harutai (Inoue, 1953) [Japan: Honshu] * O. b. kurilana (Bryk, 1942) [Russia: Kuril Islands] * O. b. rava (Vojnits, 1975) [Mongolia] Tribus Boarmiini Duponchel, 1845 (= Bistonini Stephens, 1850; = Erannini Tutt, 1896; = Phaseliini Wehrli, 1941; = Bupa­ lini Herbulot, 1963; = Cleorini Duponchel, 1845) 156 Microbiston Staudinger, 1882 M. lanaria (Eversmann, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/81 * M. phaeothorax Wehrli, 1941 [Caucasus: Dagestan, Transcaucasus: Armenia] * M. turanicus Staudinger, 1900 [Kazakhstan, Uzbekistan, Turkmenistan] 157 Apocheima Hübner, 1825 158 A. hispidaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/82 A. h. cottei (Oberthür, 1913) (= popovi Vojnits, 1972) 159 * A. h. orientis Wehrli, 1941 [Turkey, Caucasus, Trans­caucasia] * A. cinerarius (Erschoff, 1874) [Kazakhstan to China] Phigalia Duponchel, 1829 160 P. pilosaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/83 * P. buckwelli Rungs, 1952 [Morocco] 161 * Pterotocera Staudinger, 1882 162 * P. armeniacae Djakonov, 1949 [Armenia, C. Asia] * P. declinata Staudinger, 1882 [Kazakhstan]

153 Odontopera is listed under ‘Ennomini of uncertain association’ in Sihvonen & Skou (2015). Here we classify Odontopera in Odontoperini, as a sister-taxon to Gnophini, supported by multi-gene analysis (Murillo-Ramos et al. in prep.). In the analysis, Odontopera groups together with the Afro-Arabian genus Henicovalva Krüger, 2017, and numerous Neotropical genera, all these are part of a large Nacophorini assemblage, which is rather closely related to Gnophini. The molecular results allow Odontoperini to be treated valid at tribus-level, in agreement with Beljaev (2016). 154 The taxon graecarius is synonymized in Skou & Sihvonen (2015). 155 Subspecies status after Scoble (1999); both exsul and asiatica described from Sajan mts., thus validity at subspecies level doubtful. 156 For synonymy see recent molecular analyses (e.g. Wahlberg et al. 2010; Sihvonen et al. 2011; Jiang et al. 2017; Murillo-Ramos et al. 2019). 157 Subsp. rank under M. lanaria in Leraut (2009). 158 Apocheima and Phigalia are sister taxa, related to Hypomecis and Ematurga in several multi-gene analyses (Wahlberg et al. 2010; Jang et al. 2017; Murillo-Ramos et al. 2019). 159 For synonymy see text in this volume. 160 See comment under Apocheima. 161 Placed in Apocheima in Rungs (1981), in Phigalia in Scoble (1999). 162 Pterotocera synomized with Phigalia in Scoble (1999).

821

Hypomecis Hübner, 1821 163 (= danieli (Wehrli, 1936)) H. roboraria (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/84 H. punctinalis (Scopoli, 1763) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/85 (= grisearia Bastelberger, 1902; = magyarica Wehrli, 1932) * H. p. kerstinae Fischer & Lewandowski, 2004 [Cyprus, Turkey] Ematurga Lederer, 1853 164 E. atomaria (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/86 (= alpicolaria Vorbrodt & Müller-Rutz, 1917; = arenaria Candèze, 1926; = isoscelata ­Scopoli, 1763; = minuta Heydemann, 1925; = orientaria Staudinger, 1861; = zetter­ stedtaria Heydemann, 1930) * E. a. iliaria Alphéraky, 1883 [Armenia] Lycia Hübner, 1825 165 L. hirtaria (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/87 (= cataloniae Vojnits & Mészáros, 1973; = diniensis Oberthür, 1913; = istriana G ­ alvagni, 1901; = pusztae Vojnits, 1971; = uralaria Krulikovski, 1909) * L. incisarius (Lederer, 1870) [Georgia, Armenia] L. necessaria (Zeller, 1849). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/88 (= grupchei Carnelutti, 1986; = inversarius Rebel, 1903) L. lapponaria (Boisduval, 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/89 L. l. scotica (Harrison, 1916) L. isabellae (Harrison, 1914). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/90 (= strobioni Dujardin, 1965) * L. liquidaria (Eversmann, 1848) [Kazakhstan] L. pomonaria (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/91 L. zonaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/92 (= atlantica Harrison, 1938; = britannica Harrison, 1912; = rossica Harrison, 1912) L. alpina (Sulzer, 1776). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/93 L. a. carniolica (Harrison, 1912) L. graecarius (Staudinger, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/94 (= istrianus Staudinger, 1901; = odrinarius Buresch, 1915) L. florentina (Stefanelli, 1882). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/95 (= italica Harrison, 1912) Biston Leach, 1815 B. strataria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= meridionalis Oberthür, 1913) B. rosenbaueri Müller, 2018. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. achyra Wehrli, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . B. betularia (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * B. b. fumosarius (Alpherakyi, 1897) [Transcaucasia: Armenia, Azerbaijan] * B. b. alexandrinus Wehrli, 1941 [Kazakhstan, Kyrgyzstan]

6/96 6/97 6/98 6/99

163 Several multi-gene analyses revealing close relationship between Hypomecis and Ematurga, both falling in the same cluster as Phigalia and Apocheima (Jiang et al. 2017; Murillo-Ramos et al. 2019). 164 See footnote to previous genus. 165 Lycia associates in several multi-gene analyses with non-European genera (Jiang et al. 2017; Murillo-Ramos et al. 2019).

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Erannis Hübner, 1825 166 E. defoliaria (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/100 (= teriolensis Dannehl, 1933) * E. caspica Laszlo, 2003 [Azerbaijan, Iran] E. ankeraria (Staudinger, 1861) 167/���������������������������������������������������������������������������� 6/101 (= bervaensis Jablonkay, 1965) * E. a syriaca (Wehrli, 1934) [Turkey, Syria] E. declinans (Staudinger, 1879) 168/���������������������������������������������������������������������������� 6/102 * Desertobia Viidalepp, 1989 * D. heloxylonia Xue et al., 2006 [NW. China] * D. h. lawrencei Stadie & Lehmann, 2012 [Jordan] Kresnaia Leraut, 2009 K. beschkovi (Ganev, 1987). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/103 Agriopis Hübner, 1825 A. leucophaearia (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/104 A. bajaria (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/105 (= coggii Constantini, 1916) * A. b. zehrounensis Wehrli, 1940 [Morocco] 169 A. aurantiaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/106 (= cleui Leraut, 1993; = lariciaria Scholz, 1947) A. budashkini Kostjuk, 2009 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/107 A. marginaria (Fabricius, 1776). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/108 (= pallidata Turati, 1911) Calamodes Guenée, 1858 C. occitanaria (Duponchel, 1829). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/109 (= submelanaria Zerny, 1927) C. melanaria Oberthür, 1913 170 * C. m. melanaria Oberthür, 1913 [North Africa] C. m. subscudularia Turati, 1919. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/110 * C. boursini (Albers & Warnecke, 1941) [Morocco] 171 Paradarisa Warren, 1894 P. consonaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/111 Larerannis Wehrli, 1935 172 L. orthogrammaria (Wehrli, 1927). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/112

166 Erannis is listed under Erannini in Viidalepp (1996), in this volume listed as synonym of Boarmiini, following all recent molecular analyses (e.g. Wahlberg et al. 2010; Sihvonen et al. 2011; Jiang et al. 2017; Murillo-­ Ramos et al. 2019). 167 Placed in Agriopis in Viidalepp (1996). Leraut (2009) and Nahirnik & Beschkov (Acta Ent. Serbica 2016 (21): 143–146) combine ankeraria with Desertobia. Not supported, however, by COI barcodes. 168 Placed in Agriopis in Viidalepp (1996). 169 Synonymized under A. bajaria bajaria in Scoble (1999). 170 Validated at subspecies rank under C. occitanaria in Rungs (1981). 171 See Rungs (1981). 172 The position of Larerannis is tentative. Beljaev (2016) placed it between Arichanna and Phigalia.

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Chemerina Boisduval, 1840 C. caliginearia (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/113 (= adriatica Schwingenschuss & Wagner, 1927) * C. c. punctinervis Rothschild, 1914 [Algeria] Fagivorina Wehrli, 1943 F. arenaria (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/114 Spartopteryx Guenée, 1858 S. kindermannaria Staudinger, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/115 Athroolopha Lederer, 1853 A. pennigeraria (Hübner, 1813) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/116 * A. p. atlanticaria (Le Cerf, 1923) [Morocco] * A. p. latimargo Rothschild, 1914 [Morocco, western Algeria] * A. p. orientalis (Rothschild, 1925) [eastern Algeria] * A. kabylaria (Oberthür, 1878) [Algeria] 173 A. chrysitaria (Hübner, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/117 Eurranthis Hübner, 1823 174 E. plummistaria (Villers, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/118 (= albosignata Neuburger, 1907) * E. p. algeriensis (Rothschild, 1925) [Algeria] * E. p. atlanticaria (D. Lucas, 1920) [Morocco] Nychiodes Lederer, 1853 175 N. obscuraria (Villers, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/119 N. ragusaria Millière, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/120 N. andalusiaria Millière, 1865 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/121 (= estrellae Wehrli, 1934) N. notarioi Expósito, 2005. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/122 * N. mauretanica Wehrli, 1929 [Morocco to Tunisia] * N. palaestinensis Wagner, 1919 [Lebanon, Israel, Jordan] * N. persuavis Wehrli, 1929 [Lebanon, Israel] * N. muelleri Hausmann, 1991 [Jordan, Israel] N. hispanica Wehrli, 1929. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/123 N. h. torrevinagrensis Expósito, 1984 N. h. atlanticaria Schwingenschuss, 1936 176 N. waltheri Wagner, 1919. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/124 (= osthelderi Wehrli, 1929; = syriaca Wehrli, 1929; = cuencaensis Leraut, 2009 177) * N. w. transcaspia Wehrli, 1929 [Turkmenistan] * N. w. saerdabica Wehrli, 1938 [North Iran] 178 173 Subordinated as subsp. under A. pennigeraria in Scoble (1999), Validated at species rank in Leraut (2009). 174 Multi-gene analysis and genitalia reveal both genera Eurranthis and Nychiodes as closely related (cf. Murillo-­ Ramos et al. in prep.). 175 See footnote to previous genus. 176 Regarded as subsp. of N. obscuraria in Scoble (1999). 177 Leraut (2009) described this species based on single female holotype from Spain (Castilia). The genitalia of the holotype are fully identical with those of N. waltheri. However as N. waltheri is not distributed in western Europe, the female specimen may be a case of mislabelling. 178 Confirmation of this subspecies needs further study based on larger series.

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N. amygdalaria (Herrich-Schäffer, 1848). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/125 (= almensis Wehrli, 1941; = malatyaca Wehrli, 1941) N. dalmatina Wagner, 1909. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/126 * N. divergaria Staudinger, 1892 [Turkey] * N. d. fallax Wehrli, 1941 [Transcaucasus: Azerbaijan] * N. d. achtyca Wehrli, 1941 [Caucasus: Dagestan] * N. rayatica Wiltshire, 1957 [Iraq, Armenia, Turkey] * N. variabila Brandt, 1938 [Turkey, Iran] 179 Phaselia Guenée, 1858 180 P. serrularia (Eversmann, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/127 * P. s. shurensis Wehrli, 1941 [Dagestan, Georgia, Azerbaijan] * P. s. catharia Wehrli, 1941 [Armenia, Turkey?] P. algiricaria Oberthür, 1913 * P. a. algiricaria Oberthür, 1913 [Algeria] P. a. aragona Wehrli, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/128 * P. a. gigantaria Schwingenschuss, 1936 [Morocco] * P. a. relicta (Turati, 1930) [Libya: Cyrenaica] * P. joestleinae Hausmann, 1996 [Turkey] * P. deliciosaria (Lederer, 1853) [Lebanon to Egypt] * P. phoeniciaria (Turati & Krüger, 1936) [Libya: Cyrenaica] Ekboarmia Wehrli, 1943 181 E. atlanticaria (Staudinger, 1859). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/129 (= gracilis Albers & Warnecke, 1941) [Morocco] 182 * E. a. holli (Oberthür, 1909) [Algeria] * E. fascinataria (Staudinger, 1900) [Morocco, Algeria] 183 E. sagnesi Dufay, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/130 (= herrerai Expósito, 2007) 184 E. miniaria Skou, Stüning & Sihvonen, 2017. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/131 Ecleora Wehrli, 1941 185 E. solieraria (Rambur, 1834). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/132 * E. haroldaria (Oberthür, 1913) [Algeria, Morocco] 186

179 Recorded for Turkey in Koçak & Kemal (2009). 180 Phaselia is listed under Phaseliini in Viidalepp (1996; misspelled to Phaselini), in this volume listed as synonym of Boarmiini, following all recent molecular analyses (e.g. Sihvonen et al. 2011; Jiang et al. 2017; Murillo-­ Ramos et al. 2019). 181 Taxonomy of Ekboarmia follows Skou et al. (2017): Nota lep. 30: 39–63. Ekboarmia and Ecleora appear as sister taxa in multi-gene analysis (Murillo-Ramos et al. 2019). 182 Validated at subspecies rank in Scoble (1999), downgraded to synonymy in Skou et al. (2017). 183 Possibly a synonym of E. atlanticaria, see text in this volume. 184 Described at subspecies rank, downgraded to synonymy in Leraut (2009). 185 See comment under Ekboarmia. 186 subordinated as subspecies under Afriberina tenietaria in Rungs (1981).

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Afriberina Wehrli, 1943 187 A. tenietaria (Staudinger, 1900) 188/���������������������������������������������������������������������������� 6/133 * A. t. hammamrirha Wehrli, 1943 [Algeria] A. salemae Skou & Sihvonen, 2019. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/134 * A. terraria (Bang-Haas, 1907) 189 * A. t. pallescens Rothschild, 1914 [Algeria] * A. haroldi (Oberthür, 1913) [Morocco, Algeria] 190 * A. zernyi (Wehrli, 1943) [Morocco] 191 * A. rungsi (Albers & Warnecke, 1941) [Morocco] 192 Sardocyrnia Wehrli, 1943 193 S. bastelicaria (Bellier, 1862). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/135 S. fortunaria (Vázquez, 1905). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/136 Selidosema Hübner, 1823 194 S. brunnearia (Villers, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/137 (= dilucescens Wehrli, 1924 195; = ­pallidaria Staud ­ inger, 1901 196; = parenzani Haus197 mann, 1993  ) S. b. scandinaviaria Staudinger, 1901 S. b. tyronensis Cockayne, 1948 198 * S. b. syriacaria Staudinger, 1901 [Turkey: Pontus] 199 S. plumaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/138 * S. p. amani Wehrli, 1932 [Turkey] S. pyrenaearia (Boisduval, 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/139 (= oliveirata Mabille, 1876) 200 * S. p. mauretania Wehrli, 1953 [Morocco, Algeria] 201 (?= herbuloti Rungs, 1975) [Morocco] 202 S. taeniolaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/140 (= extensaria Turati, 1919) 187 In multi-gene analysis Afriberina does not have close European genera (Murillo-Ramos et al. 2019), but the taxon sampling is still rather scarce. Position tentative, placed next to Ecleora and Ekboarmia on the basis of superficially similar wing pattern. 188 The nominotypical subspecies has been described from Algeria, it is unclear if the taxon punctata Fernández, 1933 should be validated for the European populations or not. 189 Species status and potential occurrence in Europe unclear, see text (this volume) for details. 190 In Scoble (1999) validated at species rank. Under the misspelled name haroldaria subordinated as subspecies under A. tenietaria in Rungs (1981). 191 In Scoble (1999) validated at species rank, subordinated as subspecies under A. tenietaria in Rungs (1981). 192 In Scoble (1999) validated at species rank, subordinated as subspecies under A. tenietaria in Rungs (1981). 193 In multi-gene analysis Sardocyrnia + Selidosema + Peribatodes cluster together (Murillo-Ramos et al. 2019). 194 See comment under Sardocyrnia. 195 For synonymy see text in this volume. 196 Downgraded to synonymy in Flamigni et al. (2016). 197 DNA analysis revealed that “S. parenzani” is a hybrid between S. brunnearia and S. ambustaria, for details see text in this volume. 198 Subspecies extinct at its own locality (Skinner 2009). 199 Subordinated as subsp. under S. brunnearia in Scoble (1999); possibly a separate species, see text in this volume. 200 For synonymy see text in this volume. 201 Described as subspecies of S. plumaria, until recently retained conspecific with S. brunnearia. Now considered subspecies of S. pyrenaearia, tentatively, for details see text in this volume. 202 For potential synonymy see text in this volume.

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* S. atraria Powell, 1941 [Morocco] S. ambustaria (Geyer, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/141 S. erebaria Oberthür, 1883 203 * S. e. erebaria Oberthür, 1883 [Morocco, north-­western Algeria] S. e. postdentaria (Rothschild, 1914) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/142 * S. picturata Rothschild, 1914 [Morocco, Algeria] * S. tamsi Rebel, 1939 [Cyprus] * S. combustaria Püngeler, 1904 [Israel] 204 Peribatodes Wehrli, 1943 205 P. rhomboidaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/143 (= ichnusaria Ghiliani, 1852; = sublutearia Zerny, 1927; = defloraria Dannehl, 1928; = corsicaria Schawerda, 1931; = syritaurica Wehrli, 1931; = saerdabensis Wehrli, 1941) P. perversaria (Boisduval, 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/144 (= abstersaria Boisduval, 1840; = buxicolaria Mabille, 1873; = subflavaria Millière, 1876; = montserrata Wehrli, 1943; = galliberia Wehrli, 1943) P. omalensis Rajaei, Skou & Hausmann, 2019 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/145 P. umbraria (Hübner, 1809) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/146 (= syrirana Wehrli, 1943 [Turkey, Levant]; = perumbraria Leraut, 2009 206) * P. u. vaucheri (Thierry-Mieg, 1916) 207 [Morocco] (= mimeuri Rungs, 1950; = khenchelae Wehrli, 1943 [Algeria]) P. ebusaria Vallhonrat, 2012. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/147 P. secundaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/148 (= druentiaria Cleu, 1928 208; = occidentaria Lempke, 1952 209) * P. s. fallentaria Staudinger, 1879 [Turkey] 210 P. correptaria (Zeller, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/149 P. aragonis (Wehrli, 1943). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/150 P. a. gerini Leraut, 2009 [Morocco] * P. powelli (Oberthür, 1913) [Morocco, Algeria] P. ilicaria (Geyer, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/151 (= jacobiaria Fernández, 1931) * P. i. maghrebica Le Cerf, 1923 [Morocco] * P. jugurthina (Turati & Krüger, 1936) [Libya] Alcis Curtis, 1826 211 A. repandata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/152 203 Placed in Menophra in Scoble (1999). 204 Recently collected material of this rare species suggesting relationships to the Afrotropical genus Xylopteryx Guenée, 1858 (Hausmann in prep.). 205 See comment under Sardocyrnia. In the multi-gene analysis of Jiang et al. (2017) clustering with Alcis. 206 Colour combination falls within the variation range of the nominate subspecies, confirmed by Flamigni et al. (2016), here regarded as a synonym of the nominate subspecies. 207 Validated at subspecies level by Leraut (2009). Confirmation of this subspecies needs additional research based on larger series. 208 Here regarded as a synonym of the nominate subspecies based on sympatric occurrence. 209 Here regarded as a synonym of the nominate subspecies based on sympatric occurrence. 210 Valid at subspecific rank (Scoble 1999), confirmation of the validity of this subspecies needs further study. 211 In multi-gene analysis Alcis is included in a large assemblage of genera including among others Sardocyrnia, Selidosema and Peribatodes (Murillo-Ramos et al. 2019). In the multi-gene analysis of Jiang et al. (2017) in the same genetic cluster as Peribatodes.

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A. r. sodorensium (Weir, 1881) A. r. muraria Curtis, 1826 A. deversata (Staudinger, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/153 (= bastelbergeri (Hirschke, 1908) * A. subrepandata (Staudinger, 1897) [S. Kazakhstan to Tadjikistan] A. jubata (Thunberg, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/154 * A. caucasica (Wehrli, 1928) [Transcaucasia: Georgia] * A. songarica (Alpheráky, 1883) [S. Kazakhstan to China] Deileptenia Hübner, 1825 212 D. ribeata (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/155 Paraboarmia Krampl, 1994 213 P. viertlii (Bohatsch, 1883). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/156 Adactylotis Hübner, 1823 214 A. gesticularia (Hübner, 1817) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/157 * A. g. ungemachi (D. Lucas, 1933) [Morocco] A. contaminaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/158 Pseudocoremia Butler, 1877 P. suavis Butler, 1879 215/������������������������������������������������������������������������������������������ 6/159 Parectropis Sato, 1980 216 P. similaria (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/160 Aethalura McDunnough, 1920 217 A. punctulata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/161 Ascotis Hübner, 1825 218 A. selenaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/162 (= dianaria Hübner, 1817) * Ascotis fortunata (Blachier, 1887) [Canary Islands: Tenerife] * A. f. flavonigrata (Pinker, 1965) [Canary Islands: La Palma] * A. f. wollastoni (Bethune-Baker, 1891) [Madeira] * A. f. azorica (Pinker, 1971) [Açores]

212 In multi-gene analysis Deileptenia is included in an assemblage of non-European genera, which is sister to a lineage that includes among others Alcis, Sardocyrnia, Selidosema and Peribatodes (Murillo-Ramos et al. 2019). In the multi-gene analysis of Jiang et al. (2017) in the large genetic cluster including among others Alcis and Peribatodes. 213 Tentative position, requiring further research. Phthonandria combination (as in Viidalepp 1996) not supported by structural morphology (our observation) nor by molecules: type species of Phthonandria included in multi-­ gene analysis and grouping in the Gnophini (Murillo-Ramos et al. 2019). 214 Position tentative, based on similarity of male genitalia morphology between Adactylotis, Pseudocoremia and Deileptenia. 215 Not a native European fauna element, introduced from New Zealand to United Kingdom, now established there. Systematic position within Boarmiini tentative, awaiting further study. 216 In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Aethalura. 217 See footnote to previous genus. 218 In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Cleora and Ectropis.

828

Cleora Curtis, 1825 219 C. cinctaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/163 C. c. bowesi Richardson, 1952 Ectropis Hübner, 1825 220 E. crepuscularia (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/164 (= bistortata Goeze, 1781; = laricaria Doubleday, 1847 221) Arichanna Moore, 1868 222 A. melanaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/165 Bupalus Leach, 1815 223 B. piniaria (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/166 B. p. bernieri de Lajonquiére, 1958 B. p. espagnolus Eitschberger & Steiniger, 1975 * Ifrania Leraut, 2009 224 * I. michaellarius Rungs, 1941 [Morocco] Megametopon Alphéraky, 1892 225 (= Contropis Wiltshire, 1967) 226 M. grisolaria (Eversmann, 1848). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/167 * M. modestaria (Püngeler, 1914), comb. n. [C. Asia, Turkmenistan] * M. m. tagana (Wiltshire, 1967), comb. n. [Afghani­stan] * Zeuctoboarmia Prout, 1915 227 * Z. syntropha (Prout, 1931) [Jordan, Israel] 228 * Jordanisca Wehrli, 1943 229 * J. tenuisaria (Staudinger, 1900) [Israel, Jordan, Iraq] Tephronia Hübner, 1825 230 T. sepiaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/168 (= cineraria Denis & Schiffermüller, 1775; = finga­lata Millière, 1873; = teriolica ­Wehrli, 1953) T. nuragica Fiumi et al., 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/169

219 220 221 222

See footnote to previous genus. See footnote to the previous two genera. Not mentioned as subspecies in Skinner (2009). In the multi-gene analyses of Jiang et al. (2017) and Murillo-Ramos et al. (2019) in the same genetic cluster as Bupalus. 223 See footnote to previous genus. Bupalus is listed under Bupalini in Viidalepp (1996), in this volume listed as synonym of Boarmiini, following all recent molecular analyses (e.g. Wahlberg et al. 2010; Sihvonen et al. 2011; Jiang et al. 2017; Murillo-Ramos et al. 2019). 224 In previous literature combined with Bupalus. 225 Systematic position within Boarmiini tentative, awaiting further study. In Viidalepp (1996) placed next to Tephronia + Eumannia. 226 See text of this volume. 227 Systematic position within Boarmiini tentative, awaiting further study. 228 Recently collected in Israel, new for the fauna (Hausmann, in prep.). 229 Systematic position within Boarmiini tentative, awaiting further study. 230 Systematic position within Boarmiini tentative, awaiting further study.

829

* T. bytinskii (Wehrli, 1939) [Canary Islands, Morocco] 231 (= melloulica Leraut, 2009) T. codetaria (Oberthür, 1881) * T. c. codetaria (Oberthür, 1881) [Algeria, Tunisia] T. c. sicula (Wehrli, 1933). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * T. c. gerini Leraut, 2009 [Morocco] T. lhommaria (Cleu, 1928). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. l. melaleucaria Schwingenschuss, 1932 T. l. cleui Wehrli, 1953 (= diniensis Leraut, 2009) T. cyrnea (Schawerda, 1932) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= tonnara Tautel, 2015) T. lepraria Rebel, 1909. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. psyloritaria (Reisser, 1958). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. theophilaria Hausmann, 2019 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. espaniola (Schawerda, 1931) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * T. minutaria Turati, 1934 [Libya, Egypt] * T. destituta (Walker, 1870) 232 [Egypt] * T. duercki Reisser, 1933 [Morocco] * T. fatimaria Bang-Haas, 1906 [Morocco, Algeria] (= ismailaria Oberthür, 1913) * T. praerecta Wehrli, 1933 [Algeria, Tunisia] 233 * T. nigrolineata Zerny, 1934 [Morocco] 234

6/170 6/171

6/172 6/173 6/174 6/175 6/176

Eumannia Fletcher, 1979 235 E. oppositaria (Mann, 1864). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/177 * E. o. obscuraria (Staudinger, 1901) [Iran, Armenia, Turkey] E. oranaria (Staudinger, 1892) * E. o. oranaria (Staudinger, 1892) [Morocco to Tuni­sia] E. o. castiliaria Staudinger, 1892 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/178 E. o. cebennaria Chrétien, 1898 (= hubertcleui Leraut, 2009) E. o. alpinaria Leraut, 2009 * E. arenbergeri Hausmann, 1995 [Cyprus] * E. neoppositaria Beljaev & Vasilenko, 1998 [Turkmenistan] * E. fumosata Hausmann, 1995 [Jordan, Israel] * E. syriaca (Turati & Krüger, 1936) [Libya] Phyllometra Boisduval, 1840 236 P. culminaria (Eversmann, 1843). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/179 P. gracilaria Boisduval, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/180 * P. teneraria (Staudinger, 1892) [central Asia] 231 In Scoble (1999) combined with Eumannia, see remarks under Tephronia, this volume. 232 Described as “Tephrina”, combined with Tephronia in Scoble 1999; unclear taxon, possibly a senior synonym of T. minutaria? 233 For validity of this taxon at species rank see this volume. 234 For validity of this taxon at species rank see this volume. 235 Systematic position within Boarmiini tentative, awaiting further study. 236 Systematic position within Boarmiini tentative, awaiting further study.

830

* P. argentaria (Bang-Haas, 1910) [Tunisia] (= planaria Chrétien, 1911) 237 * P. proutiana Turati, 1936 [Libya] * Habermania Viidalepp, 1988 238 * H. oxygonaria (Püngeler, 1900) [central Asia] Genera of uncertain tribus association (see remarks under each genus in GME5) Ithysia Hübner, 1825 239 I. pravata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6/181 Lignyoptera Lederer, 1853 L. fumidaria (Hübner, 1825). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/57 L. thaumastaria Rebel, 1901. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/58 Opisthograptis Hübner, 1823 O. luteolata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/64 Toulgoetia Herbulot, 1946 T. cauteriata (Staudinger, 1859). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/72 Enanthyperythra Wehrli, 1937 E. legataria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/74 Eumera Staudinger, 1892 E. regina Staudinger, 1892. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/82 * E. hoeferi Wehrli, 1934 [Turkey] * E. h. transcaucasica Wehrli, 1940 [Transcaucasus, Armenia] * E. mulier Prout, 1929 [Cyprus] * E. turcosyrica Wehrli, 1932 [Turkey, Eastern Mediterranean] * Apocolotois Wehrli, 1936 * A. smirnovi (Romanoff, 1885) [Caucasus, Armenia] Asovia Alphéraky, 1908 A. maeoticaria (Alphéraky, 1876). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/97 Lhommeia Wehrli, 1939 L. biskraria (Oberthür, 1885). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/103 * L. b. bleusei (Thierry-Mieg, 1905) [Algeria] * L. b. cinnamomaria (Rothschild, 1914) [Algeria] * L. b. illiturata (Warren, 1897) [South-East Africa] * L. b. viridaria (Rothschild, 1914) [Algeria] Onychora Meyrick, 1892 O. agaritharia (Dardoin, 1842). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5/138 * O. a. atlasica Leraut, 2009 [Morocco] 237 In Scoble (1999) still validated at species rank. 238 Systematic position within Boarmiini tentative, awaiting further study. 239 Systematic position near the so far unassigned Lignyoptera probable, supported by male and female genitalia (P. Sihvonen). In the multi-gene analysis of Murillo-Ramos et al. (2019) clustering with the “Apeirini + Hypochrosini + Epionini” complex of tribes, as a separate lineage.

831

O. a. hepaticaria (Schwingenschuss, 1930) * O. a. mauretanicaria (Staudinger, 1901) [Mauretania?, Algeria] * O. a. rungsi Leraut, 2009 [Morocco] * Boarmioides * B. colpias (D. Lucas, 1932) [Morocco] 240 * Xenobiston Warren & Rothschild, 1903 241 * X. casta Warren & Rothschild, 1903 [Egypt, Levant]

Subfamily Sterrhinae Meyrick, 1892 Tribus Sterrhini Meyrick, 1892 Anthometra Boisduval, 1840 A. plumularia Boisduval, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/1 Emmiltis Hübner, 1825 E. pygmaearia (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/2 Cleta Duponchel, 1845 C. ramosaria (Villers, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/3 * C. transiens Prout, 1935 [Algeria, Tunisia] C. perpusillaria (Eversmann, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/4 * C. cinneretharia Amsel, 1935 [Levant] C. filacearia (Herrich-Schäffer, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/5 Idaea Treitschke, 1825 The aureolaria species-group I. serpentata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/6 I. luteolaria (Constant, 1863). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/7 I. aureolaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/8 I. flaveolaria (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/9 I. muricata (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/10 The rufaria species-group I. determinata (Staudinger, 1876). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/11 I. litigiosaria (Boisduval, 1840) 242/�������������������������������������������������������������������������������� 2/12 I. mutilata (Staudinger, 1876). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/13 * I. hausmanni (Trusch & Müller, 2001) [Tunisia, Libya] I. lusohispanica Herbulot, 1991. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/14 * I. lambessata (Oberthür, 1887) [Morocco to Tunisia] I. sardoniata (Homberg, 1912) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/15 I. korbi (Püngeler, 1916) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/16 240 In Rungs (1981) combined with Semiothisa. 241 Habitus suggesting position near Dyscia (Gnophini). 242 Leraut (2009) mentions Idaea agrestaria (de Joannis, 1891) (sic! = misspelling of agraria) and upgrades it from synonymy of Idaea litigiosaria (cf. GME2) to species rank vaguely stating differences in male genitalia (“genitalia are different”), but without describing these differences and without examining the possibility of clinous distribution of characters. Since these taxa still await a thorough differential analysis, agrestaria is herewith downgraded again to synonymy.

832

I. mediaria (Hübner, 1819). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. m. flavidior (Rothschild, 1914) [Algeria] I. leipnitzi Hausmann, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. bengasiaria (Turati, 1926) [Libya] * I. renataria (Oberthür, 1878) [Morocco to Libya] I. rufaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. consanguiberica Rezbanyai-Reser & Expósito, 1992. . . . . . . . . . . . . . . . . . . . . . . . . . * I. debiliata (Sterneck, 1934) [Morocco] I. admiranda Hausmann, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. consanguinaria (Lederer, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. c. turatii (Sohn-Rethel, 1929) * I. c. consecrata (Staudinger, 1897) [Levant] I. ossiculata (Lederer, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sericeata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. s. calvaria Wehrli, 1927 (= subrecta Prout, 1935) 243 * I. allardiata (Mabille, 1869) [Algeria to Libya] I. macilentaria (Herrich-Schäffer, 1847) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= antiquaria Herrich-Schäffer, 1847) * I. numidaria (Lucas, 1849) [Algeria] I. ochrata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. o. albida (Zerny, 1936) The rusticata species-group I. nevadata (Wehrli, 1934). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. figuraria (Bang-Haas, 1907). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= subcompleta Fernández, 1931) I. alicantaria (Reisser, 1963). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. oberthueri (Lucas, 1917) [Tunisia] I. nigrolineata (Chrétien, 1910). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. completa (Staudinger, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. c. apostolica Hausmann, 1994 [Cyprus] * I. erlacheri Hausmann, 2003 [Tunisia] I. intermedia (Staudinger, 1879). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. rusticata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= vulpinaria Herrich-Schäffer, 1852) I. mustelata (Gumppenberg, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. filicata (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= levequei Leraut, 2005) 244 I. troglodytaria (Heydenreich, 1851). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. laevigata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/17 2/18 2/19 2/20 2/21 2/22 2/23 2/24

2/25 2/26

2/27 2/28 2/29 2/30 2/31 2/32 2/33 2/34 2/35 2/36 2/37

243 In Leraut (2009) upgraded from synonymy of Idaea sericeata calvaria to species rank vaguely stating that “genitalia are different”, but without describing these differences. Herewith downgraded again to the above mentioned synonymy based on the analysis presented in Hausmann (2004). 244 For Corsica, Leraut (2009) reports both the occurrences of subsp. levequei Leraut, 2005 and nominate subspecies with the remark that subsp. levequei “could be a separate species”. Own research on such individual forms with forewing tip pointed and postmedial line clearly visible do not reveal any constant difference in genitalia and neither show genetic divergences (COI barcode fragment, n=25) from the other forms. Therefore downgraded to synonymy.

833

(= couloniata Balestre, 1907) 245 * I. vilaflorensis (Rebel, 1911) [Canary Islands] * I. charitata (Rebel, 1914) [Canary Islands] (= curvata Weisert, 2005) 246 * I. nigra Hausmann & Blaesius, 2007 [Canary Islands] I. efflorata Zeller, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/38 (= extarsaria Herrich-Schäffer, 1852) I. attenuaria (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/39 I. incalcarata (Chrétien, 1913) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/40 The typicata species-group I. typicata (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/41 I. t. hornigaria (Staudinger, 1901) * I. dyraria (Zerny, 1934) [Morocco] I. alyssumata (Himmighoffen & Millière, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/42 I. a. reynaldiata (Rouast, 1876) I. a. selenensis Mazel & Varenne, 2017 247 I. barbuti Tautel & Lévêque, 2013 248/������������������������������������������������������������������������ 6/185 I. urcitana (Agenjo, 1952). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/43 I. moniliata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/44 * I. ludovicaria (Culot, 1918) [Morocco to Tunisia] * I. unicalcarata (Prout, 1922) [Morocco, Algeria] The spissilimbaria species-group I. spissilimbaria (Mabille, 1888) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/45 * I. capnaria (Püngeler, 1909) [Levant] * The microptera species-group * I. microptera (Warren & Rothschild, 1905) [Egypt, Levant] * I. sordida (Rothschild, 1913) [Morocco to Libya] * I. s. nili (Wiltshire, 1949) [Egypt, Levant] * I. sordidior (Wiltshire, 1949) [Egypt] * I. granulosa (Warren & Rothschild, 1905) [Egypt, Levant] * I. miserrima (Turati, 1930) [Libya] * I. scabraria (Chrétien, 1909) [Algeria, Tunisia] * I. sanctaria (Staudinger, 1900) [Levant] * I. s. outayana (Wehrli, 1931) [Morocco to Tunisia] (= bucephalaria Chrétien, 1909) * I. s. transcatenulata (Rothschild, 1915) [Algeria] * I. ortali Hausmann, 1991 [Levant] * I. romeii (Turati, 1930) [Libya] * The volloni species-group * I. volloni (Lucas & Joannis, 1907) [Canaries to Levant] 245 In Leraut (2009) erroneously listed as synonym of I. belemiata. 246 Described from Canary island Hierro without differential diagnosis from I. charitata. In genitalia well matching I. charitata. 247 Recently described based on material from southern France (prov. Gard to Pyrenees). The classification of the Iberian populations (currently under the nominotypical subspecies) remains unresolved. 248 New description for Europe (Portugal), see information in this volume.

834

The circuitaria species-group I. circuitaria (Hübner, 1819). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/46 I. c. expositoi Hausmann, 1994 I. rainerii Hausmann, 1994 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/47 * I. mimosaria (Guenée, 1858) [Turkey, Levant] * I. m. wimmeri Hausmann, 1994 [Cyprus] The incisaria species-group I. incisaria (Staudinger, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/49 I. i. albarracina (Reisser, 1933) 249/������������������������������������������������������������������������ 2/48 * I. i. praecisa (Reisser, 1933) [Morocco] * I. i. pulverulenta (Reisser, 1933) [Morocco to Libya] I. textaria (Lederer, 1861) 250/������������������������������������������������������������������������� 6/186 (2/49l) I. calunetaria (Staudinger, 1859). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/50 (= dorycniata Bellier, 1862) I. c. valesiaria (Püngeler, 1888) * I. c. episticta (Wehrli, 1930) [Morocco, Algeria] * I. mareotica (Draudt, 1912) [Morocco to Levant] (= fuscularia Turati, 1930) * I. allongata (Staudinger, 1897) [Levant] * I. a. sublongaria (Staudinger, 1899) [Levant] * I. pectinata (Sterneck, 1933) [Levant] The obsoletaria species-group * I. hathor (Wiltshire, 1949) [Egypt, Levant] I. belemiata (Millière, 1868). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= substraminata Prout, 1913) I. b. helianthemata (Millière, 1870) (= euphorbiata Balestre, 1906) (= oberthuriata Balestre, 1907) (= cavenacata Chrétien, 1929) I. elongaria (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. e. pecharia (Staudinger, 1863) * I. maxima (Wehrli, 1934) [Morocco] * I. palmata (Staudinger, 1901) [Canary Islands] * I. substriata (Turati, 1930) [Libya] * I. peluraria (Reisser, 1939) [Turkey, Cyprus] * I. persidis (Wiltshire, 1966) [Turkey] I. palaestinensis (Sterneck, 1933). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. obsoletaria (Rambur, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. o. lilaceola Hausmann, 2003 I. o. dierli Hausmann, 1991 I. o. dionigii Hausmann, 1991 I. o. rufularia (Herrich-Schäffer, 1847) * I. o. algeriensis (Bethune-Baker, 1888) [Algeria] * I. epaphrodita (Wehrli, 1934) [Levant]

2/51

2/52

2/53 2/54

249 For downgrading to subspecies rank, see information in this volume. 250 New records for Europe (Samos), see information in this volume.

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* I. e. eremita (Wiltshire, 1949) [Egypt] * I. wittmeri (Wiltshire, 1982) * I. w. mienisi Hausmann, 1991 [Levant] I. obliquaria (Turati, 1913). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/55 The inquinata species-group I. inquinata (Scopoli, 1763) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= herbariata Fabricius, 1798) * I. i. adherbariata (Staudinger, 1898) [Levant] * I. saida (Wiltshire, 1968) [Levant] * I. holliata (Homberg, 1909) [Levant] 251 * I. banghaasi (Prout, 1934) [Levant] * I. affinitata (Bang-Haas, 1907) [Levant] I. blaesii Lenz & Hausmann, 1992. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. dilutaria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. fuscovenosa (Goeze, 1781) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. f. corsula (Schawerda, 1929) I. robiginata (Staudinger, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. lutulentaria (Staudinger, 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. terminolineata (Rothschild, 1933) [Morocco, A ­ lgeria] I. humiliata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. h. davidi Gaston & Redondo, 2005 252 I. bigladiata Herbulot, 1975. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. b. alberti Hausmann, 2004 [Algeria, Tunisia] I. politaria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. roseofasciata (Christoph, 1882) [Turkey, Trans­caucasus] * I. maronitaria (Zerny, 1933) [Levant]

2/56

2/57 2/58 2/59 2/60 2/61 2/62 2/63 2/64

The longaria species-group I. longaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/65 * I. abnorma (Pinker, 1960) [Canary Islands] * I. atlantica (Stainton, 1859) [Madeira] (= I. illuminata Prout, 1939) * I. margaritacea (Turati & Krüger, 1936) [Libya] The nexata species-group I. nexata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/66 The consolidata species-group I. descitaria (Christoph, 1893) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. vesubiata (Millière, 1873). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. libycata (Bartel, 1906) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. consolidata (Lederer, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. joannisiata (Homberg, 1911) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/68 2/69 2/70 2/71 2/72

251 In Leraut (2009) downgraded from species rank to subsp. of Idaea inquinata without presenting facts (e.g. figures of the examined material), based on the erroneous opinion to have examined “the type” and ignoring the comprehensive revision of the species-group (Hausmann 2005) with lectotype designation. Taxonomic change not accepted. 252 New description for Europe (Spain). For downgrading to subspecies rank, see information in this volume.

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I. j. ibericata (Wehrli, 1927) 253 I. acutipennis Hausmann & Honey, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/73 * I. jacobi Hausmann, 1991 [Levant] * I. striolata (Staudinger, 1892) [Levant] * I. detritaria (Staudinger, 1897) [Levant] The seriata species-group I. seriata (Schrank, 1802). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= virgularia sensu Hübner, 1799) I. s. canteneraria (Boisduval, 1840) I. minuscularia (Ribbe, 1912). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= herbuloti Agenjo, 1952) I. ibizaria Mentzer, 1980. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. albitorquata (Püngeler, 1909). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. a. madoniensis Hausmann, 1993 * I. a. maculifera Hausmann, 1994 [Cyprus] I. carvalhoi Herbulot, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. fathmaria (Oberthür, 1876) [Morocco, Algeria] * I. muelleri Hausmann, 2003 [Caucasus] * I. maderae (Bethune-Baker, 1891) [Madeira] (= zargi Bethune-Baker, 1891)

2/74 2/75 2/76 2/77 2/78

The camparia species-group I. camparia (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/79 I. c. sodaliaria (Herrich-Schäffer, 1852) The subsericeata species-group I. subsericeata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= zernyi Lucas, 1934) * I. maurusia (Turati, 1924) [Libya] I. pallidata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. mancipiata (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= bustilloi Agenjo, 1967) I. m. repagulata (Prout, 1934) * I. proclivata (Fuchs, 1902) [Turkey, Transcaucasus] (= osthelderi Wehrli, 1932) I. sylvestraria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. dimidiata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. d. antitaurica (Wehrli, 1931) [Turkey, Levant] I. subsaturata (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. s. lecerfiata (Homberg, 1910) [Morocco to Libya; Levant?; Cyprus?] I. trigeminata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * I. t. tenuirussata (Zerny, 1933) [Cyprus, Levant] I. biselata (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/80 2/81 2/82

2/83 2/84 2/85 2/86 2/87

The fractilineata species-group I. fractilineata (Zeller, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/88 253 In Leraut (2009) as Idaea joannisiata f. ibericata suggesting downgrading from subspecies of joannisiata to synonymy. To be maintained at subspecies rank (cf Hausmann 2004).

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I. f. subrufaria (Staudinger, 1899) * I. inclinata (Lederer, 1855) [Libya to Turkey, Cyprus] * I. merklaria (Oberthür, 1885) [Algeria] The cervantaria species-group I. cervantaria (Millière, 1869). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/90 * I. c. carneotincta (Zerny, 1936) [Morocco] (= okbaria Chrétien, 1911) [Algeria?, Tunisia] 254 I. contiguaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/91 I. c. britanniae (Müller, 1936) I. rupicolaria (Reisser, 1927). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/92 The deitanaria species-group I. deitanaria (Reisser & Weisert, 1977). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/93 The infirmaria species-group I. saleri Domínguez & Baixeras, 1992. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/94 I. tineata (Thierry-Mieg, 1911) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/95 (= antennata Wehrli, 1931) * I. t. praetineata Hausmann, 1994 [Levant] I. infirmaria (Rambur, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/96 * I. i. mitescens Scoble, 1999 [Morocco to Tunisia] I. rhodogrammaria (Püngeler, 1913) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/97 I. lobaria (Chrétien, 1909) 255/������������������������������������������������������������������������������������ 6/187 (= balestraria Lucas, 1909) * I. bacalladoi (Pinker, 1974) [Canary Islands] * I. alutaceata (Rungs, 1945) [Morocco] * I. purpurariata (Pinker, 1974) [Canary Islands] * I. fuerteventurensis (Pinker, 1974) [Canary Islands] *The gracilipennis species-group * I. gracilipennis (Warren, 1901) [Turkey, Levant] * I. purpureomarginata (Bohatsch, 1880) [Turkey, Levant] * I. p. stefani Hausmann, 1991 [Levant] * I. subpurpurata (Staudinger, 1900) [Libya, Levant, Turkey] The ostrinaria species-group I. ostrinaria (Hübner, 1813) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/98 * I. o. demarginata (Schwingenschuss, 1938) [Turkey] I. metohiensis (Rebel, 1900). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/99 * I. m. anatolica (Schwingenschuss, 1938) [Turkey] The eugeniata species-group I. eugeniata (Millière, 1870). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/100 * I. e. algeriaca (Culot, 1917) [Morocco to Tunisia] I. distinctaria (Boisduval 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/101 (= ruficostata Zeller, 1849) 254 In Leraut (2009) downgraded from species rank to subsp. of Idaea cervantaria with statement to have “examined the type”, referring, however, to a paralectotype of MNHN, Paris. Examination of the lectotype in coll. Herbulot/ZSM confirms the proposed synonymy. 255 New records for Europe (Spain), see information in this volume.

838

I. predotaria (Hartig, 1951) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/102 (= griseanova Rezbanyai-Reser, 1987) The nitidata species-group I. nitidata (Herrich-Schäffer, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/103 The aversata species-group I. emarginata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/104 I. dromikos Hausmann, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/105 I. simplicior (Prout, 1934) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/106 * I. omari Hausmann & Bläsius, 2008 [Morocco] I. rubraria (Staudinger, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/107 (= bilinearia Fuchs, 1889) I. aversata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/108 * I. a. indeviata (Prout, 1929) [Algeria, Tunisia] I. gelbrechti Hausmann, 2003 256/�������������������������������������������������������������������������������� 6/188 I. degeneraria (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/109 I. d. erschoffi (Christoph, 1872) I. straminata (Borkhausen, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/110 (= inornata Haworth, 1809) I. deversaria (Herrich-Schäffer, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/111 (?= maritimaria Bruand, 1846) I. d. fallax Hausmann, 2003 I. squalidaria (Staudinger, 1882). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/112 * I. neglecta Hausmann & Werno, 2003 [Canary ­Islands] incertae sedis * I. protrusa (Turati, 1926) [Libya] * I. bicertaria (Staudinger, 1892) [Tunisia] Brachyglossina Wagner, 1914 B. hispanaria (Püngeler, 1913). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/113 B. exilaria (Guenée, 1858) 257/�������������������������������������������������������������������������������������� 2/89 * B. erubescens (Hausmann, 2003) [Algeria, Tunisia] 258 * B. truschi (Hausmann, 2003) [Morocco] 259 B. manicaria (Herrich-Schäffer, 1852) 260/������������������������������������������������������������������ 2/67 (= fatimata Staudinger, 1895) * B. brevitarsata (Hausmann, 1991) [Levant] * B. consociata (Staudinger, 1900) [Turkey, Levant] * B. improbata (Staudinger, 1898) [Levant] * B. hassan Herbulot, 1974 [Algeria] * B. tantalidis (Turati, 1924) [Algeria?, Tunisia, Libya] * B. mauritanica (Bethune-Baker, 1885) [Algeria] 256 New records for Europe (Spain), see information in this volume. 257 For new combination with genus Brachyglossina, see information in this volume. 258 Tunisian sister species of B. exilaria, for new combination with genus Brachyglossina, see information in this volume. 259 Moroccan sister species of B. exilaria, for new combination with genus Brachyglossina, see information in this volume. 260 For new combination with genus Brachyglossina, see information in this volume.

839

* B. mzabensis Prout, 1938 [Algeria] * B. ochrolutearia (Turati, 1934) [Libya] * B. acidalaria Wagner, 1914 [Tunisia] * B. macracantha Prout, 1938 [Algeria] * B. maroccana (Wehrli, 1930) [Morocco] * B. paroranaria Wehrli, 1938 [Algeria] * B. p. suboranaria Wehrli, 1938 [Algeria] * B. seitzi Prout, 1938 [Algeria] * B. vindicata Prout, 1938 [Algeria] * B. fulta Prout, 1938 [Algeria] * B. culoti (Wehrli, 1930) [Morocco, Algeria] (= pseudoranaria Zerny, 1936) * B. oranaria (Bang-Haas, 1907) [Algeria, Morocco] * B. staudingeri (Prout, 1932) [Levant] * B. sonyae (Wiltshire, 1990) [Levant] * B. williamsi Wiltshire, 1949 [Egypt, Levant] * Tineigidia Sterneck, 1934 * T. eremica Amsel & Sterneck, 1934 [Levant] * T. aegyptiaca Wiltshire, 1949 [Egypt] Limeria Staudinger, 1892 L. macraria Staudinger, 1892. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/114 Tribus Scopulini Duponchel, 1845 Oar Prout, 1913 261 O. reaumuraria (Millière, 1864). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/115 O. pratana (Fabricius, 1794) 262 /���������������������������������������������������������������6/189 (2/115d-h) * O. p. oppressa (Walker, 1870) [Levant, Egypt, Arabia] * O. p. baezi Hausmann, 2004 [Canary Islands, Morocco] Cinglis Guenée, 1858 263 C. humifusaria (Eversmann, 1837) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/116 C. andalusiaria Wagner, 1935 264/������������������������������������������������������������������������������ 2/117 Scopula Schrank, 1802 Subgenus Scopula Schrank, 1802 265 The immorata species-group S. immorata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/118 S. i. duercki Sheljuzhko, 1955 261 Genus Oar subordinated under Scopula in Sihvonen (2005), subsequently again treated at genus level (e.g. Fiumi & Guidi 2011; Hausmann 2006a; 2009; Hausmann et al. 2011; 2013; Redondo et al. 2009; 2015). 262 New records for Europe (Lampedusa), see information in this volume. 263 Genus Cinglis subordinated under Scopula in Sihvonen (2005), subsequently again treated at genus level (e.g. Leraut 2009; Redondo et al. 2009; Hausmann et al. 2011; 2013). 264 In Redondo et al. (2015) as ‘Scopula reducta Thierry-Mieg, 1915‘, an unused epithet of doubtful nomen­ clatural availability (thus not listed in Scoble 1999). 265 Genus Scopula here subdivided into three subgenera (following GME2) which will probably not stand a ­thorough revision of this megadiverse genus at global level (cf. Sihvonen 2005).

840

S. i. riloensis (Züllich, 1936) S. tessellaria (Boisduval, 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= tabianaria Turati, 1905) S. t. guillaumei Leraut, 2005 266 S. corrivalaria (Kretschmar, 1862). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. caricaria (Reutti, 1853) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= virginalis Fourcroy, 1785) S. nemoraria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. umbelaria (Hübner, 1813) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigropunctata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * S. nepheloperas (Prout, 1916) [Egypt, Sinai]

2/119 2/120 2/121 2/122 2/123 2/124

The virgulata species-group S. virgulata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/125 S. v. rossica (Djakonov, 1926) The ornata species-group S. ornata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. concinnaria (Duponchel, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. c. universaria (Zerny, 1927) S. orientalis (Alphéraky, 1876). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. decorata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. d. violata (Thunberg, 1784) S. d. congruata (Zeller, 1847) S. drenowskii Sterneck, 1941. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. honestata (Mabille, 1869) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. subtilata (Christoph, 1867). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. submutata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. s. roseonitens (Wagner, 1926) S. s. gedrensis Hausmann, 2003 S. s. nivellearia (Oberthür, 1922) * S. s. taurilibanotica (Wehrli, 1932) [Turkey, Cyprus, Levant] * S. transcaspica Prout, 1935 [Turkey, Transcaucasus] S. vigilata (Wagner, 1926). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/133

The rubiginata species-group S. rubiginata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. ochraceata (Staudinger, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. turbidaria (Hübner, 1819) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. turbulentaria (Staudinger, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/134 2/135 2/136 2/137

2/126 2/127 2/128 2/129 6/190 2/130 2/131 2/132

Subgenus Calothysanis Hübner, 1823 The incanata species-group S. decolor (Staudinger, 1897). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/138 * S. d. flaccata (Staudinger, 1897) [Levant] S. imitaria (Hübner, 1799) 267/������������������������������������������������������������������������������������ 2/145 266 Validity of subspecies awaiting confirmation (type locality of nominotypical subspecies may be quite close) and delimitation, including ample material and DNA barcodes. 267 Systematic position beside S. decolor suggested by shared DNA barcodes, see Hausmann et al. (2013).

841

S. i. syriacaria (Culot, 1918) * S. gastonaria (Oberthür, 1876) [Algeria268] * S. g. luteofasciata (Rothschild, 1913) [Algeria] S. rubellata (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. scalercii Hausmann, 2003. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. beckeraria (Lederer, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. b. rebeli (Prout, 1913) * S. b. hermonicola Hausmann, 1997 [Levant] S. incanata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. i. ibericata (Reisser, 1935) S. marginepunctata (Goeze, 1781). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= pastoraria Joannis, 1891) 269 S. luridata (Zeller, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= fulminataria Turati, 1927) * S. bimaculata (Turati & Krüger, 1936) [Libya] (?= barcearia Turati & Krüger, 1936) S. immutata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. frigidaria (Möschler, 1860) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/139 2/140 2/141 2/142 2/143 2/144

2/146 2/147

The ternata species-group S. ternata Schrank, 1802. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/148 S. floslactata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/149 S. f. scotica (Cockayne, 1951) S. subpunctaria (Herrich-Schäffer, 1847) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/150 S. flaccidaria (Zeller, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/151 S. emutaria (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/152 S. minorata (Boisduval, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/153 (= ochroleucaria Herrich-Schäffer, 1847) (= inustaria Herrich-Schäffer, 1847) (= medioumbraria Turati, 1930) * S. andresi (Draudt, 1912) [Egypt] * S. adelpharia (Püngeler, 1894) [Levant] * S. a. pharaonis Sterneck, 1933 [Egypt] S. donovani (Distant, 1892) 270 /�������������������������������������������������������������������������������� 6/191 * S. irrorata (Bethune-Baker, 1891) [Madeira] * S. guancharia (Alphéraky, 1889) [Canary Islands] * S. g. illustris Pinker, 1968 [Canary Islands] * S. g. mus Pinker, 1968 [Canary Islands] * S. g. uniformis Pinker, 1968 [Canary Islands] * S. g. punctabilineatella (Lucas, 1937) [Morocco] 271 268 Position near S. decolor suggested by P. Sihvonen (own observation), based on male genitalia structure. 269 In Leraut (2009) validated as subspecies of S. incanata. The type has been “examined” (dissected?). The figure of the “type” (holotype?) shows a specimen which is more reminiscent of S. marginepunctata than of S. inca­ nata. Until confirmation by genitalia dissection the taxon should be kept in synonymy of S. marginepunctata. 270 Recorded as new for Europe, see information in this volume. 271 In Leraut (2009) raised to species rank without any comment. Species rank not excluded but requiring detailed study combining morphological and molecular data, including the eastern Canary Islands populations of S. guancharia and considering the fact that in the genus Scopula the length of cerata underlies a complicated pattern of variation and polymorphisms.

842

The immistaria species-group S. cajanderi (Herz, 1903). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. divisaria (Christoph, 1893). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= latelineata sensu Staudinger, 1901) S. arenosaria (Staudinger, 1879) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. albiceraria (Herrich-Schäffer, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * S. lehmanni Hausmann, 1991 [Levant] S. immistaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. i. beshkovi Gelbrecht & Hausmann, 1997

2/154 2/155 2/156 2/157 2/158

Subgenus Glossotrophia Prout, 1913 272 The confinaria species-group S. confinaria (Herrich-Schäffer, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. c. prouti (Hausmann, 1993) S. c. aetnaea (Prout, 1935) S. c. scoblei (Hausmann, 1993) * S. diffinaria (Prout, 1913) [Turkey, Transcaucasus] * S. uberaria (Zerny, 1933) [Cyprus, Levant] S. alba (Hausmann, 1993). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. a. milleri (Hausmann, 2004) S. a. brunellii (Hausmann, 1993) * S. a. africana (Hausmann, 1993) [Tunisia] S. mentzeri (Hausmann, 1993) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rufomixtaria (Graslin, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= annae Mentzer, 1990) * S. r. sahariensis (Hausmann, 1993) [Algeria, Tunisia] * S. chalcographata (Brandt, 1938) * S. c. sinaica (Rebel, 1948) [Levant]

2/159

2/160

2/161 2/162

The sacraria species-group 273 S. sacraria (Bang-Haas, 1910) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/163 * S. s. semitata (Prout, 1913) [Levant] * S. s. fumata (Hausmann, 1993) [Cyprus] * S. s. taurica (Wehrli, 1930) [Turkey, Transcaucasus] The asellaria species-group 274 S. asellaria (Herrich-Schäffer, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/164 S. a. romanaria (Millière, 1869) S. a. isabellaria (Millière, 1868) S. a. dentatolineata (Wehrli, 1926) * S. a. gerstbergeri (Hausmann, 1993) [Canary Islands] * S. a. lenzi (Hausmann, 1993) [Morocco] * S. a. philipparia (Prout, 1913) [Algeria, Tunisia] * S. a. tripolitana (Turati, 1929) [Algeria to Libya]

272 Validated at genus rank in Hausmann (2004), subordinated under Scopula in Sihvonen (2005). 273 In Hausmann (2004) “subgenus Libanonia Hausmann, 1993”, requires downgrading to synonymy. 274 In Hausmann (2004) “subgenus Parenzanella Hausmann, 1993”, requires downgrading to synonymy.

843

The romanarioides species-group 275 * S. romanarioides (Rothschild, 1913) [Algeria] * S. alfierii (Wiltshire, 1949) [Egypt, Levant] Scopula incertae sedis * S. moralesi (Rungs, 1945) [Morocco] 276 Holarctias Prout, 1913 277 H. rufinaria (Staudinger, 1861). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/165 Problepsis Lederer, 1853 P. ocellata (Frivaldszky, 1845). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/166 Tribus Rhodostrophiini Prout, 1935 Rhodostrophia Hübner, 1823 The jacularia species-group R. jacularia (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/167 The vibicaria species-group R. vibicaria (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/168 R. v. strigata (Staudinger, 1871) The calabra species-group * R. auctata (Staudinger, 1879) [Turkey, Transcaucasus] R. pudorata (Fabricius, 1794). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. p. perezaria (Oberthür, 1875) R. p. sicanaria (Zeller, 1852) * R. sieversi (Christoph, 1882) R. cretacaria Rebel, 1916. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. calabra (Petagna, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . R. c. separata Prout, 1935 * R. c. transcaucasica Prout, 1920 [Caucasus, Trans­caucasus] R. tabidaria (Zeller, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * R. t. cypriaria Rebel, 1916 [Cyprus] R. discopunctata Amsel, 1935. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= tabidaria sensu Guenée, 1858)

2/169

2/170 2/171 2/172 2/173

The terrestraria species-group * R. terrestraria (Lederer, 1869) [European Russia?; Turkey, Transcaucasus] * R. t. dispar (Staudinger, 1892) [Central Asia] 278 The badiaria species-group R. badiaria (Freyer, 1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/174 * R. iranica Schwingenschuss, 1939 [Transcaucasus] * R. bahara Brandt, 1938 [Turkey, Levant] * R. vastaria (Christoph, 1876) [W. Kazakhstan] 275 In Hausmann (2004) “subgenus Microglossotrophia Hausmann, 1993”, requires downgrading to synonymy. 276 Previously combined with Pseudocinglis Hausmann, 1994, the latter synonymized with Scopula in Sihvonen (2005). 277 Synonymy with Scopula suggested by Sihvonen (2005), validated at genus rank by Beljaev (2016). 278 In Leraut (2009) downgraded from species rank to subspecies of R. terrestraria. Preliminarily acceptable, but awaiting type studies and detailed discussion.

844

The cuprinaria species-group * R. cuprinaria (Christoph, 1876) [Transcaucasus; Turkey] * R. peripheres (Prout, 1938) [Transcaucasus] Tribus Timandrini Stephens, 1850 Timandra Duponchel, 1829 T. comae Schmidt, 1931. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/175 (= amata sensu Staudinger, 1871) * T. c. palmonii Hausmann, 1997 [Levant] T. griseata Petersen, 1902. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/176 T. rectistrigaria (Eversmann, 1851). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/177 Tribus Cosymbiini Prout, 1911 Cyclophora Hübner, 1822 The pendularia species-group C. pendularia (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. albipunctata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. albiocellaria (Hübner, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. lennigiaria (Fuchs, 1883). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. l. mauretanica Reisser, 1933 [Morocco] C. ariadne (Reisser, 1939) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. serveti Redondo & Gaston, 1999. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. annularia (Fabricius, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. puppillaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. p. granti (Prout, 1935) [Açores] * C. p. lilacinipes (Schaus & Cockerell, 1923) [­Madeira] * C. maderensis (Bethune-Baker, 1891) [Madeira] (= lundbladi Bryk, 1940) * C. azorensis (Prout, 1920) [Azores] C. quercimontaria (Bastelberger, 1897). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. ruficiliaria (Herrich-Schäffer, 1855) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

2/178 2/179 2/180 2/181 2/182 2/183 2/184 2/185

2/186 2/187

The porata species-group C. hyponoea (Prout, 1935) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/188 C. porata (Linnaeus, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/189 The punctaria species-group C. suppunctaria (Zeller, 1847). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/190 C. punctaria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/191 C. p. fritzae Hausmann, 2003 C. linearia (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/192 * Pseudosterrha Warren, 1888 * P. rufistrigata (Hampson, 1896) [Egypt, Levant] * P. paulula (Swinhoe, 1887) [Egypt, Levant] * Chlorerythra Warren, 1895 * C. rubriplaga Warren, 1895 * C. r. sinaica Wiltshire, 1949 [Egypt: Sinai] 845

Tribus Rhodometrini Agenjo, 1952 Rhodometra Meyrick 1892 R. sacraria (Linnaeus, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/193 (= debiliaria Rothschild, 1914) Casilda Agenjo, 1952 C. antophilaria (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/194 C. a. rosearia (Treitschke, 1828) C. consecraria (Staudinger, 1871) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/195 (= intermediaria Turati, 1930) Ochodontia Lederer, 1853 O. adustaria (Fischer de Waldheim, 1840). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2/196 Tribus Lythriini Herbulot, 1963 279 Lythria Hübner, 1823 280 L. plumularia (Freyer, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. purpuraria (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. cruentaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. sanguinaria (Duponchel, 1842). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/1 3/2 3/3 3/4

Subfamily Larentiinae Duponchel, 1845 281 Tribus Chesiadini Herbulot, 1963 282 Schistostege Hübner, 1825 S. nubilaria (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/225 S. decussata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/226 (= forsteri Vojnits, 1973; = treitsschkei Kovacs, 1957; = dioszeghyi Kovacs, 1957; = transiens Stauder, 1920; = rumelica Rebel & Zerny, 1931) S. d. dinarica (Schawerda, 1913) S. d. flavata Barajon, 1952 S. d. lugubrata Hartig, 1971 Amygdaloptera Gumppenberg, 1887 A. testaria (Fabricius, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/227 Odezia Boisduval, 1840 O. atrata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/228 O. a. meridionalis Reisser, 1935 279 Lythriini closely related with Rhodometrini, but tribus rank formally maintained by Õunap et al. (2008; 2009) and Murillo-Ramos et al. (2019). 280 Transferred from Larentiinae to Sterrhinae in Õunap et al. (2008), not yet considered in Leraut (2009). 281 Order of tribes adapted to Õunap et al. (2016). The ‘basal’ tribes (in sister relationship to the rest of the Larentiinae) are positioned at the head. In this checklist the basal (hypothetically ancestral) node of Cidariini + rest of Xanthorhoini lineage is inverted compared with the tree geometry in Õunap et al. (2016) (both options just ‘mirrored’, thus congruent), thus maintaining the order of tribes as presented in Hausmann & Viidalepp (2012). 282 In Õunap et al. (2016) the most ‘basal’ (European) tribe, in sister relationship to the rest of the Larentiinae. Only the Neotropical tribe Dyspteridini results in a more basal position.

846

Docirava Walker, 1863 D. dervenaria (Mentzer, 1981). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/229 * D. mundata (Staudinger, 1892) [Turkey, Levant] * D. mundulata (Guenée, 1858) [Turkey, Levant] D. musculata (Staudinger, 1892) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/230 * Lehmannodes Rajaei & Stüning, 2013 * L. guneyi (Riemis, 1992) [Turkey] Carsia Hübner, 1825 C. sororiata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/231 C. s. anglica Prout, 1937 C. s. imbutata (Hübner, 1813) * C. perpetuata (Lederer, 1870) [Caucasus, Georgia] (= caucasica Niesiolowski, 1937) C. lythoxylata (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/232 * C. l. christophi (Prout, 1937) [Caucasus, Transcaucasus, Turkey] Aplocera Stephens, 1827 A. columbata (Metzner, 1845) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bohatschi (Püngeler, 1914) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. corsalta (Schawerda, 1928) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= sardalta Bytinski-Salz, 1934) * A. fraudulentata (Herrich-Schäffer, 1861) [Turkey] A. plagiata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. p. hausmanni Expòsito Hermosa, 1998 A. vivesi Expòsito Hermosa, 1998. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. efformata (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= britonata Leraut, 1995) * A. arenbergeri Weisert, 2004 [Algeria, Tunisia] A. cretica (Reisser, 1974) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. praeformata (Hübner, 1826). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= gibeauxi Leraut, 1995) A. p. urbahni Dufay, 1981 * A. uniformata (Urbahn, 1971) [Caucasus, Transcaucasus, Turkey] A. simpliciata (Treitschke, 1835). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. s. pierretaria (Guillemot, 1856) A. s. balcanica (Züllich, 1936) A. s. graeciata (Staudinger, 1901) A. annexata (Freyer, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= boisduvaliata Duponchel, 1831) * A. obsitaria (Lederer, 1853) [Turkey] * A. o. anatolica (Wehrli, 1931) [Turkey] * A. o. evanescens (Wehrli, 1931) [Turkey, Armenia, Azerbaijan] * A. opificata (Lederer, 1870) [Turkey, Armenia, Azerbaijan] * A. fraternata (Herrich-Schäffer, 1861) [Turkey] * A. numidaria (Herrich-Schäffer, 1852) [Turkey, Georgia, Armenia]

3/233 3/234 3/235 3/236 3/237 3/238 3/239 3/240

3/241

3/242

847

Chesias Treitschke, 1825 C. legatella (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. capriata Prout, 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. isabella Schawerda, 1915. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rufata (Fabricius, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. r. cinereata Staudinger, 1901 C. r. pinkeri Schawerda, 1939 (?= zuellichi Schawerda, 1939) * C. plumbeata Staudinger, 1901 [NW. Africa] C. angeri Schawerda, 1919. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. leuconeura (Prout, 1931) [Turkey] (= sureyata Rebel, 1933) * C. rhegmatica Prout, 1937 [Cyprus] C. linogrisearia Constant, 1888. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/243 3/244 3/245 3/246

3/247

3/248

* Chesistege Viidalepp, 1990 * C. korbi (Bohatsch, 1910) [Armenia, Turkey] * C. k. taurica (Wehrli, 1938) [Turkey, Levant] * Grammochesias Prout, 1937 * G. rotroui Prout, 1937 [Algeria] * G. hippocastanarioides (Rothschild, 1914) [Algeria] Lithostege Hübner, 1825 L. bosporaria (Herrich-Schäffer, 1848) 283 /�������������������������������������������������������������� 3/249 * L. witzenmanni Standfuss, 1892 [Turkey] L. coassata (Hübner, 1825). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/250 (= stepparia Boisduval, 1848) * L. c. ochraceata Staudinger, 1897 [Turkey, Transcaucasus, Caucasus] L. farinata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/251 * L. narynensis Prout, 1938 [W. Kazakhstan?, Caucasus] L. griseata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/252 L. g. cycnaria Guenée, 1858 L. duponcheli Prout, 1938. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/253 L. clarae Gastón & Redondo, 2004. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/254 * L. cinerata Turati, 1924 [Libya, Egypt] * L. c. apicata Prout, 1938 [NW. Africa] * L. palaestinensis Amsel, 1935 [Turkey, Cyprus, Levant] * L. ancyrana Prout, 1938 [Turkey] L. infuscata (Eversmann, 1837). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/255 * L. flavicornata (Zeller, 1847) [Transcaucasus, Turkey] * L. odessaria (Boisduval, 1848) [Transcaucasus, Caucasus] L. castiliaria Staudinger, 1877. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/256 L. fissurata (Mabille, 1888). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/257 * L. biermis Prout, 1922 [Morocco] (= lecerfi Lucas, 1937) * L. notata Bang-Haas, 1906 [N. Africa, S. Levant, Arabia]

283 Possibly extinct from the territory of Europe.

848

Tribus Trichopterygini Warren, 1894 284 Celonoptera Lederer, 1862 (= Sparta Staudinger, 1862) C. mirificaria Lederer, 1862. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/258 C. m. paradoxaria (Staudinger, 1862) Epilobophora Inoue, 1943 E. sabinata (Geyer, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/259 E. s. teriolensis (Kitt, 1932) * E. kostjuki Tikhonov, 1994 [Caucasus, Georgia, Turkey?] Oulobophora Staudinger, 1892 O. internata (Püngeler, 1888). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/260 O. externaria (Herrich-Schäffer, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/261 O. e. turcosyrica Wehrli, 1934 Lobophora Curtis, 1825 L. halterata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/262 * Episauris Rebel, 1898 * E. kiliani Rebel, 1898 [Canary Islands] Pterapherapteryx Curtis, 1825 P. sexalata (Retzius, 1783) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/263 Nothocasis Prout, 1936 N. sertata (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/264 N. rosariae Scalercio, Infusino & Hausmann, 2016. . . . . . . . . . . . . . . . . . . . . . . . . . . 6/192 Acasis Duponchel, 1845 A. viretata (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/265 A. appensata (Eversmann, 1842). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/266 A. a. callaina Hausmann & Huemer, 2011 Trichopteryx Hübner, 1825 T. polycommata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/267 T. p. grisea (Djakonov, 1926) T. carpinata (Borkhausen, 1794). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/268 Not assigned to tribus 285 Anticollix Prout, 1938 A. sparsata (Treitschke, 1828) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/224

284 In Õunap et al. (2016) the second-most ‘basal’ (European) tribe, in sister relationship to the rest of the Larentiinae, except for the Neotropical tribe Dyspteridini and the Chesiadini (see above) which result in a more basal position. 285 Position according to Õunap et al. (2016) and Beljaev (2016).

849

Tribus Asthenini Warren, 1894 286 Minoa Treitschke, 1825 M. murinata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/180 M. m. monochroaria Herrich-Schäffer, 1848 Asthena Hübner, 1825 A. albulata (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/181 * A. percandidata (Christoph, 1893) [Caucasus, Trans­caucasus, Turkey] A. lactularia (Herrich-Schäffer, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/182 (= nymphulata Guenée, 1858) A. anseraria (Herrich-Schäffer, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/183 * A. a. candidissima (Staudinger, 1897) [Transcaucasus] Euchoeca Hübner, 1823 E. nebulata (Scopoli, 1763) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/184 Hydrelia Hübner, 1825 H. sylvata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/185 (= testaceata Donovan, 1810) H. flammeolaria (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/186 Venusia Curtis, 1839 V. cambrica Curtis, 1839 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/187 V. blomeri (Curtis, 1832). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/188 Tribus Scotopterygini Warren, 1895 287 Scotopteryx Hübner, 1825 S. coarctaria (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/9 * S. c. infuscata (Staudinger, 1871) [Turkey] S. vicinaria (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/10 (= roesleri Vojnits, 1973) * S. v. hyrcanaria (Staudinger, 1892) [Georgia, Armenia, Azerbaijan] * S. subvicinaria (Staudinger, 1892) [Caucasus, Turkey] S. burgaria (Eversmann, 1843). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/11 * S. perplexaria (Staudinger, 1892) [Armenia] * S. elbursica (Bytinski-Salz & Brandt, 1935) [Armenia, Azerbaijan] * S. kuznetzovi (Wardikjan, 1957) [Turkey, Armenia] * S. pinnaria (Christoph, 1888) [Caucasus, Georgia, Armenia] * S. p. miljanovskii (Viidalepp, 1977) [Caucasus] S. mucronata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/12 S. m. umbrifera (Heydemann, 1925) S. m. scotica (Cockayne, 1940) S. luridata (Hufnagel, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/13 286 Position according to Õunap et al. (2016). From that analysis, Asthenini result as a sister group to the large bulk of Larentiinae tribes of “Eupitheciini lineage” + “Xanthorhoini lineage”. 287 Upgraded from synonymy of Xanthorhoini to tribe rank in Õunap et al. (2016) and Beljaev (2016). Belonging to the “Xanthorhoini lineage” in Õunap et al. (2016). In this checklist the basal (hypothetically ancestral) nodes of “Eupitheciini lineage” + “Xanthorhoini lineage” as well as of Cidariini + rest of Xanthorhoini lineage are inverted compared with the tree geometry in Õunap et al. (2016) (both options just ‘mirrored’, thus congruent), thus maintaining the order of tribes as presented in Hausmann & Viidalepp (2012).

850

S. l. plumbaria (Fabricius, 1775) S. ignorata Huemer & Hausmann, 1998. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. peribolata (Hübner, 1817) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * S. p. chouika (Oberthür, 1909) [NW. Africa] S. proximaria (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bipunctaria (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. b. cretata (Prout, 1937) S. octodurensis (Favre, 1903). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. o. lozerae (Herbulot, 1957) S. o. nevadina (Wehrli, 1927) (= ibera Prout, 1937) S. olympia Rezbanyai-Reser, 2003. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * S. aelptes (Prout, 1937) [Turkey, Caucasus, Trans­caucasus] S. moeniata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. angularia (Villers, 1789). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= diniensis Neuburger, 1906) S. coelinaria (Graslin, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. obvallaria (Mabille, 1867) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. alfacaria (Staudinger, 1859). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= albarracina Zerny, 1927) * S. a. transmarina (Zerny, 1934) [NW. Africa] * S. alpherakii (Erschov, 1877) [Caucasus, Trans­caucasus, Turkey] * S. diana (Riemis, 1997) [Turkey] * S. langi (Christoph, 1885) [Armenia] S. chenopodiata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/14 3/15 3/16 3/17 3/18

3/19 3/20 3/21 3/22 3/23 3/24

3/25

Tribus Cataclysmini Herbulot, 1962 288 Phibalapteryx Stephens, 1829 P. virgata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/5 Cataclysme Hübner, 1825 C. riguata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/6 C. dissimilata (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/7 C. uniformata (Bellier, 1862). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/8 * C. multistriata Rothschild, 1914 [Algeria] * C. subtilisparsata Wehrli, 1932 [Turkey] 289 Tribus Xanthorhoini Pierce, 1914 Psychophora Kirby, 1824 P. sabini (Kirby, 1824) * P. s. sabini (Kirby, 1824) [North America] P. s. frigidaria (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/26 * P. cinderella Viidalepp, 2001 [Nowaya Zemlya]

288 Suggested (but not formally changed) as potentially subordinated under a wider concept of Xanthorhoini by Õunap et al. (2016). Preliminarily we maintain the taxon at tribe level in accordance with Hausmann & ­Viidalepp (2012). 289 Recognized as bona species in Stadie et al. (2014).

851

Orthonama Hübner, 1825 O. vittata (Borkhausen, 1794). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/27 (= lignata Hübner, 1799) O. obstipata (Fabricius, 1794). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/28 (= fluviata Hübner, 1799) Xanthorhoe Hübner, 1825 X. decoloraria (Esper, 1806). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= munitata Hübner, 1809) X. d. hethlandica (Prout, 1901) X. d. arcticaria (Keferstein, 1835) X. fluctuata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. f. iberaria (Staudinger, 1871) * X. f. multistriga (Oberthür, 1910) [Algeria, Tunisia] * X. f. herculeana (Zerny, 1934) [Morocco] X. disjunctaria (la Harpe, 1860). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. sardisjuncta Hausmann, 2011. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * X. acutangulata (Christoph, 1887) [Georgia, Armenia, Azerbaijan] * X. bigeminata (Christoph, 1885) [Georgia, Armenia, Azerbaijan] X. oxybiata (Millière, 1872). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. skoui Viidalepp & Hausmann, 2003. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. friedrichi Viidalepp & Skou, 2003. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * X. pseudogaliata (Staudinger, 1898) [Turkey, Levant] * X. inconsiderata (Staudinger, 1892) [Turkey] * X. pederi Hausmann & Friedrich, 2011 [Cyprus] X. incursata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. i. altarhaetica Wolfsberger, 1953 X. i. cindrelaria (Dannehl, 1927) X. uralensis Choi, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= derzhavini auct. nec Djakonov, 1931) X. annotinata (Zetterstedt, 1839). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. a. pseudoannotinata Vasilenko, 2007 X. majorata Heydemann, 1936 * X. m. majorata Heydemann, 1936 [C. Asia] X. m. pseudomajorata Vasilenko, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. biriviata (Borkhausen, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. spadicearia (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. vidanoi Parenzan & Hausmann, 1994. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. ferrugata (Clerck, 1759) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. abrasaria (Herrich-Schäffer, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. designata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. d. islandicaria (Staudinger, 1871) X. d. faeroensis (Wolff, 1929) X. montanata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= lapponica Staudinger, 1871) X. m. shetlandica (Weir, 1880) X. iberica (Staudinger, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . X. quadrifasiata (Clerck, 1759) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= quadrifasciata auct.) 852

3/29

3/30

3/31 3/32 3/33 3/34 3/35

3/36 3/37 3/38

3/39 3/40 3/41 3/42 3/43 3/44 3/45 3/46 3/47 3/48

* X. rectifasciaria (Lederer, 1853) [Georgia] * X. r. rectior (Wiltshire, 1976) [Turkey] * X. conspectaria (Mann, 1859) [Madeira] * X. inaequata (Warren, 1905) [Açores] * X. rupicola (Wollaston, 1858) [Madeira] (= conspectaria Mann, 1859) Protorhoe Herbulot, 1951 P. unicata (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/49 (= avetianae (Wardikjan, 1974) [Armenia]) 290 P. corollaria (Herrich-Schäffer, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/50 * P. beshkovi (Rajaei, Stadie & Hausmann, 2017) [Turkey] * P. drechseli (Rajaei, Stadie & Hausmann, 2017) [Turkey] Juxtephria Viidalepp, 1976 J. consentaria (Freyer, 1846). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/51 Catarhoe Herbulot, 1951 C. hortulanaria (Staudinger, 1879). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. h. palaestinensis (Staudinger, 1895) [Cyprus, Israel] C. basochesiata (Duponchel, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= cupreata Herrich-Schäffer, 1838) C. putridaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= bulgariata Millière, 1868) C. mazeli Viidalepp, 2008. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. permixtaria (Herrich-Schäffer, 1856). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. semnana Wiltshire, 1970 [Turkey, Transcaucasus] C. cuculata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. c. lucasi (Herbulot, 1984) [Algeria] * C. c. decolor (Schwingenschuss, 1939) [Turkey, Caucasus, Transcaucasus] C. rubidata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. r. fumata (Eversmann, 1844) * C. arachne (Wiltshire, 1967) [Armenia]

3/52 3/53 3/54 3/55 3/56 3/57 3/58

Costaconvexa Agenjo, 1949 C. polygrammata (Borkhausen, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/59 * C. centrostrigaria (Wollaston, 1858) [Canary Islands, Madeira, Açores] Camptogramma Stephens, 1831 C. bilineata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= atlantica Staudinger, 1892) C. b. bohatschi (Prout, 1914) * C. b. numidica (Rothschild, 1925) [NW. Africa] C. bistrigata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. grisescens (Staudinger, 1892). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. scripturata (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. s. albidaria (Sohn-Rethel, 1929) C. s. poliata (Schawerda, 1913) C. s. rilica (Prout, 1938)

3/60

3/61 3/62 3/63

290 In Hausmann & Viidalepp (2012) listed as valid species, for synonymy see Rajaei et al. (2017).

853

Epirrhoe Hübner, 1825 E. tristata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/64 E. pupillata (Thunberg, 1788). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/65 E. tartuensis Möls, 1965. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/66 E. alternata (Müller, 1764) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/67 (= sociata Borkhausen, 1794; = obscurata South, 1898; = islandica Prout, 1915; = latevittata Turati, 1913) E. rivata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/68 E. hastulata (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/69 E. molluginata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/70 E. galiata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/71 * E. g. orientata (Staudinger, 1901) [Turkey, Trans­caucasus] E. balearia Fischer, 2011. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/72 E. timozzaria (Constant, 1884). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/73 E. sandosaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/74 * E. s. bertrandi (Rothschild, 1912) [NW. Africa] * E. s. libycaria (Turati, 1930) [Libya] * E. mosulensis (Schawerda, 1924) [Turkey, Levant] Tribus Euphyiini Herbulot, 1962 291 Euphyia Hübner, 1825 E. biangulata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= picata Hübner, 1813) E. b. baltica (Prout, 1939) E. vallantinaria (Oberthür, 1890). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. unangulata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. u. gracilaria (Bang-Haas, 1906) [W. Siberia] E. adumbraria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. frustata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. f. griseoviridis (Kitt, 1926) E. f. fulvocinctata (Staudinger, 1871) * E. maximiliana (Reisser, 1933) [Morocco] E. mesembrina (Rebel, 1927). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. sintenisi (Staudinger, 1892) [Turkey, Caucasus, Georgia, Armenia] * E. chalusata Wiltshire, 1970 [Azerbaijan, Turkey]

3/75 3/76 3/77 3/78 3/79

3/80

Tribus Larentiini Duponchel, 1845 292 Mattia Viidalepp, 2012 M. callidaria (Joannis, 1891). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/81 (= adlata Staudinger, 1895) Antilurga Herbulot, 1951 A. alhambrata (Staudinger, 1859). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/82 * A. a. altatlas Herbulot, 1981 [Morocco] * A. cabrerai (Pinker, 1962) [Canary Islands]

291 Position and tribe status confirmed by Õunap et al. (2016). 292 Position and tribe status confirmed by Õunap et al. (2016).

854

Earophila Gumppenberg, 1887 E. badiata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/83 E. b. fennokarelica (Kaisila, 1945) * E. b. tellensis (Herbulot, 1957) [Algeria] * E. semna Prout, 1929 [Algeria] * E. beduina (Turati, 1930) [Libya] Anticlea Stephens, 1831 A. derivata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/84 Mesoleuca Hübner, 1825 M. albicillata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/85 Larentia Treitschke, 1825 L. clavaria (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/87 L. c. datinaria (Oberthür, 1890) L. c. pallidata (Staudinger, 1901) L. malvata (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/88 * Herbulotina Pinker, 1969 * H. grandis (Prout, 1914) [Canary Islands] * H. g. grancanariae (Pinker, 1963) [Canary Islands] * H. g. lapalmae (Pinker, 1963) [Canary Islands] * H. maderae Pinker, 1971 [Madeira] * H. feliciaria (Lucas, 1907) [Morocco, Algeria] * H. f. hebes (Rungs, 1950) [Morocco] * H. carolata (Lucas, 1938) [Morocco] * H. berberina (Herbulot, 1981) [Morocco] Entephria Hübner, 1825 E. polata (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/89 * E. p. transsibirica Vasilenko, 1990 [N. Siberia] E. byssata Aurivillius, 1891 293/������������������������������������������������������������������������������������ 3/90 * E. punctipes (Curtis, 1835) [North America] * E. p. brullei (Levebvre, 1836) [Greenland] E. nobiliaria (Herrich-Schäffer, 1852). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/91 E. n. borearia (Prout, 1914) E. flavata (Osthelder, 1929). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/92 * E. rjabovi Tikhonov, 1994 [Caucasus, Armenia] * E. ignorata (Staudinger, 1892) [Caucasus, Turkey] * E. calcephila Tikhonov, 1994 [Caucasus] * E. tejmurovi Tikhonov, 1994 [Caucasus] * E. muscosaria (Christoph, 1893) [Caucasus] E. cyanata (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/93 (= gerennae Gyulai, 1984) E. c. petronensis Herbulot, 1968 E. c. leucocyanata (Reisser, 1935)

293 Recently raised from synonymy of E. punctipes to species rank by Aarvik et al. (2017), see also information in this volume.

855

E. c. bubaceki (Reisser, 1926) (= altivolans Wehrli, 1926) E. c. acyana (Prout, 1938) E. caeruleata (Guenée, 1858). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. flavicinctata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. f. septentrionalis (Warnecke, 1934) E. f. ruficinctata (Guenée, 1858) E. f. corsaria (Schawerda, 1928) E. f. veletaria (Wehrli, 1926) * E. f. elbrusensis Tikhonov, 1994 [Caucasus] E. contestata (Vorbrodt, 1913). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. infidaria (la Harpe, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. caesiata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. c. hethlandicaria (Bang-Haas, 1910) E. c. norvegica (Strand, 1902) E. c. glaciata (Keferstein, 1831)

3/94 3/95

3/96 3/97 3/98

Spargania Guenée, 1858 294 S. luctuata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/99 S. l. borealis (Petersen, 1902) Not assigned to tribus Pelurga Hübner, 1825 295 P. comitata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/86 Tribus Hydriomenini Meyrick, 1892 296 Hydriomena Hübner, 1825 H. furcata (Thunberg, 1784). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. f. saga (Prout, 1938) H. impluviata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. i. insulicolata (Schawerda, 1933) H. sanfilensis (Stauder, 1915). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. ruberata (Freyer, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/100 3/101 3/102 3/103

Tribus Stamnodini Forbes, 1948 297 Stamnodes Guenée, 1858 S. depeculata (Lederer, 1870) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/104 * S. d. narzanica Alphéraky, 1877 [Transcaucasus, Turkey]

294 Spargania listed under Euphyiini in Beljaev (2016). 295 Position within the Larentiini (cf. Hausmann & Viidalepp 2012) not confirmed by later molecular analyses. Basal and isolated position within the ‘Xanthorhoini lineage’ postulated by Õunap et al. (2016). 296 Position and tribe status confirmed by Õunap et al. (2016). 297 Position and tribe status confirmed by Õunap et al. (2016).

856

Tribus Cidariini Duponchel, 1845 298 Almeria Agenjo, 1949 A. kalischata (Staudinger, 1870). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/105 * A. k. rubrotincta (Zerny, 1934) [Morocco] Heterothera Inoue, 1943 H. serraria (Lienig, 1846). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/106 H. taigana (Djakonov, 1926). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/107 Pennithera Viidalepp, 1980 P. firmata (Hübner, 1822). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/108 P. f. tavoilloti (Mazel, 1998) P. f. tyrrhenica Tautel & Billi, 2009 P. ulicata (Rambur, 1834) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/109 Thera Stephens, 1831 T. cognata (Thunberg, 1792). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= geneata Feisthamel, 1835) * T. variolata (Staudinger, 1899) [NW. Africa] T. variata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. v. balcanicola Lattin, 1951 * T. v. subtaurica (Wehrli, 1932) [Turkey, Cyprus, Levant] T. britannica (Turner, 1925). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= albonigrata Gornik, 1942) T. vetustata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= stragulata Hübner, 1809) T. cembrae (Kitt, 1912) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. c. mugo Burmann & Tarmann, 1983 T. obeliscata (Hübner, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. o. tristrigaria (Donovan, 1808) T. juniperata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * T. j. loeberbaueri Friedel, 1969 [Turkey, Caucasus] T. cupressata (Geyer, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/110 3/111 3/112 3/113 3/114 3/115 3/116 3/117

Plemyria Hübner, 1825 P. rubiginata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/118 (= bicolorata Hufnagel, 1767 (homonym); = plumbata Curtis, 1837) Cidaria Treitschke, 1825 C. fulvata (Forster, 1771) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/119 * C. distinctata Staudinger, 1892 [Kazakhstan, SW. Siberia, C. Asia] Polythrena Guenée, 1858 P. coloraria (Herrich-Schäffer, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/120

298 Position and tribe status confirmed by Õunap et al. (2016). In this checklist the basal (hypothetically ancestral) node of Cidariini + rest of Xanthorhoini lineage is inverted compared with the tree geometry in Õunap et al. (2016) (both options just ‘mirrored’, thus congruent), thus maintaining the order of tribes as presented in Hausmann & Viidalepp (2012).

857

* Trichodezia Warren, 1895 * T. haberhaueri (Lederer, 1864) [Caucasus, E. Trans­caucasus, Turkey] * T. h. ledereri (Staudinger, 1901) [Armenia] Electrophaes Prout, 1923 E. corylata (Thunberg, 1792) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/121 Cosmorhoe Hübner, 1825 C. ocellata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/122 Eustroma Hübner, 1825 E. reticulata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/123 * E. mardinata (Staudinger, 1895) [Turkey] Eulithis Hübner, 1821 E. prunata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. p. teberdensis (Alberti, 1969) [Turkey, Caucasus] E. testata (Linnaeus, 1761) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. populata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. mellinata (Fabricius, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pyropata (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. peloponnesiaca (Rebel, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. roessleraria (Staudinger, 1870) [Turkey] * E. r. pseudoledereri (Schwingenschuss, 1939) [Turkey]

3/124 3/125 3/126 3/127 3/128 3/129

Gandaritis Moore, 1868 G. pyraliata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/130 Ecliptopera Warren, 1894 E. capitata (Herrich-Schäffer, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/131 E. silaceata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/132 (= ‡ posticata Fabricius, 1794) Chloroclysta Hübner, 1825 C. siterata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/133 C. miata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/134 C. m. clara (Thierry-Mieg, 1915) * C. m. buzurga Wiltshire, 1970 [Turkey, Transcaucasus, Caucasus, Levant] * C. guriata (Emich, 1873) [Transcaucasus] Dysstroma Hübner, 1825 D. citrata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/135 (= pythonissata Millière, 1870) D. c. unicolorata (Staudinger, 1901) D. pseudimmanata Heydemann, 1929 299 /������������������������������������������������������������������ 3/136 D. truncata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/137 D. t. concinnata (Stephens, 1831) D. infuscata (Tengström, 1869). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/138 D. latefasciata (Blöcker, 1908). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/139 299 According to Beljaev (2016) this taxon is misidentified in GME3 (fig. 136a) and refers to D. truncata trans­ baicalensis, but true D. pseudimmanata also occurring in Europe (Urals).

858

Colostygia Hübner, 1825 C. aptata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/140 C. olivata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/141 C. hilariata (Pinker, 1953). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/142 C. wolfschlaegerae (Pinker, 1953) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/143 C. fitzi (Schawerda, 1914). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/144 * C. fezae Hausmann, 2011 [Turkey, Caucasus] C. cyrnea (Wehrli, 1925). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/145 C. pectinataria (Knoch, 1781). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/146 C. aqueata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/147 C. a. deviridata Hausmann, 2012 (= ‡ pyrenaeata Bubacek, 1924 (homonym); = ‡ neva­densis Reisser, 1931 (homonym)) C. a. hercegovinensis (Rebel, 1901) C. stilpna (Prout, 1924). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/148 C. s. mounieri (Herbulot, 1945) C. turbata (Hübner, 1799) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/149 C. t. fuscolimbata (Tengström, 1875) C. t. pyrenaeata (Oberthür, 1882) C. kollariaria (Herrich-Schäffer, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/150 (= larentiaria Bruand, 1850) C. laetaria (la Harpe, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/151 C. puengeleri (Stertz, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/152 C. p. varonaria (Vorbrodt, 1913) C. p. sauteri Rezbanyai-Reser, 1977 C. p. bavaricaria (Löberbauer, 1955) C. austriacaria (Herrich-Schäffer, 1852) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/153 C. a. noricaria (Löberbauer, 1955) C. a. gremmingeri (Schawerda, 1942) C. a. hoefneri (Schawerda, 1942) C. a. distans Krampl & Marek, 1991 C. kitschelti (Rebel, 1934). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/154 C. tempestaria (Herrich-Schäffer, 1856). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/155 C. multistrigaria (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/156 C. m. olbiaria (Millière, 1865) C. sericeata (Schwingenschuss, 1926). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/157 C. s. holli (Prout, 1938) Coenotephria Prout, 1914 C. salicata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. s. latentaria (Curtis, 1830) C. ablutaria (Boisduval, 1840) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. a. probaria (Herrich-Schäffer, 1852) C. a. hangayi (Vojnits, 1986) C. christlae Hausmann & Leipnitz, 2012. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * C. florianii Hausmann, 2011 [Turkey, Lebanon] C. tophaceata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. t. jurassica (Vorbrodt, 1914) C. antonii Hausmann, 2011. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. schneideraria (Lederer, 1855) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

3/158 3/159 3/160 3/161 3/162 3/163

859

C. s. eteocretica (Rebel, 1906) * C. s. taurica (Staudinger, 1901) [Turkey] Nebula Bruand, 1846 N. nebulata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/164 (= vallesaria la Harpe, 1853) N. n. albicans (Sohn-Rethel, 1929) N. pirinica (Reisser, 1936) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/165 N. senectaria (Herrich-Schäffer, 1852) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/166 (= ludificata Staudinger, 1870; = sultana Schwingenschuss, 1939; = decipiata Staudinger, 1901) * N. juvenilata (Zerny, 1933) [Levant] * N. mantelorum (Hausmann, 1997) [Sinai] * N. approximata (Staudinger, 1879) N. achromaria (la Harpe, 1853). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/167 N. carlae Hausmann, 2011 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/168 N. ibericata (Staudinger, 1871). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/169 N. i. fiumii Hausmann, 2011 N. i. numidiata (Staudinger, 1892) * N. rastremata (Turati & Krüger, 1936) [Libya] * N. impunctata (Staudinger, 1879) [Turkey] * N. propagata (Christoph, 1893) [Caucasus] * N. apiciata (Staudinger, 1892) [Georgia, Azerbaijan] Lampropteryx Stephens, 1831 L. suffumata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/170 L. s. arctica (Sparre-Schneider, 1895) L. otregiata (Metcalfe, 1917). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/171 Tribus Operophterini Packard, 1876 300 Malacodea Tengström, 1869 M. regelaria Tengström, 1869. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/172 Operophtera Hübner, 1825 O. fagata (Scharfenberg, 1805) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/173 O. brumata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/174 Epirrita Hübner, 1822 E. dilutata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/175 E. terminassianae Wardikjan, 1974 301 /���������������������������������������������������������������������� 3/176 E. christyi (Allen, 1906). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/177 E. autumnata (Borkhausen, 1794) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/178 E. a. altivagata (Hartig, 1938) E. filigrammaria (Herrich-Schäffer, 1848). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/179

300 In Õunap et al. (2016) confirmed at tribe level and resulting as part of the “Eupitheciini lineage”. 301 Spelling of author name in the original description: ‘Wardikjan’. In GME3 spelled as ‘Vardikjan’.

860

Tribus Solitaneini Leraut, 1980 302 Solitanea Djakonov, 1924 S. mariae (Stauder, 1921) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/206 Baptria Hübner, 1825 B. tibiale (Esper, 1804). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/207 B. t. borealis Lankiala, 1937 B. t. fennica Lankiala, 1937 Tribus Rheumapterini Herbulot, 1962 303 Hospitalia Agenjo, 1950 H. flavolineata (Staudinger, 1883) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/191 Rheumaptera Hübner, 1822 304 R. hastata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/192 R. h. thulearia (Herrich-Schäffer, 1848) R. h. nigrescens (Cockerell, 1889) R. subhastata (Nolcken, 1870). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/193 R. s. moestata (Nolcken, 1970) Hydria Hübner, 1822 H. undulata (Linnaeus, 1758) 305/�������������������������������������������������������������������������������� 3/194 H. cervinalis (Scopoli, 1763) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/195 H. c. simplonica (Wackerzapp, 1890) H. c. hawelkae (Schawerda, 1915) * H. ithys (Prout, 1937) [Morocco, Algeria] H. incertata (Staudinger, 1882). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/196 H. gudarica (Dufay, 1983) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/197 H. montivagata (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/198 H. m. andalusica (Ribbe, 1912) (= moscardonica Laever, 1983) * H. m. hyrcana (Staudinger, 1871) [Georgia, Armenia, Azerbaijan] Tribus Phileremini Pierce, 1914 306 Philereme Hübner, 1825 307 P. vetulata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/189 P. transversata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/190 * P. senescens (Staudinger, 1892) [Transcaucasus, Turkey]

302 In Õunap et al. (2016) resulting as sister group of Operophterini (but only Baptria included in the study). 303 Tribe status and position confirmed by Õunap et al. (2016), but subordinated under Triphosini in Beljaev (2016). 304 Rheumaptera listed under Triphosini in Beljaev (2016). 305 Combined with genus Rheumaptera in Beljaev (2016). 306 Tribe status and position confirmed by Õunap et al. (2016), but subordinated under Triphosini in Beljaev (2016). 307 Philereme listed under Triphosini in Beljaev (2016).

861

Tribus Triphosini Tutt, 1896 308 Triphosa Stephens, 1829 T. sabaudiata (Duponchel, 1830) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * T. agnata Le Cerf, 1918 [Turkey] 309 T. petronata Le Cerf, 1918. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * T. taochata Lederer, 1870 [Caucasus, Transcaucasus, Turkey] T. dubitata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. tauteli Leraut, 2008 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= herzeti Leraut, 2008) * T. dyriata Powell, 1941 [Morocco, Algeria]

3/199 3/200 3/201 3/202

Pareulype Herbulot, 1951 P. berberata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/203 P. b. sineliturata (Culot, 1919) (= cinerea Herbulot, 1977) P. b. maindroni Herbulot, 1977 P. b. nevadensis (Rebel, 1916) * P. b. mauretanica (Reisser, 1934) P. lasithiotica (Rebel, 1906). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/204 P. casearia (Constant, 1884). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/205 Tribus Melanthiini Duponchel, 1845 310 Coenocalpe Hübner, 1825 C. lapidata (Hübner, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/208 C. millierata (Staudinger, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/209 * C. m. zerhounaria (Oberthür, 1921) [Morocco] Horisme Hübner, 1825 H. vitalbata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. corticata (Treitschke, 1835). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. tersata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. radicaria (la Harpe, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= laurinata Schawerda, 1919) H. r. insularis Bytinski-Salz, 1937 H. exoletata (Herrich-Schäffer, 1838). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. predotai Bytinski-Salz, 1937 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. scorteata (Staudinger, 1901) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. aemulata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. incurvaria (Erschov, 1877) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. calligraphata (Herrich-Schäffer, 1838). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . H. c. taeniolata (Eversmann, 1843) * H. milvaria (Christoph, 1893) [Caucasus, Trans­caucasus] * H. intricata Staudinger, 1882 [Kazakhstan] H. aquata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

308 Tribe status and position suggested by Schmidt (2015) and Õunap et al. (2016). 309 T. agnata is regarded as junior synonym of T. sabaudiata by Wanke et al. (2019). 310 Tribe status and position confirmed by Õunap et al. (2016).

862

3/210 3/211 3/212 3/213 3/214 3/215 3/216 3/217 3/218 3/219

3/220

Melanthia Duponchel, 1829 M. procellata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/221 M. alaudaria (Freyer, 1846). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/222 M. mandshuricata (Bremer, 1864) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3/223 Tribus Perizomini Herbulot, 1961 Pseudobaptria Inoue, 1982 P. bogumilaria (Rebel, 1904). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/1 Mesotype Hübner, 1825 M. didymata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/2 M. d. hethlandica (Rebel, 1910) M. parallelolineata (Retzius, 1783). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/3 M. verberata (Scopoli, 1763). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/4 Perizoma Hübner, 1825 * P. gigas Wiltshire, 1976 [Turkey] P. affinitata (Stephens, 1831). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/5 P. alchemillata (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/6 P. hydrata (Treitschke, 1829). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/7 * P. parahydrata Alberti, 1969 [Caucasus] P. lugdunaria (Herrich-Schäffer, 1855) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/8 P. bifaciata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/9 P. flavosparsata (Wagner, 1926). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/10 P. f. uniformata Mentzer, 1974 P. f. cleui Leraut, 2009 311 P. minorata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/11 P. m. ericetata (Stephens, 1831) P. m. norvegica Prout, 1906 P. blandiata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/12 P. b. perfasciata Prout, 1915 P. albulata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/13 P. a. thules (Weir, 1880) P. flavofasciata (Thunberg, 1792) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/14 P. obsoletata (Herrich-Schäffer, 1838). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/15 P. o. avilaria Reisser, 1936 312 P. juracolaria (Wehrli, 1919) 313/�������������������������������������������������������������������������������� 6/193 P. j. reisseri (Schawerda, 1932) 314 P. incultaria (Herrich-Schäffer, 1848) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/16 P. barrassoi Zahm, Cieslak & Hausmann, 2006 315 /������������������������������������������������������ 6/194 311 Validity at subspecies rank awaiting further confirmation. No differences in genitalia. 312 Validated at subspecies rank by Berard et al. (2005), tentatively raised to species rank by Taurand & Tautel (2019), identity and taxonomic status requiring further integrated taxonomic study including DNA barcode analysis. 313 New record for Europe in Berard et al. (2005). 314 Taxon reisseri Schawerda, 1932 validated at subspecies rank in Leraut (2009) under his “P. alpicolaria” (sensu Leraut 2009 nec Herrich-Schäffer, 1848; identity of the latter: P. obsoletata) which is based on a misidentification and should refer to P. juracolaria. ‘Perizoma reisseri (Schawerda, 1932)’ tentatively raised to species rank by Taurand & Tautel (2019), identity and taxonomic status requiring further integrated taxonomic study including DNA barcode analysis. 315 Described from central Italy after publication of GME4 (Zahm et al. 2006), see information in this volume.

863

Martania Mironov, 2000 M. taeniata (Stephens, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/17 Gagitodes Warren, 1893 G. sagittata (Fabricius, 1787) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/18 Tribus Eupitheciini Pierce, 1914 Gymnoscelis Mabille, 1868 G. rufifasciata (Haworth, 1809) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/19 * G. insulariata (Stainton, 1859) [Canary Islands] * G. lundbladi Prout, 1939 [Madeira, Canary Islands] Chloroclystis Hübner, 1825 C. v-ata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/20 Pasiphila Meyrick, 1883 P. chloerata (Mabille, 1870). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/21 P. rectangulata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/22 P. debiliata (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/23 Eupithecia Curtis, 1825 The haworthiata species-group E. haworthiata Doubleday, 1856. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/24 The tenuiata species-group E. tenuiata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/25 E. inturbata (Hübner, 1817). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/26 The abietaria species-group E. abietaria (Goeze, 1781) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/27 E. analoga Djakonov, 1926. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/28 The linariata species-group E. linariata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pulchellata Stephens, 1831. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. p. intermedia Dietze, 1913 E. pyreneata Mabille, 1871 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. laquaearia Herrich-Schäffer, 1848 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. albidulata Staudinger, 1892 [Great Caucasus]

4/29 4/30 4/31 4/32

The ultimaria species-group E. ultimaria Boisduval, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/33 * E. golearia Lucas, 1939 [Algeria] E. usbeca Viidalepp, 1992. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/34 E. minusculata Alphéraky, 1882. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/35 * E. armeniaca Wardikjan, 1970 [Turkey, Armenia] 316 (= djakonovi Wardikjan, 1961) 317

316 Revised by Mironov (2013). 317 Revised by Mironov (2013).

864

The undata species-group E. pantellata Millière, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. p. canariata Pinker, 1965 E. plumbeolata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. p. reikjavikaria (Staudinger, 1871) E. p. lutosaria Bohatsch, 1893 E. pygmaeata (Hübner, 1799). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. nigritaria Staudinger, 1879 [Turkey] * E. puengeleri Dietze, 1913 [Turkey to Caucasus] E. undata (Freyer, 1842) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fennoscandica Knaben, 1949 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. boryata (Rebel, 1906) [Canary Islands] * E. gomerensis (Rebel, 1917) [Canary Islands]

4/36 4/37 4/38 4/39 4/40

The silenata species-group E. silenata Assmann, 1848. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/41 The venosata species-group E. carpophagata Staudinger, 1871. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/42 E. venosata (Fabricius, 1787). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/43 E. v. ochracae Gregson, 1886 318 E. v. fumosae Gregson, 1887 E. v. hebridensis Curtis, 1944 E. v. plumbea Huggins, 1962 E. schiefereri Bohatsch, 1893. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/44 E. silenicolata Mabille, 1867. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/45 * E. amasina Bohatsch, 1893 [Turkey] * E. achyrdaghica Wehrli, 1929 [Turkey] * E. rosai Pinker, 1962 [Canary Islands] * E. pinkeri Mironov, 1991 [Turkey] E. alliaria Staudinger, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/46 E. dissertata (Püngeler, 1905) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/47 E. carpophilata (Püngeler, 1905) 319 /������������������������������������������������������������������������ 6/195 The interruptofasciata species-group E. cocciferata Millière, 1864. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/48 E. abbreviata Stephens, 1831. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/49 E. dubiosa Dietze, 1910 320/���������������������������������������������������������������������������������������� 6/196 E. lentiscata Mabille, 1869. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/50 E. dodoneata Guenée, 1858. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/51 (= andrea Weisert, 2005) 321 E. reisserata Pinker, 1976. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/52 E. r. levarii Hausmann, 1991 [Levant] E. massiliata Millière, 1865 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/53 * E. maerkerata Schütze, 1938 [Jordan, Saudi Arabia] 318 319 320 321

Misspelled as ochracea in Leraut (2009). New record for Europe, see information in this volume. New record for Europe in Samos, see information in this volume. For synonymy see information in this volume.

865

E. extremata (Fabricius, 1787) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. scopariata (Rambur, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. rusicadaria Dietze, 1910 [Algeria, Tunisia] E. pusillata (Denis & Schiffermüller, 1775) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. p. scoriata Staudinger, 1857 E. sardoa Dietze, 1910. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. ericeata (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= euxinata Bohatsch, 1893 [Turkey]) 322 E. phoeniceata (Rambur, 1834). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. oxycedrata (Rambur, 1833). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= adscriptaria Staudinger, 1871 [Turkey]) 323 E. rosmarinata Dardoin & Millière, 1865. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. rhoisata Chrétien, 1917 [North Africa]

4/54 4/55 4/56 4/57 4/58 4/59 4/60 4/61

The tripunctaria species-group E. barteli Dietze, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/62 E. tripunctaria Herrich-Schäffer, 1852. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/63 The fletcherata species-group E. virgaureata Doubleday, 1861 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/64 The tantillaria species-group E. tantillaria Boisduval, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/65 E. liguriata Millière, 1884. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/66 The lariciata species-group E. lariciata (Freyer, 1842). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/67 The lanceata species-group E. lanceata (Hübner, 1825). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/68 The egenaria species-group E. groenblomi Urbahn, 1969 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. selinata Herrich-Schäffer, 1861. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. actaeata Walderdorff, 1869. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. mesogrammata Dietze, 1908 [Turkey] (= kunzi Pinker, 1976) 324 E. quercetica Prout, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= buxata Pinker, 1958) * E. tenerifensis (Rebel, 1906) [Canary Islands] * E. latipennata Prout, 1914 [Madeira] E. egenaria Herrich-Schäffer, 1848. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. repentina Vojnits & de Laever, 1978 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. schuetzeata Pinker, 1961 [Canary Islands] * E. stertzi (Rebel, 1911) [Canary Islands] E. orana Dietze, 1910. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. tricerata Wiltshire, 1990 [Saudi Arabia]

4/69 4/70 4/71 4/72

4/73 4/74 4/75

322 Synonymized in Mironov (2013). 323 For synonymy see Mironov (2013). 324 Downgraded to synonymy of mesogrammata in Mironov (2013). Possible validation at subspecies rank requiring further study.

866

* E. t. sperlichi Hausmann, 1991 [Jordan] * E. jizlensis Wiltshire, 1986 [Saudi Arabia] * E. j. muelleri Hausmann, 1991 [Jordan] E. pimpinellata (Hübner, 1813) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/76 E. karadaghensis Mironov, 1988. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/77 * E. maspalomae Pinker, 1961 [Canary Islands] The suboxydata species-group E. suboxydata Staudinger, 1897. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/78 The simpliciata species-group E. simpliciata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/79 The sinuosaria species-group E. sinuosaria (Eversmann, 1848). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/80 The innotata species-group * E. gelidata Möschler, 1860 E. g. hyperboreata Staudinger, 1861. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/81 E. nanata (Hübner, 1813) 325 /�������������������������������������������������������������������������������������� 4/82 E. n. zebrata Wolff, 1929 E. n. gelidatoides Warnecke, 1951 E. innotata (Hufnagel, 1767). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/83 E. ochridata Schütze & Pinker, 1968 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/84 E. unedonata Mabille, 1868. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/85 * E. parallelaria Bohatsch, 1893 [Turkey, Levant] 326 E. chalikophila Wehrli, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/86 The graphata species-group * E. brunneata Staudinger, 1900 [Turkey] E. graphata (Treitschke, 1828). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/87 E. g. setaceata Dietze, 1903 E. g. sproengertsi Dietze, 1910 E. g. olympica Tuleschkov, 1951 E. g. hesperia Wehrli, 1926 E. g. carpetanensis Redondo & Gaston, 2009 327 * E. albofasciata Staudinger, 1879 [Turkey] 328 * E. antigraphata Mironov, 2013 [Turkey] * E. perigraphata Mironov, 2013 [Turkey] E. riparia Herrich-Schäffer, 1851 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/88 E. furcata Staudinger, 1883 329/���������������������������������������������������������������������������������� 6/197 * E. elissa Dietze, 1910 [Tunisia] * E. farinosa Dietze, 1913 [Algeria]

325 Leraut (2009) validates the taxon angusta Prout, 1938 at subspecies rank (British Isles and Atlantic coast of France), based on different wing colour. In GME4 this taxon was retained as synonym because of clinous character transitions. 326 Raised to species rank in Mironov (2013), described by Bohatsch (1893) as var. of E. unedonata. 327 Recently described for populations from central Spain. 328 Raised to species rank in Mironov (2013). 329 Recently discovered in Greece, see information in this volume.

867

E. inconspicuata Bohatsch, 1893 (= thurnerata Schütze, 1958) 330/�������������������������������������������������������������������������� 4/89 * E. pseudoscriptaria Rothschild, 1914 [Morocco] 331 * E. novata Dietze, 1904 [Turkey] 332 E. gemellata Herrich-Schäffer, 1861. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/90 E. gypsophilata Skou, Mironov & Rietz, 2017 333 /������������������������������������������������������ 6/198 E. mystica Dietze, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/91 (= wehrlii Wagner, 1931 [Turkey]) 334 E. cooptata Dietze, 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/92 * E. subsequaria Herrich-Schäffer, 1852 [Turkey] (= dubiosata Wagner, 1929) 335 * E. tesserata Brandt, 1938 [Jordan, Iran] * E. convallata Brandt, 1938 [Turkey, Iran, Afghanistan] 336 (= terricolor Vojnits, 1988) * E. siata Brandt, 1938 [Turkey, Iran] 337 The breviculata species-group E. spissilineata (Metzner, 1846) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cuculliaria (Rebel, 1901). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. unitaria Herrich-Schäffer, 1852. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. breviculata (Donzel, 1837). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. cerussaria (Lederer, 1855). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

4/93 4/94 4/95 4/96 4/97

The irriguata species-group E. irriguata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/98 E. i. eriguata Staudinger, 1871 E. i. staudingeri Bohatsch, 1893 * E. i. mauretanica Dietze, 1910 [Morocco] E. antalica Mironov, 2001. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/99 * E. marasa Wehrli, 1932 [Turkey] E. indigata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/100 * E. i. hamamata Pinker, 1976 [Turkey] E. conterminata (Lienig & Zeller, 1846). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/101 E. pindosata Weidlich, 2008 338 /�������������������������������������������������������������������������������� 6/199 The distinctaria species-group E. distinctaria Herrich-Schäffer, 1848. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/102 E. senorita Mironov, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/103

330 Synonymized in Mironov (2013), see information in this volume. 331 Validated at species rank by Leraut (2009), stating potential synonymy with E. thurnerata, awaiting thorough analysis and confirmation. 332 This taxon may be a synonym of Eupithecia gemellata (Mironov pers. comm.) or the eastern sister species in this species complex. 333 Recently described from Spain, see information in this volume. 334 Synonym according to Mironov (2013). 335 Synonym according to Mironov (2013). 336 Revised and recorded for Turkey in Mironov (2013). 337 Recorded for Turkey in Mironov (2013). 338 Recently described from Greece, see information in this volume.

868

The centaureata species-group E. extraversaria Herrich-Schäffer, 1852 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/104 E. centaureata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/105 E. limbata Staudinger, 1879. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/106 E. l. tomillata (Chrétien, 1904) (= irritaria Staudinger, 1892 [Turkey]) 339 * E. schwingenschussi Zerny, 1934 [Morocco] * E. deverrata Dietze, 1910 [Tunisia] * E. lecerfi Prout, 1928 [Morocco, Algeria] E. mandarinca Mironov, 2001 340/������������������������������������������������������������������������������ 4/107 * E. desertorum Dietze, 1910 [Algeria, Tunisia] The despectaria species-group E. mekrana Brandt, 1941. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/108 * E. tenellata Dietze, 1906 [Morocco to Saudi Arabia] * E. despectaria Lederer, 1853 [Turkey, Central Asia] 341 The variostrigata species-group E. variostrigata Alphéraky, 1876. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/109 (= albosparsata Joannis, 1891 [Turkey]) 342 * E. falkneri Vojnits, 1978 [Algeria] The nevadata species-group E. insigniata (Hübner, 1790). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/110 The gueneata species-group E. trisignaria Herrich-Schäffer, 1848. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. gueneata Millière, 1862. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= separata Staudinger, 1879 [Turkey]) 343 E. g. plantei Herbulot, 1981 [Morocco] * E. syriacata Staudinger, 1879 [Turkey to Levant] E. gratiosata Herrich-Schäffer, 1861. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. scalptata Christoph, 1885. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

4/111 4/112

4/113 4/114

The cretaceata species-group E. veratraria Herrich-Schäffer, 1848. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/115 E. v. eynensata de Graslin, 1863 E. v. arctica Viidalepp, 1974 E. cretaceata Packard, 1874 E. c. fenestrata Millière, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/116 * E. subfenestrata Staudinger, 1892 [Caucasus, Trans­caucasus] The satyrata species-group E. praealta Wehrli, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/117 E. intricata (Zetterstedt, 1839). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/118 339 Synonym according to Scoble (1999) and Mironov (2013). 340 In GME4 (Mironov 2003) erroneously synonymized with E. nachadira Brandt, 1941 (described from south-eastern Iran). Revised in this volume (see text; Mironov, pers.comm.). 341 Recorded for Turkey in Mironov (2013). 342 Validated at species rank in Scoble (1999), synonymized in Mironov (2013). 343 Validated at species rank in Scoble (1999), synonymized in Mironov (2013).

869

E. i. hibernica Mere, 1964 E. i. millieraria Wnukowsky, 1929 (= pfeifferi Wehrli, 1929 [Caucasus, Transcaucasus, Turkey]) 344 * E. tribunaria Herrich-Schäffer, 1852 [Great Caucasus] E. satyrata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/119 E. s. curzoni Gregson, 1884 E. cauchiata (Duponchel, 1830). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/120 E. pernotata Guenée, 1858. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/121 E. p. enictata Pellmyr & Mikkola, 1984 E. extensaria (Freyer, 1845). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/122 E. e. occidua Prout, 1914 E. absinthiata (Clerck, 1759). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/123 E. goossensiata Mabille, 1869 345/������������������������������������������������������������������������������ 6/200 E. expallidata Doubleday, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/124 E. valerianata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/125 * E. arenbergeri Pinker, 1976 [western Caucasus, Turkey] The russeliata species-group E. assimilata Doubleday, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. vulgata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. v. cyrneata Schawerda, 1933 E. v. clarensis Huggins, 1962 E. biornata Christoph, 1867. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. sutiliata Christoph, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. immundata (Lienig & Zeller, 1846). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

4/126 4/127 4/128 4/129 4/130

The addictata species-group E. thalictrata (Püngeler, 1902). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/131 E. addictata Dietze, 1908 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/132 The druentiata species-group E. exiguata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. druentiata Dietze, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. moecha Dietze, 1904. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. denotata (Hübner, 1813) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. d. hellenata Schütze, 1959 (= pseudocastigata Pinker, 1976 [Turkey]) 346 The millefoliata species-group E. weissi Prout, 1938. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. pauxillaria Boisduval, 1840 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. santolinata Mabille, 1871. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. millefoliata Rössler, 1866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. spadiceata Zerny, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

4/133 4/134 4/135 4/136

4/137 4/138 4/139 4/140 4/141

The semigraphata species-group E. icterata (de Villers, 1789) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4/142 344 Downgraded to synonymy in Mironov (2013). Possible validation at subspecies rank requiring further study. 345 Raised to species rank in Hausmann et al. (2011), see information in this volume. 346 Synonymized in Mironov (2013).

870

E. succenturiata (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. s. artvinata Mironov, 2001 [Turkey] 347 * E. tarfata (Lucas, 1907) [Algeria] * E. marginata Staudinger, 1892 [Cyprus, Turkey to Caucasus] E. semigraphata Bruand, 1851. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. impurata (Hübner, 1813). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. poecilata Püngeler, 1888. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. fuscicostata Christoph, 1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. denticulata (Treitschke, 1828) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. d. anatolicata Pinker, 1976 [Turkey] 348 E. subumbrata (Denis & Schiffermüller, 1775). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. s. lacteolata Dietze, 1906 [Turkey, Armenia] 349 E. orphnata Petersen, 1909. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . E. subfuscata (Haworth, 1809). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * E. gelbrechti Mironov, 2013 [Turkey] * E. niveata Mironov, 2013 [Turkey]

4/143

4/144 4/145 4/146 4/147 4/148 4/149 4/150 4/151

Position in species-group uncertain * E. arenitincta Prout, 1938 [Algeria] * E. glaisi Lucas, 1937 [Algeria] * E. insignifica Rothschild, 1914 [Algeria] * E. ogilviata (Warren, 1905) [Açores] * E. yathomi Hausmann, 1991 [Jordan] * E. atlanticata Pinker, 1971 [Madeira] * E. dearmata Dietze, 1904 [Turkey] 350 * E. bastelbergeri Dietze, 1910 [Turkey] 351 * E. mima Mironov, 1989 [Turkey] 352 * E. ochrovittata Christoph, 1887 [Turkey, Transcaucasus] 353 * E. terrenata Dietze, 1904 [Turkey] 354

347 cf. Mironov (2013). 348 cf. Mironov (2013). 349 Downgraded from species rank (Scoble 1999; Mironov 2003) to subspecies rank in Mironov (2013). 350 cf. Mironov (2013), transferred from genus Gymnoscelis. 351 cf. Mironov (2013). 352 cf. Mironov (2013). 353 cf. Mironov (2013). 354 cf. Mironov (2013).

871

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Index to scientific names of the Systematic Account, Vol. 6 The index gives page references to the main treatment. Numbers in semibold refer to the species numbers and figures. abruptaria – 230, 70 abstersaria – 420 achyra – 302, 98 Adactylotis – 448 adriatica – 335 Aethalura – 457 Afriberina – 386 Agriopis – 315 albarinata – 75 albarracina – 528 albosignata – 345 Alcis – 436 alexandrinus – 304 algeriensis – 345 algiricaria – 372, 128 almasa – 222 almensis – 366 alpicolaria – 272 alpina – 290, 93 alpinaria – 509 alpinata – 129, 35 alpmaritima – 147, 43 alticolaria – 148, 44 amani – 403 ambiguata – 103, 26 ambustaria – 408, 141 Amorphogynia – 275 amygdalaria – 366, 125 andalusiaria – 357, 121 andereggaria – 192, 59 andrea – 548 androgynus – 63 ankeraria – 310, 101 annegreteae – 242, 76 Apocheima – 259 Apochima – 53 aragona – 372 aragonensis – 70 aragonis – 432, 150 arenaria – 273 arenaria – 337, 114 argillacearia – 67 Arichanna – 469 Ascotis – 460 assoi – 88, 20

Athroolopha – 341 atlantica – 288 atlanticaria – 341 atlanticaria – 345 atlanticaria – 360 atlanticaria – 375, 129 atomaria – 272, 86 aurantiaria – 320, 106 avilarius – 70 bajaria – 318, 105 baldensis – 154, 46 barbuti – 526, 185 barrassoi – 543, 194 bastelbergeri – 439 bastelicaria – 396, 135 Bellachima – 53 bellieri – 68 belzebuth – 144, 41 benestriata – 122 bentelii – 138, 39 berenicidaria – 234, 72 bernieri – 472 bervaensis – 310 beschkovi – 314, 103 betularia – 303, 99 Biston – 297 Bistonini – 254 bistortata – 466 Boarmiini – 253 bowesi – 463 Brachyglossina – 533 britannica – 288 brunnearia – 400, 137 budashkini – 322, 107 budensis – 122 Bupalini – 254 Bupalus – 471 bureschi – 136 buxicolaria – 420 byssata – 538 bytinskii – 479, 484 caelibaria – 183, 56 Calamodes – 326 calceata – 67 caliginearia – 335, 113

canaliculata – 132, 36 carniolica – 290 carpathica – 142 carpophilata – 545, 195 Cassymini – 40 castiliaria – 509 cataloniae – 277 catharia – 370 cebennaria – 509 certhiatus – 125 chalybaeus – 149 Charissa – 66, 67 Chemerina – 334 Chondrosoma – 56 chrysitaria – 343, 117 cinctaria – 463, 163 cineraria – 481 Cleora – 462 Cleorini – 253 Cleorodes – 244 cleui – 321 cleui – 490 Cnestrognophos – 82 codetaria – 486 coggii – 318 cognataria – 303 Colotoini – 52 confertata – 75 consonaria – 330, 111 constantina – 521, 183 contaminaria – 450, 158 Contropis – 474 coracina – 135, 38 correptaria – 430, 149 corsica – 91, 21 corsicaria – 417 Costignophos – 70 cottei – 259 crenulata – 70, 12 crepuscularia – 466, 164 cretacaria – 235 Crocota – 212 cuencaensis – 363 culminaria – 513, 179 cyrnea – 493, 172

903

dagestanensis – 38 dalmata – 230 dalmatina – 367, 126 danieli – 266, 84 Dasypteroma – 251 davidi – 529 declinans – 312, 102 defloraria – 417 defoliaria – 307, 100 Deileptenia – 444 deversata – 439, 153 dianaria – 460 Dicrognophos – 41 difficilis – 77, 15 dilucescens – 400 dilucidaria – 196, 60 dinicola – 240 diniensis – 276 diniensis – 490 dioszeghyi – 122 dioszeghyi – 135 distinctaria – 244 dodoneata – 548 dognini – 206 dolomitica – 169, 53 donovani – 536, 191 drenowskii – 534, 190 dresnayi – 237 druentiaria – 428 dubiosa – 237 dubiosa – 546, 196 dumetata – 30, 1 Dyscia – 557 Dysgnophos – 77 ebusaria – 427, 147 Ecleora – 383 Ectropis – 465 Ekboarmia – 374 Elophos – 181 Ematurga – 272 Entephria – 538 Erannini – 254 Erannis – 307 erebaria – 410, 142 erubescens – 533 espagnolus – 472 espaniola – 504, 176 estachyi – 215 estrellae – 357 Euchrognophos – 87 Eumannia – 476, 505 Eupithecia – 545

904

Eurranthis – 345 exilaria – 533 extensaria – 407 Fagivorina – 336 fallentaria – 428 faroulti – 36 fascinataria – 378 faucium – 148 fidelensis – 233 fiduciaria – 56, 8 fingalata – 481 flabellaria – 54, 7 florentina – 295, 95 fortunaria – 397, 136 francothuringiaca – 122 fumosarius – 303 furcata – 548, 197 furvata – 61, 9 galliberia – 420 gedrensis – 148 gelbrechti – 531, 188 gerini – 432 gerini – 486 gesticularia – 448, 157 gigantaria – 372 Glacies – 127 glaucinaria – 116, 31 Gnophini – 58 Gnophos – 61 goossensiata – 553, 200 govii – 557, 20 gracilaria – 514, 180 gracilis – 375 graecaria – 125, 34 graecarius – 293, 94 granataria – 246, 78 grisearia – 269 grisolaria – 474, 167 grupchei – 279 gulsensis – 123 gypsophilata – 550, 198 hammamrirha – 388 hanoviensis – 276 Harrisonodes – 275 harterti – 237, 73 herbuloti – 103 herbuloti – 405 herrerai – 380 hibernica – 31 hirtaria – 276, 87 hispanica – 360, 123 hispidaria – 259, 82

hoefneri – 189 holli – 375 hubertcleui – 509 humiliata – 529 Hypomecis – 265 Hyposcotis – 66 ichnusaria – 417 Idaea- 526 idrisarius – 102 iliaria – 272 ilicaria – 434, 151 impectinata – 75 incerta – 244 incisaria – 528 inconspicuata – 550 innuptaria – 166, 51 insignata – 524 intermedia – 122 inversarius – 279 isabellae – 283, 90 isoscelata – 272 istriana – 276 istrianus – 293 italica – 295 italohelveticus – 73, 13 Ithysia – 516 jacobiaria – 434 japygiaria – 232, 71 jaspidaria – 244 jubata – 442, 154 juracolaria – 540, 193 kabylaria – 341 Kemtrognophos – 102 kerstinae – 269 khenchelae – 424 kindermannaria – 339, 115 kovacsi – 75 Kresnaia – 313 Kuchleria – 524 kusdasi – 142 lafauryata – 68 lanaria – 257, 81 lappona – 135 lapponaria – 281, 89 Larerannis – 332 laricaria – 466 lariciaria – 320 latimargo – 341 lepraria – 497, 173 Leucobrephos – 519 leucophaearia – 316, 104 lhommaria – 489, 171

lichenaria – 244, 77 lobaria – 529, 187 lutescentaria – 63 Lycia – 275 Macariini – 49 macguffini – 80 maghrebica – 434 magyarica – 269 malatyaca – 366 mandarinca – 553 manicaria – 533 margaritata – 35, 2 marginaria – 324, 108 marmararia – 54 marsicaria – 63 maura – 230 mauretania – 405 Megalycinia – 248 Megametopon – 474 melaleucaria – 490 melanaria – 328, 110 melanaria – 470, 165 melloulica – 480, 484 menadiara – 524, 184 mendicaria – 201 Menophra – 228 meridionalis – 298 Microbiston – 257 middendorfii – 520, 182 millefoliata – 556 mimeuri – 424 miniaria – 382, 131 minorasiaticus – 125 minuta – 272 modestaria – 474 mongolicum – 520 montserrata – 420 mucidaria – 99, 25 muraria – 436 mutilata – 83, 17 Myinodes – 521 myrtillata – 63 nachadira – 553 necessaria – 279, 88 necopinatus – 189 Neognophina – 115 nigrolineata – 480, 484 nivea – 63 niveata – 220, 67 niveiscythes – 556, 201 noricana – 142, 40 notarioi – 359, 122

Nothocasis – 538 nubilata – 75 nuragica – 485, 169 Nychiodes – 347 nycthemeraria – 240, 75 Oar – 534 obfuscata – 63, 10 obscuraria – 352, 119 obscuraria – 506 obscurata – 67, 11 occidentaria – 428 occitanaria – 327, 109 Odontognophos – 29 odrinarius -293 oliveirata – 405 omalensis – 422, 145 onustaria – 109, 28 operaria – 189, 58 ophthalmicata – 103 oppositaria – 506, 177 oranaria – 508, 178 orientalis – 341 orientaria – 272 orientis – 260 Ortaliella – 49 orthogrammaria – 333, 112 osthelderi – 363 Ourapteryx – 556 pallidaria – 400 pallidata – 324 Paraboarmia – 446 Paradarisa – 330 Parectropis – 455 parenzani – 400, 408 Parietaria – 195 peletieraria – 217, 65 peloponnesiaria – 84, 18 pennigeraria – 341, 116 pentheri – 86, 19 Peribatodes – 412 Perizoma – 540 perlinii – 138 perspersata – 49, 6 perumbraria – 425 perversaria – 420, 144 petrina – 86 pfeifferi – 119, 32 Phaselia – 369 Phaseliini – 254 Phigalia – 262 Phthonandria – 225 Phyllometra – 476, 512

pilosaria – 263, 83 pindosata – 551, 199 piniaria – 472, 166 plumaria – 403, 138 plummistaria – 345, 118 pomonaria – 285, 91 popovi – 260 postdentaria – 410 powelli – 252 powelli – 433 praerecta – 480, 484 pratana – 534, 189 pravata – 517, 181 predotae – 107, 27 pretiosaria – 526 propinquaria – 222 Prosopolophini – 52 Pseudocoremia – 451 pseudonoricana – 135 pseudotinctaria – 219, 66 Psodos – 127 psyloritaria – 500, 174 pullaria – 67 pullata – 75, 14 pullularia – 103 punctinalis – 269, 85 punctinervis – 335 punctipes – 538 punctulata – 458, 161 pusztae – 277 pyrenaea – 157 pyrenaearia – 405, 139 pyrenaica – 146, 42 pyrenaica – 70 quadrifaria – 157, 48 ragusaria – 354, 120 reisseri – 540 relicta – 372 remmi – 112, 29 repandata – 436, 152 retyezatensis – 138 rhomboidaria – 417, 143 Rhopalognophos – 115 ribeata – 444, 155 roboraria – 266, 84 rosariae – 538, 192 rosenbaueri – 300, 97 rossica – 288 rothschildi – 98 saerdabensis – 417 saerdabica – 363 sagnesi – 380, 130

905

salemae – 391, 134 Sardocyrnia – 395 sartata – 41, 4 scandinaviaria – 400 schwingenschussi – 132 Sciadia – 159 Scopula – 534 scotica – 281 secundaria – 428, 148 selenaria – 460, 162 Selidosema – 399 sepiaria – 481, 168 septaria – 165, 50 serotinaria – 206, 63 serotinoides – 207 serraria – 109 serraria – 249, 79 serrularia – 370, 127 sheljuzhkoi – 103 shurensis – 370 siciliana – 44, 5 sicula – 487, 170 similaria – 455, 160 slovenica – 168, 52 sociaria – 222, 68 sodorensium – 436 solieraria – 384, 132 Spartopteryx – 339 spitzi – 156, 47 sproengertsi – 199, 61 spurcaria – 183 staudingeri – 92, 22 strataria – 298, 96 strictaria – 226, 69

906

strobioni – 284 suavis – 453, 159 subflavaria – 420 sublutearia – 417 submelanaria – 327 subscudularia – 328 subtaurica – 94, 23 sudetica – 157 supinaria – 122, 33 Synopsia – 222 syriaca – 310 syriaca – 363 syriacaria – 400 syrirana – 424 syritaurica – 417 taeniolaria – 407, 140 tagana – 474 taurusica – 161 teberdensis – 78 tenebraria – 161, 49 tenietaria – 388, 133 Tephronia – 476, 479 teriolensis – 307 teriolica – 481 terraria – 390 textaria – 528, 186 thaumasia – 252, 80 theophilaria – 501, 175 thuriferaria – 239, 74 thurnerata – 550 tinctaria – 214, 64 tonnara – 493 torrevinagrensis – 361 transcaspia – 363

transiens – 135 Trilobignophos – 125 truschi – 533 trypanaria – 234 tundranoides – 135 turfosaria – 80, 16 tyronensis – 400 umbraria – 424, 146 undulosa – 237 ungemachi – 448 unicoloraria – 177, 55 unitaria – 222 uralaria – 276 urmensis – 78 variegata – 96, 24 vaucheri – 424 viertlii – 446, 156 vintejouxi – 215 vittaria – 201, 62 waltheri – 363, 124 wehrlii – 151, 45 werneri – 134, 37 wettsteini – 556 wockearia – 161 Yezognophos – 194 zacharia – 38, 3 zehrounensis – 318 zeitunaria – 114, 30 zelleraria – 172, 54 Zernyia – 246 zetterstedtaria – 273 zirbitzensis – 187, 57 zonaria – 288, 92