Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae) [1st ed.] 9783030361730, 9783030361747

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Jan Klimaszewski · E. Richard Hoebeke  Benoit Godin · Anthony Davies · Kayla I. Perry  Caroline Bourdon · Neville Winchester

Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae)

Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae)

Jan Klimaszewski • E. Richard Hoebeke • Benoit Godin • Anthony Davies • Kayla I. Perry • Caroline Bourdon • Neville Winchester

Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae)

Jan Klimaszewski Natural Resources Canada Laurentian Forestry Centre Québec, QC, Canada Benoit Godin Whitehorse, YT, Canada Kayla I. Perry Department of Entomology The Ohio State University Columbus, OH, USA Neville Winchester Department of Biology University of Victoria Victoria, BC, Canada

E. Richard Hoebeke Georgia Museum of Natural History and Department of Entomology University of Georgia Athens, GA, USA Anthony Davies Agriculture and Agri-Food Canada Canadian National Collection of Insects, Arachnids and Nematodes Ottawa, ON, Canada Caroline Bourdon Natural Resources Canada Laurentian Forestry Centre Québec, QC, Canada

ISBN 978-3-030-36173-0 ISBN 978-3-030-36174-7 https://doi.org/10.1007/978-3-030-36174-7

(eBook)

# Her Majesty the Queen in Right of Canada 2020 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

We are honored to dedicate this body of work to our colleague Steve Ashe for his many authoritative contributions to aleocharine systematics through his impressive scientific publication record and his love of little rove beetles. His unwavering commitment to promoting and strengthening our knowledge of the Aleocharinae is unsurpassed.

Preface

Worldwide, aleocharine beetles are among the most poorly known and difficult-to-identify groups of Coleoptera. Here, we present the first comprehensive synopsis of all known valid aleocharine rove beetle species (Coleoptera, Staphylinidae) from British Columbia, Canada. Two hundred twentyseven confirmed and valid species from the province are presented and discussed here. This taxonomic account includes one new BC genus (not including new genus from CA not recorded from BC), 15 new species, 16 new generic records, and 37 (excluding new species) new provincial and 6 state records, in 79 genera and 14 tribes. For each species, the illustrations include color habitus and genital diagnostic structures of both sexes. Tribes and subtribes are arranged in phylogenetic order as it is currently recognized, and genera and subgenera are listed alphabetically within each tribe or subtribe. Species are listed alphabetically or in species groups to better reflect their relationships. Species distribution is listed by provinces and territories in Canada and states in the United States, and the geographic origin of each species is categorized as native, Holarctic, adventive, or undetermined (either adventive or Holarctic). Collection and habitat data are presented for each species, including collecting period and collecting methods. A faunal analysis and discussion on BC aleocharine in a broader context of North America is provided. Two lists of Canadian and BC species with their currently known distribution in North America are presented at the end of the book (Tables A.1 and A.2 in Appendix). TAXONOMIC SYNOPSIS. “Salinamexus” giulianii Moore was incorrectly included in Bryobiota and is here assigned to a newly erected genus, Ashella Klimaszewski, gen. n., of the tribe Liparocephalini Fenyes. Ashella is, so far, not recorded from BC. Bryobiota Casey is redefined based on morphology of the type species. We record fifteen new species: Atheta copleyi Klimaszewski, sp. n.; Atheta godini Klimaszewski, sp. n.; Atheta wheelerae Klimaszewski, sp. n.; Boreostiba pseudolaticollis Klimaszewski and Godin, sp. n.; Atheta winchesteri Klimaszewski, sp. n.; Calodera bennetti Klimaszewski, sp. n.; Dinaraea inexpectata Klimaszewski, sp. n.; Geostiba horwoodae Klimaszewski and Godin, sp. n.; Gnypeta baranowskii Klimaszewski, sp. n.; Leptusomorpha claudiae Klimaszewski, sp. n.; Liogluta scudderi Klimaszewski, sp. n.; Neothetalia robergei Klimaszewski, sp. n., Philhygra charlottae Klimaszewski, sp. n.; Philhygra terrivaga Klimaszewski, sp. n.; Stictalia kranabetteri Klimaszewski and Godin, sp. n. vii

viii

One new genus is erected, Leptusomorpha Klimaszewski and Hoebeke of the tribe Homalotini. Thirty-seven new BC-specific provincial records are noted (excluding new species): Amischa analis (Gravenhorst), Atheta alesi Klimaszewski and Brunke, Atheta capsularis Klimaszewski, Atheta brunswickensis Klimaszewski, Atheta lucifera Bernhauer, Atheta munsteri Bernhauer, Atheta pseudoklagesi Klimaszewski and Webster, Atheta pseudometlakatlana Klimaszewski and Godin, Atheta ripariides Newton, Atheta terranovae Klimaszewski and Langor, Boreostiba parvipennis (Bernhauer), Dinaraea angustula (Gyllenhal), Dinaraea subdepressa (Bernhauer), Dochmonota rudiventris (Eppelsheimer), Blepharhymenus illectus Casey, Brachyusa helenae (Casey), Bryothinusa catalinae Casey, Liogluta trapezicollis Lohse, Lypoglossa franclemonti Hoebeke, Mocyta discreta (Casey), Nehemitropia lividipennis (Mannerheim), Philhygra terrestris Klimaszewski and Godin, Schistoglossa hampshirensis Klimaszewski, Strigota ambigua (Erichson), Strigota obscurata Klimaszewski and Brunke, Gymnusa grandiceps Casey, Oligota parva Kraatz, Neoisoglossa agnita (Casey), Ocyusa canadensis Lohse, Oxypoda canadensis Klimaszewski, Oxypoda convergens Casey, Oxypoda irrasa Mäklin, Oxypoda orbicollis Casey, Parocalea pseudobaicalica Lohse, Phloeopora arctica Lohse, Phloeopora canadensis Klimaszewski and Langor, and Phloeopora oregona Casey. In addition, sixteen new generic provincial records are (including new genus): Amischa C.G. Thomson, Blepharhymenus Solier, Bryothinusa Casey, Calodera Mannerheim, Dinaraea C.G. Thomson, Dochmonota C.G. Thomson, Geostiba C.G. Thomson, Leptusomorpha Klimaszewski and Hoebeke, Meotica Mulsant and Rey, Nehemitropia Lohse, Neoisoglossa Casey, Ocyusa Kratz, Oligota Mannerheim, Parocalea Bernhauer, Phloeopora C.G. Thomson, and Strigota Casey. We note, three New Synonyms (first name being valid): Atheta (Datomicra) celata (Erichson, 1837) ¼ Datomicra wrangleri Casey, 1910 ¼ Pseudota nanulina Casey, 1911; Stictalia brevicornis Casey, 1906 ¼ S. arcuata Casey, 1906; S. californica (Casey, 1885) ¼ S. densicollis Casey, 1906. Twelve New Lectotypes are designated: Acrimea acerba Casey, 1911 (¼Aleochara (Tinotus) acerba (Casey)); Atheta holmbergi Bernhauer (Atheta (Microdota) holmbergi Bernhauer); Atheta munsteri Bernhauer (¼Atheta (Dimetrota) munsteri Bernhauer); Atheta relicata Casey (¼Atheta (Lamiota) relicata Casey); Datomicra wrangleri Casey (¼Atheta (Datomicra) celata (Erichson)); Ousipalia pacifica Casey; Stictalia arcuata Casey (¼S. brevicornis Casey); Stictalia carlottae Casey; Stictalia densicollis Casey (¼S. californica (Casey)); Trichiusa columbica Casey; Pseudota nanulina Casey (¼Atheta (Datomicra) celata (Erichson)); Datomicra surgens Casey (¼Atheta (Microdota) surgens (Casey)). Nomen nudum: Stictalia notata (Mäklin, 1852), type lost at ZMH. Species with unconfirmed status are: two Palearctic species, Atheta subrugosa (Kiessenwetter) and A. basicornis (Mulsant and Rey), and records in BC were not confirmed in this study and most likely represent misidentifications.

Preface

Preface

ix

These two species are illustrated here for further reference. Brachyusa americana (Fenyes), known only from damaged male holotype, was not available for study but was included in this book with illustrations of the median lobe of aedeagus and paramere, provided by Seevers (1978). Québec, QC, Canada Athens, GA, USA Whitehorse, YT, Canada Ottawa, ON, Canada Columbus, OH, USA Québec, QC, Canada Victoria, BC, Canada

Jan Klimaszewski E. Richard Hoebeke Benoit Godin Anthony Davies Kayla I. Perry Caroline Bourdon Neville Winchester

Acknowledgements

This book is dedicated to Steve Ashe, and other prominent entomologists mainly from British Columbia, as a mark of gratitude for their entomological contributions to North America and the province. We named several new species for BC entomologists, as indicated in the text. We received photographs of several prominent American entomologists from Lee Herman (AMNH). Our colleague Reginald Webster (Fredericton, New Brunswick) provided most of collection and habitat data for transcontinental species recorded from New Brunswick. We appreciate taxonomic advice from Al Newton and his generosity in sharing his knowledge (FMNH). We thank Kee-Jeong Ahn, Chungnam National University, Daejeon, Republic of Korea, who provided taxonomic advice on coastal aleocharines and some images and drawings. The first author would like to thank his Director General Dominic St-Pierre and Research Director Elizabeth Gauthier for supporting this project under the Coleoptera Collection program. The first author thanks his wife, Patricia Corvera, for her support, encouragement, and participation in discovering nature landmarks on Vancouver Island. Diane Paquet (LFC) helped us with formatting manuscripts and resolving many technical issues. Credit for images goes mainly to Sylvain Roberge (LFC), and Caroline Bourdon (LFC). The third author thanks Claudia Copley for providing many uncatalogued specimens from the Royal British Columbia Museum collection, as well as Leah Ramsey and David Fraser for their hospitality and access to their property for collecting, and my wife Denise Horwood for participating in many collection surveys. The following curators and individuals provided specimens including types and ordinary specimens, including those from Europe, and we gratefully acknowledge their assistance: V. Assing (VAC), C.C. Grinter (CAS), C. Copley and R. Bennett (RBCM), C. Maier (FMNH), J. Mattila (ZMH), K. Needham (UBC), D. Sikes (UAM), A. Solodovnikov (UCC), and F. Shockley (USNM). Funding for this project was provided by Natural Resources Canada. Neville Winchester would like to thank Brenda Costanzo for her input into the chapter on unique habitats in BC and the use of several photographs.

xi

Contents

1

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

1 3

2

Prominent Forebearers of Modern Aleocharine Systematics in North America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

5 8

Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia . . . . . . . . . . . . . . . . . . . . Pre-Casey Era (1925) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . .

9 10 10 11 14

. . . . . . .

17 17 17 17 18 18 19

. . . .

21 22 22 42

5

British Columbia: Hotspot of Canadian Biodiversity . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

45 52

6

Faunal Analysis and Discussion . . . . . . . . . . . . . . . . . . . . . . Dominant Aleocharinae Tribes in British Columbia . . . . . . . . . Adventive Species of Aleocharinae . . . . . . . . . . . . . . . . . . . . . Predictions for Aleocharinae Diversity in British Columbia . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

53 53 57 58 61

3

4

Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Format . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Institution Codes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diagnostic Features . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . External Body Structures Important for Identification . . . . . . . . Genital Structures Important for Identification . . . . . . . . . . . . . Microdissections and Preparation of Genital Structures for Examination and Diagnostics . . . . . . . . . . . . . . . . . . . . . . . . . Image Production . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Collection Methods and Habitats . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . .

xiii

xiv

7

Contents

List of Recorded Species of British Columbian Aleocharinae and Their Composition . . . . . . . . . . . . . . . . . . . . . . . . . . . . Family STAPHYLINIDAE Latreille, 1802 (Rove beetles) . . . . Subfamily ALEOCHARINAE Fleming, 1821 . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

63 63 63 71

Key to Tribes of Aleocharine Occurring in British Columbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Reference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

75 87

9

Tribe Gymnusini Heer, 1839 . . . . . . . . . . . . . . . . . . . . . . . . Tribe Gymnusini Heer, 1839 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Gymnusa Gravenhorst, 1806 . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

89 89 89 96

10

Tribe Aleocharini Fleming, 1821 . . . . . . . . . . . . . . . . . . . . . Tribe Aleocharini Fleming, 1821 . . . . . . . . . . . . . . . . . . . . . . Genus Aleochara Gravenhorst, 1802 . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. 97 . 97 . 97 . 139

11

Tribe Oxypodini C.G. Thomson, 1859 . . . . . . . . . . . . . . . . . Tribe Oxypodini C.G. Thomson, 1859 . . . . . . . . . . . . . . . . . . Genus Blepharhymenus Solier, 1849 . . . . . . . . . . . . . . . . . . . . Genus Meotica Mulsant and Rey, 1873 . . . . . . . . . . . . . . . . . . Genus Crataraea C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . Genus Calodera Mannerheim, 1830 . . . . . . . . . . . . . . . . . . . . Genus Devia Blackwelder, 1952 . . . . . . . . . . . . . . . . . . . . . . . Genus Gnathusa Fenyes, 1910 . . . . . . . . . . . . . . . . . . . . . . . . Genus Mniusa Mulsant and Rey, 1875 . . . . . . . . . . . . . . . . . . Genus Ocyusa Kraatz, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Oxypoda Mannerheim, 1830 . . . . . . . . . . . . . . . . . . . . . Ocalea Group of Genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Alfocalea Klimaszewski, 2004 . . . . . . . . . . . . . . . . . . . Genus Betocalea Klimaszewski, 2004 . . . . . . . . . . . . . . . . . . . Genus Megocalea Klimaszewski, 2004 . . . . . . . . . . . . . . . . . . Genus Metocalea Klimaszewski, 2004 . . . . . . . . . . . . . . . . . . Genus Neoisoglossa Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . Genus Neothetalia Klimaszewski, 2004 . . . . . . . . . . . . . . . . . . Genus Ocalea Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Parocalea Bernhauer, 1902 . . . . . . . . . . . . . . . . . . . . . Genus Phloeopora Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . .

141 141 143 144 145 146 147 148 150 151 152 168 169 170 171 172 173 175 180 181 183 232

12

Tribe Tachyusini C. G. Thomson, 1859 . . . . . . . . . . . . . . . . Tribe Tachyusini C.G. Thomson, 1859 . . . . . . . . . . . . . . . . . . Genus Brachyusa Mulsant and Rey, 1873 . . . . . . . . . . . . . . . . Genus Gnypeta C.G. Thomson, 1859 . . . . . . . . . . . . . . . . . . . Genus Paradilacra Bernhauer, 1909 . . . . . . . . . . . . . . . . . . . . Genus Tachyusa Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . .

235 235 235 237 243 244 259

8

Contents

xv

13

Tribe Hypocyphtini Laporte, 1835 . . . . . . . . . . . . . . . . . . . . Tribe Hypocyphtini Laporte, 1835 . . . . . . . . . . . . . . . . . . . . . Genus Cypha Leach, 1819 . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Holobus Solier, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Oligota Mannerheim, 1830 . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . .

261 261 261 262 263 268

14

Tribe Myllaenini Ganglbauer, 1895 . . . . . . . . . . . . . . . . . . . Tribe Myllaenini Ganglbauer, 1895 . . . . . . . . . . . . . . . . . . . . . Genus Myllaena Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

269 269 269 277

15

Tribe Diglottini Jakobson, 1909 . . . . . . . . . . . . . . . . . . . . . . Tribe Diglottini Jakobson, 1909 . . . . . . . . . . . . . . . . . . . . . . . Genus Bryothinusa Casey, 1904 . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

279 279 279 282

16

Tribe Liparocephalini Fenyes, 1918 . . . . . . . . . . . . . . . . . . . Tribe Liparocephalini Fenyes, 1918 . . . . . . . . . . . . . . . . . . . . Genus Amblopusa Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . . . Genus Paramblopusa Ahn and Ashe, 1996a . . . . . . . . . . . . . . Genus Diaulota Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Liparocephalus Mäklin, 1853 . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . .

283 283 284 285 286 287 294

17

Tribe Autaliini C. G. Thomson, 1859 . . . . . . . . . . . . . . . . . . Tribe Autaliini C. G. Thomson, 1859 . . . . . . . . . . . . . . . . . . . Genus Autalia Leach, 1819 . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

295 295 295 301

18

Tribe Homalotini Heer, 1839 . . . . . . . . . . . . . . . . . . . . . . . . Tribe Homalotini Heer, 1839 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Leptusa Kraatz, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Leptusomorpha Klimaszewski and Hoebeke, gen. n. . . . Genus Stictalia Casey, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Encephalus Stephens, 1832 . . . . . . . . . . . . . . . . . . . . . Genus Gyrophaena Mannerheim, 1830 . . . . . . . . . . . . . . . . . . Genus Silusa Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . .

303 303 304 305 306 310 310 315 335

19

Tribe Placusini Mulsant and Rey, 1871 . . . . . . . . . . . . . . . . Tribe Placusini Mulsant and Rey, 1871 . . . . . . . . . . . . . . . . . . Genus Placusa Erichson, 1837 . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

337 337 337 350

20

Tribe Athetini Casey, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . Tribe Athetini Casey, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Adota Casey, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Aloconota C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . Genus Amischa C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . . Genus Atheta C.G. Thomson, 1858 [sensu lato] . . . . . . . . . . . .

. . . . . .

351 351 355 357 359 360

xvi

Contents

Atheta Incertae Sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Dinaraea C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . . Genus Earota Mulsant and Rey, 1873b . . . . . . . . . . . . . . . . . . Genus Geostiba C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . . Genus Goniusa Casey, 1906 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Liogluta C.G. Thomson, 1858 . . . . . . . . . . . . . . . . . . . Genus Lypoglossa Fenyes, 1918 . . . . . . . . . . . . . . . . . . . . . . . Genus Mocyta Mulsant and Rey, 1873b . . . . . . . . . . . . . . . . . Genus Nehemitropia Lohse, 1971 . . . . . . . . . . . . . . . . . . . . . . Genus Ousipalia Gozis, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Paraleptonia Klimaszewski, 2002 . . . . . . . . . . . . . . . . . Genus Pelioptera Kraatz, 1857 . . . . . . . . . . . . . . . . . . . . . . . . Genus Philhygra Mulsant and Rey, 1873a . . . . . . . . . . . . . . . . Genus Pontomalota Casey, 1885 . . . . . . . . . . . . . . . . . . . . . . Genus Psammostiba Yosii and Sawada, 1976 . . . . . . . . . . . . . Genus Schistoglossa Kraatz, 1856 . . . . . . . . . . . . . . . . . . . . . . Genus Seeversiella Ashe, 1986 . . . . . . . . . . . . . . . . . . . . . . . . Genus Strigota Casey, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Tarphiota Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Thinusa Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Trichiusa Casey, 1893 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Paragoniusa Maruyama and Klimaszewski, 2004 . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . .

404 413 418 419 420 421 429 431 434 435 437 438 439 446 447 449 454 455 457 458 460 462 566

21

Tribe Falagriini Mulsant and Rey, 1873 . . . . . . . . . . . . . . . . Tribe Falagriini Mulsant and Rey, 1873 . . . . . . . . . . . . . . . . . . Genus Bryobiota Casey, 1893 [Revised Concept] . . . . . . . . . . . Genus Cordalia Jacobs, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Falagria Leach, 1819 . . . . . . . . . . . . . . . . . . . . . . . . . Genus Myrmecocephalus MacLeay, 1871 . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . .

571 571 571 573 574 575 582

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Tribe Lomechusini Fleming, 1821 . . . . . . . . . . . . . . . . . . . . Tribe Lomechusini Fleming . . . . . . . . . . . . . . . . . . . . . . . . . . Genus Xenodusa Wasmann, 1894 . . . . . . . . . . . . . . . . . . . . . . Genus Myrmoecia Mulsant and Rey, 1873 . . . . . . . . . . . . . . . . Genus Zyras Stephens, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . .

583 583 583 585 585 591

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 593 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 623

1

Introduction

The family Staphylinidae is the most diverse family of Coleoptera in North America north of Mexico (Poole and Gentili 1996), with over 4000 valid names (Newton et al. 2001), and much of this diversity is associated with habitats found in forest ecosystems (Pohl et al. 2008). Rove beetles occur in all habitat types with the exception of extremely dry habitats (Newton et al. 2001). At the landscape scale, species composition is dictated by forest type and species have strong affinities with forest microhabitats where moisture, temperature, substrate type, and debris accumulation are variables that promote high diversity in rove beetles (Pohl et al. 2008). The subfamily Aleocharinae is one of the largest lineages of all the beetles (Coleoptera) worldwide. This group includes approximately 62 tribes, 1310 described and probably valid genera and nearly 17,000 described species (Newton, unpublished database; Klimaszewski et al. 2018). The immense number of described and formally named aleocharine rove beetles only hints at the true diversity of this subfamily, with many thousands of species, and numerous higher taxa, remaining to be described throughout the world, especially in tropical regions (Ashe 2007). This seemingly endless diversity, the minute size of most adults, and the virtual lack of illustrated keys and descriptions of species for most geographical regions make the Aleocharinae one of the most taxonomically challenging groups of beetles (Ashe 2007). Aleocharines are widely distributed in North

America and occur in almost all terrestrial habitats, but the majority of species are forest dwelling where they occur in leaf litter, under bark, in fungi, in moss, and in the nests of ants, termites, mammals, and birds. Elsewhere, they inhabit seashores, edges of water bodies, wetlands and prairies (Klimaszewski et al. 2018). Aleocharine rove beetles represent key ecological or environmental indicators of changes in managed forests due to their highly specific microhabitat associations, their response to disturbances, their high local abundance, and their ease of collection (Paquin and Dupérré 2002; Pohl et al. 2008; Venier et al. 2017; Klimaszewski et al. 2018). As indicators, the presence or absence of particular taxa of these minute rove beetles may reveal details about the overall health of ecosystems and the impact of human activity on them. For this body of work we focus on British Columbia, the western most province of Canada. This province has a large diverse land area, bounded by the Pacific coastline, mountains, and islands, and is characterized by a wide range of ecosystems and habitats (Meidinger and Pojar 1991), harboring an exceptionally rich biota (Cannings and Cannings 1996). It is alleged to have more species than any other province (or territory) in Canada, with both rarity and richness “hotspots” located in three areas of the province, namely the South Okanagan, the Lower Mainland, and southeastern Vancouver Island

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_1

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plus the Gulf Islands (Scudder 2004). Some of the largest remaining tracts of intact ancient coniferous forests in North America are found in British Columbia (Winchester 1997). Although the forests are the dominant vegetation type, there also are extensive areas of tundra, wetlands, grasslands, and rolling scrub. British Columbia has several genera of aleocharines that are geographically unique to its inventory list and not found in the remaining provinces and territories of Canada. The tribe Liparocephalini, includes four genera (Amblopusa, Paramblopusa, Diaulota, and Liparocephalus) and five species that are primarily found in the intertidal zones of the Pacific shores from Korea, Japan, through Alaska to southern California and Mexico (Ahn and Ashe 1996). In the tribe Athetini, Pontomalota is another intertidal zone, seashore genus that includes two species from the Pacific coast, occurring from Alaska to Baja California (Ahn and Ashe 1992). The athetine genera Adota Casey, 1910 and Psammostiba Yosii and Sawada, 1976—both including seashore species common in decomposing seaweed on the beach—are restricted to the Pacific coast of North America and to the coasts of the northern Pacific, respectively (Gusarov 2003), but each include species recorded from British Columbia. Work on the Aleocharinae in the province is dependent on the availability of specialist taxonomists. No ecoregion, ecosystem, or habitat in British Columbia has been thoroughly inventoried for invertebrates, let alone staphylinid rove beetles. Here, we present the first comprehensive synopsis of all aleocharine rove beetle species (Coleoptera, Staphylinidae) known from British Columbia. For every species, illustrations are provided, including colour habitus and genital diagnostic structures of both sexes. Two hundred and twenty-seven confirmed and valid species from British Columbia are here presented and discussed. This taxonomic summary includes one new genus, 15 new species, 16 new generic records, and 37 (excluding new species) new BC provincial and 5 USA state records, in 79 genera and 14 tribes and provides the necessary taxonomic tools for their identification.

1

Introduction

Several metrics can be used to assess the Aleocharinae of Canada and particularly British Columbia. Thomas Casey was responsible for naming and describing 200 aleocharine species that occur in Canada, and 160 of these, approximately 80% of the total, have been retained as valid species-level taxa. As a result, Casey’s descriptive efforts account for about 30% of all named species in British Columbia, a considerably large and notable contribution. During the past three decades, an historical examination of the recorded inventories of aleocharines in British Columbia illustrates a compelling and not too surprising increasing trend in the number of species recorded. Bousquet’s checklist recorded a total of 106 species of Aleocharinae in British Columbia, Gouix and Klimaszewski (2007) listed 153 species from the province, and by 2013 the second edition of the checklist (Bousquet et al. 2013) listed 176 species, a 66% increase from 1991. With the completion of this treatment, 227 species of aleocharines are currently documented for the province, a 28% increase since 2013. Since the 1991 checklist, the number of recorded aleocharines in British Columbia has increased by about 114%, more than doubling. For all Staphylinidae, Bousquet et al. (2013) records 779 species from British Columbia alone, second in overall species richness in Canada next only to Ontario (865 spp.). With the 227 species treated herein, the Aleocharinae rove beetles constitute nearly 29% of all species of Staphylinidae recorded for the province. Nearly one-half are represented by the large, genus-rich tribe Athetini (101 spp. in 29 genera), followed (in descending number of species) by the Oxypodini (44 spp. in 18 genera), Aleocharini (23 spp., 1 genus: Aleochara), Homalotini (15 spp. in 6 genera), and Tachyusini (12 spp. in 4 genera). For additional details on these dominant tribes in British Columbia, the reader is referred to Chap. 6. This new body of work and its companion piece (Aleocharine rove beetles of eastern Canada; Klimaszewski et al. 2018) will fill a void where before it was nearly impossible to identify most aleocharine rove beetles with

1

Introduction

accuracy. Armed with habitus illustrations of the adults and detailed photographic depictions of the male and female genital structures along with the terminal segments, identification is now made possible. It is our expectation that the identification tools and information provided herein will inspire future taxonomic and ecological studies of the Aleocharinae in Canada. Further inventory work and examination of specimens in collections will likely greatly increase numbers of species in future checklists for the province. Future studies will also help gauge whether particular groups of species are at risk and will continue to reveal the captivating biologies of this diverse and complex group of beetles.

References Ahn KJ, Ashe JS (1992) Revision of the intertidal aleocharine genus Pontomalota (Coleoptera: Staphylinidae) with a discussion of its phylogenetic relationships. Entomol Scand 23:347–359 Ahn KJ, Ashe JS (1996) Revision of the intertidal aleocharine genus Amblopusa Casey and description of the new genus Paramblopusa (Coleoptera: Staphylinidae). J N Y Entomol Soc 103:138–154 Ashe JS (2007) Aleocharinae. Version 25 April 2007. http://tolweb.org/Aleocharinae/9777/2007.04.25. In: The tree of life web project. http://tolweb.org/ Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013) Checklist of beetles (Coleoptera) of Canada and Alaska: second edition. Pensoft Series Faunistica No. 109. Pensoft Publishers, Sofia-Moscow. 402 pp Cannings R, Cannings S (1996) British Columbia: a natural history. Greystone Books, Vancouver, BC. 310 pp

3 Gouix N, Klimaszewski J (2007) Catalogue of aleocharine rove beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae). Pensoft, Sofia. 165 pp Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharine), with synonymic notes. Zootaxa 353:1–134 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. 902 pp Meidinger D, Pojar J (eds) (1991) Ecosystems of British Columbia. Special Report Series No. 6. British Columbia Ministry of Forests, Victoria, BC, 330 pp Newton AF, Thayer MK, Ashe JS, Chandler DS (2001) Staphylinidae. In: Arnett RH, Thomas MC (eds) American Beetles, vol 1. CRC Press, Boca Raton, FL, pp 272–418 Paquin P, Dupérré N (2002) Beetles of the boreal forest: a faunistic survey carried out in western Québec. Proc Entomol Soc Ont [2001] 132:57–98 Pohl G, Langor D, Klimaszewski J, Work T, Paquin P (2008) Rove beetles (Coleoptera: Staphylinidae) in northern Nearctic forests. Can Entomol 140:415–436 Poole RW, Gentili P (comps) (1996) Volume 1: Coleoptera and Strepsiptera. Nomina Insecta Nearctica: a check list of the insects of North America. Entomological Information Services, Rockville, MD, 827 pp Scudder, GGE (2004) Rarity and richness hotspots in British Columbia. In: Proceedings of the species at risk 2004 pathways to recovery conference, 2–6 March 2004, Victoria, BC, 6 pp Venier LA, Work TT, Klimaszewski J, Morris DM, Bowden JJ, Kwiaton MM, Webster K, Hazlett P (2017) Ground-dwelling arthropod response to fire and clearcutting in jack pine: implications for ecosystem management. Can J For Res 47(12):1614–1631 Winchester NN (1997) The arboreal superhighway: arthropods and landscape dynamics. Can Entomol 129(4):595–599

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Prominent Forebearers of Modern Aleocharine Systematics in North America

Today’s staphylinid specialists stand on the shoulders of some of the early and contemporary giants who have contributed substantially to our understanding of aleocharine systematics in North America. Based on their life’s work and achievements, a number of deceased individuals are worthy of recognition here. We have selected five beetle specialists who have had, in our estimation, the most profound impact on aleocharine systematics in the past century and a quarter. Each has contributed substantially to the exploration and study of genus- and species-level taxa, some of which are reported from British Columbia, the principal focus of this book. The biographical sketches available in Smetana and Herman (2001: 40–159) were invaluable in producing the synopses below.

*** The Lasting Influence of Thomas Lincoln Casey (1857–1925) (Fig. 2.1a) on North American aleocharine systematics is legendary and he is discussed at greater length below in context with the historical review of research on the Aleocharinae. Max Bernhauer (1866–1946) (Fig. 2.1b), a German coleopterist, first turned his attention to the family Staphylinidae in the late 1890s, and developed into one of the leading researchers on the group. Early in his career, he published papers on the Aleocharinae, including a monograph on

the Palearctic species of Leptusa (Bernhauer 1900) and a synthesis of the tribe Aleocharini of the Palearctic region (Bernhauer 1901, 1902) (Smetana and Herman 2001). He published several early papers describing new aleocharine beetles from North America (Bernhauer 1905, 1906, 1907, 1909). As one of the leading world staphylinidologists, these collective works became standard references and identification tools for that time. Beginning in 1910, in conjunction with European workers Karl Schubert and Otto Scheerpeltz, he produced a world catalog of the Staphylinidae within the JunkSchenkling Coleopterorum Catalogus, a reference still consulted today as a “database” on the Staphylinidae (Smetana and Herman 2001). He described over 5000 species and in excess of 340 genera of Staphylinidae during his productive career (Capinera 2008). His extensive collection of rove beetles is housed in the Field Museum of Natural History in Chicago. Adalbert Fenyes (1863–1937) (Fig. 2.1c), another noted staphylindologist and Hungarian physician by training, started his studies of the subfamily Aleocharinae beginning around 1905 and thereafter was recognized as an authority on the group. Historical accounts indicate that he completed a manuscript for a monograph of the North American Aleocharinae, including 766 coloured figures, but because of its size it was never published (Smetana and Herman

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_2

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Prominent Forebearers of Modern Aleocharine Systematics in North America

2001). A copy of his notable work on aleocharine beetles is found in the Fenyes Collection in P. Wytman’s Genera Insectorum dated 1920–1921 (Pratt 2013). He also left behind an extensive collection of Aleocharinae (in the California Academy of Sciences, San Francisco) (Pratt 2013) containing over 19,000 specimens, representing some 1800 species (Smetana and Herman 2001). Gustav Adolf Lohse (1910–1994) (Fig. 2.1d), German born and a dentist by profession, developed into an accomplished taxonomic coleopterist and became a widely recognized expert on central European beetle groups and was the driving force behind the series “Die Käfer Mitteleuropas.” He published numerous papers on central European staphylinids, and particularly on the Aleocharinae. Late in his career, he published several seminal papers on North American Aleocharinae, and particularly those with arctic and Holarctic distributions (Lohse et al. 1990; Smetana and Herman 2001). He also worked on a review of Canadian species of Gnypeta but was unable to complete the project (Klimaszewski et al. 2008). Charles Hamilton Seevers (1907–1965) (Fig. 2.1e), recognized at the height of his career as an internationally acclaimed specialist on the systematics of Aleocharinae, contributed substantially to our knowledge of rove beetles, publishing on termitophilous and myrmecophilous Staphylinidae (Seevers 1957, 1965) of which a large percentage are represented by genera of Aleocharinae, and his published studies on the mushroom-inhabiting Gyrophaenae of North America and Europe (Seevers 1951). Perhaps his most acclaimed paper was published posthumously in 1978—“A generic and tribal revision of North American Aleocharinae.” Although this research was primarily concerned

with genera of North America, its overall importance and relevance extends well beyond this faunal region. He was one of the first to attempt integrating the nomenclature and classification of North American taxa with that of Eurasia (R. Wenzel, in Smetana and Herman 2001). Seevers described 215 species and 42 genera in the Staphylinidae (Smetana and Herman 2001). James Stephen “Steve” Ashe (1947–2005) (Fig. 2.1f), one of the world’s experts on the subfamily Aleocharinae and our esteemed staphylinidologist colleague, will be remembered primarily as one of the outstanding beetle systematists of his generation. Steve loved natural history, was a keen observer of nature, and delighted in discovering a beetle new to him (Timm 2006). During the height of his professional career, Steve published over 100 peerreviewed papers, including many large monographs, and numerous web pages for the Tree of Life (eg., Ashe and Maus 1998) and other projects (Lingafelter et al. 2006). His main taxonomic interest was the Aleocharinae, and his published works included a generic revision and phylogeny of the Gyrophaenina (Ashe 1984a, b), a phylogeny of the Bolitocharina (Ashe 1992), and studies of the genera Tachiona (Ashe and Wheeler 1988, Ashe 1990, 1993) and Gansia (Ashe and Lingafelter 1996). Smetana and Herman (2001) noted that the long-range goals of Steve Ashe were to make the Aleocharinae more accessible to researchers by producing a database of images of aleocharine staphylinids and by providing a framework for a phylogeny of the genera and tribes of the subfamily. In the end, he described at least 103 new species and 17 new genera during his shortened life, and as evidenced by his colleagues respect for him as a researcher and educator, 25 new species were named in his honor (patronyms), many of them Staphylinidae (Lingafelter et al. 2006).

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Prominent Forebearers of Modern Aleocharine Systematics in North America

Fig. 2.1 (a–f) Photographs of early and contemporary researchers who have had the most profound impact on aleocharine systematics in North America: (a) Thomas Lincoln Casey (1857–1925); (b) Max Bernhauer

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(1866–1946); (c) Adalbert Fenyes (1863–1937); (d) Gustav Adolf Lohse (1910–1994); (e) Charles Hamilton Seevers (1907–1965); (f) James Stephen “Steve” Ashe (1947–2005)

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Prominent Forebearers of Modern Aleocharine Systematics in North America

References Ashe JS (1984a) Major features of the evolution of relationships between gyrophaenine staphylinid beetles (Coleoptera: Staphylinidae: Aleocharinae) and fresh mushrooms. In: Wheeler Q, Blackwell M (eds) Fungus/insect relationships: perspectives in ecology and evolution. Columbia University Press, New York Ashe JS (1984b) Generic revision of the subtribe Gyrophaenina (Coleoptera: Staphylinidae: Aleocharinae) with review of the described subgenera and major features of evolution. Quest Entomol 20(3):129–349 Ashe JS (1990) New species, phylogeny and natural history of Tachiona Sharp 1883 (Coleoptera: Staphylinidae: Aleocharinae). Trop Zool 3(2):225–235 Ashe JS (1992) Phylogeny and revision of genera of the subtribe Bolitocharina (Coleoptera: Staphylinidae: Aleocharinae). Univ Kans Sci Bull 54(10):335–406 Ashe JS (1993) New species and records of Tachiona Sharp (Coleoptera: Staphylinidae, Aleocharinae) with a revised key to known species. Ent Scand 24:231–239 Ashe JS, Lingafelter S (1996) Revision of Gansia Sharp of Mexico and Central America (Coleoptera: Staphylinidae: Aleocharinae). J N Y Entomol Soc [1995] 103:251–280 Ashe JS, Maus C (1998) The tree of life web project, Aleocharini. Available at http://tolweb.org/Aleocharini. Accessed 3 July 2019 Ashe JS, Wheeler QD (1988) Revision of Tachiona Sharp (Coleoptera: Staphylinidae: Aleocharinae) with a description of the larva of T. latipennis new species, and a preliminary assessment of generic relationships. J N Y Entomol Soc 96:176–199 Bernhauer M (1900) Die Staphyliniden-Gattung Leptusa Kraatz, nebst einer analytischen Bestimmungstabelle der paläarktischen Arten. Verh Zool Bot Ges Wien 50:399–432 Bernhauer M (1901) Die Staphyliniden der paläarktischen Fauna. Verh Zool Bot Ges Wien 51:430–506 Bernhauer M (1902) Die Staphyliniden der paläarktischen Fauna. I. Tribus: Aleocharini. (II. Theil.). Verh Kaiserlich-Königlichen Zool Bot Ges Wien 52:87–284 Bernhauer M (1905) Neue Aleocharinen aus Nordamerika. Deut Entomol Z 1905(2):249–256

Bernhauer M (1906) Neue Aleocharinen aus Nordamerika. (II. Teil). Deut Entomol Z 1906(2):337–348 Bernhauer M (1907) Neue Aleocharini aus Nordamerika (Col.) (3. Stück). Deut Entomol Z 1907(4):381–405 Bernhauer M (1909) Neue Aleocharinen aus Nordamerika. (Col.) (4. Stück). Deut Entomol Z 1909(4):515–528 Capinera J (2008) Encyclopedia of entomology, 2nd edn. Springer Science & Business Media, Heidelberg. 2061 pp Klimaszewski J, Savard K, Pelletier G, Webster R (2008) Species review of the genus Gnypeta Thomson from Canada, Alaska and Greenland (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics and distribution. ZooKeys 2:11–84 Lingafelter SW, Anderson RS, Timm B, Falin Z, Jameson ML, Newton AF, Ball GE, Ahn K-J, Leschen R (2006) In: Memoriam, James Stephen “Steve” Ashe (1947–2005). Coleopt Bull 60(1):1–12 Lohse GA, Klimaszewski J, Smetana A (1990) Revision of Arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44(2):121–202 Pratt NF (2013) Dr. Adalbert Fenyes: a biographical sketch of a twentieth century entomologist. Pan-Pac Entomol 89(1):1–6 Seevers CH (1951) A revision of the North American and European staphylinid beetles of the subtribe Gyrophaenae (Aleocharinae, Bolitocharini). Fieldiana: Zoology 32(10):655–762 Seevers CH (1957) A monograph on the termitophilous Staphylinidae (Coleoptera). Fieldiana: Zoology 40(1):1–334 Seevers CH (1965) The systematics, evolution and zoogeography of staphylinid beetles associated with army ants (Coleoptera, Staphylinidae). Fieldiana: Zoology 47:139–351 Smetana A, Herman L (2001) Brief history of taxonomic studies of the Staphylinidae including biographical sketches of the investigators, pp 17–159. In: Herman L (ed) Catalog of the Staphylinidae (Insecta: Coleoptera). 1758 to the end of the second millennium. I. Introduction, history, biographical sketches, and omaliine group. Bull Am Mus Nat Hist 265:1–650 Timm RM (2006) In memoriam: James S. (“Steve” Ashe, 1947–2005). J Kansas Entomol Soc 79(2):89–91

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Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia

A serious dialogue cannot take place on a historical review of taxonomic research on the North American Aleocharinae without the mention of one of the early students of Coleopterology in this country, Thomas Lincoln Casey (Fig. 2.1a). Casey was one of the most prolific describers of species-level taxa in the subfamily Aleocharinae. Casey’s taxonomic philosophy, however, was the topic of appreciable criticism by his peers. He was considered as a “splitter” as opposed to a “lumper,” for he recognized more species, rather than less. One colleague (W. Dwight Pierce) once accused Casey of being a splitter, to which he responded, “Pierce, I am just a generation ahead of the rest of you” (Mallis 1971: 262). His many detractors charged that the morphological characters he relied upon were too trivial, failing to take into account intraspecific variation, to support the naming of new species-level taxa, and that he ignored or made little attempt to refer to previous published literature (Mallis 1971). As a result, many of Casey’s new species eventually fell into synonymy. This is evident in other beetle groups he studied, such as the Carabidae (Lindroth 1969). For example, of the 902 Casey “types” of Carabidae, only 81 of those remained valid species after close study by Carl

A comparable historical review of research on taxa of eastern Canadian Aleocharinae was completed by Klimaszewski et al. (2018). A significant number of aleocharine genera and species occurring in eastern Canada are also found in British Columbia.

Lindroth (Majka and Sikes 2009). Similarly, Casey’s taxonomic studies of the aleocharine genus Gyrophaena, a mushroom specialist and related genera were thoroughly re-examined by Charles Seevers (1951), and the result was much the same. Casey had described 47 new species in this group of genera and Seevers (1951) synonymized 20 of them. Casey may best be remembered for producing more synonyms than any other individual. The North American beetle fauna was largely unknown before the late 1800s, despite the efforts of other North American coleopterists of that era, such as Thomas Say, John LeConte, and George Horn. The slate was essentially clean for Thomas Casey to initiate his descriptive studies of staphylinids and other beetles. Casey published his first major treatment on staphylinids in 1884–1885, with his “Contributions to the descriptive and systematic coleopterology of North America, Parts I and II.” In the early 1900s, Casey published monographs on the North American Aleocharinae exclusively (Casey 1906, 1910, 1911). His most meaningful body of work was his privately published Memoirs on the Coleoptera (1910–1924) (Essig 1972). Notwithstanding his inability to recognize and accept intraspecific variation and several other taxonomic shortcomings, Casey still was one of the first to provide comprehensive taxonomic studies of staphylinid rove beetles, in particular the Aleocharinae, in the United States and

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_3

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3 Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia

Canada. Among the fauna of Aleocharinae of Canada, many of his species concepts still survive today and represent the majority of specieslevel taxa within some genera (i.e., Aleochara, Oxypoda, and many Athetini genera and species) (Majka and Sikes 2009). Below, we provide an historical account of taxonomic research of the Aleocharinae of North America, beginning from the early and mid 1800s to present day. For the majority of taxa treated by specialists over this time span, distribution records of aleocharine species occurring in Canada, particularly in British Columbia and elsewhere in the Pacific Northwest, were documented. Using the Casey years (era) as a point of reference or benchmark in this historical analysis, we have divided our treatment into three time periods: the pre-Casey era, the Casey era, and the post-Casey era.

Pre-Casey Era (1925) From the late 1920s through the 1940s, little taxonomic progress was made in aleocharine studies despite numerous publications on taxonomy, classification, developmental stages, ecology, and distributions of species of this subfamily. A few minor papers contributed to our taxonomic knowledge base during this three-decade time period. Included here is an obscure paper that provided a short list of aleocharine species, among other rove beetles, recorded from Alaska and adjacent parts of the Yukon Territory (Fall 1926). Some of these same taxa are listed from British Columbia herein. An early account of intertidal aleocharines (Liparocephalus and allied genera) occurring on shorelines of the northern Pacific coast of North America was provided by Chamberlin and Ferris (1929). The 1950s saw only a modicum of published papers on the aleocharines of Canada. The first person to provide a substantial improvement to our taxonomic knowledge of the Canadian aleocharine fauna was Charles Seevers in 1951 with a revision of North American and European Gyrophaena and allied genera (Seevers 1951). A review of the Canadian Gyrophaenina by Klimaszewski et al. (2009b) provided a modern update to Seever’s (1951) study. Another attempt to characterize and revise the group of intertidal genus-level taxa of the “Phytosi” (¼tribes Liparocephalini and Athetini, in part, of authors) of the Pacific coast was addressed in an early paper by Moore (1956). Melville Hatch (1957) added many B.C. records in Part II (the Staphyliniformia) of his seminal multivolume series “The Beetles of the Pacific Northwest,” in which he documented species and genera occurring in northern California, Oregon, Washington, and southern British Columbia. This publication has served as an invaluable historical resource for distribution records of the Pacific Northwest. While the 1960s did not see any significant number of taxonomic publications on Canadian aleocharines, the 1970s did see a profusion of published studies on the fauna of North American

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Aleocharinae. Revisions of the charismatic myrmecophilous genus Xenodusa (Hoebeke 1976) and the distinctive wetland-dwelling genera Gymnusa and Deinopsis (Klimaszewski 1979, 1982b) were completed. In 1978, a revision of the North American genera and tribes of Aleocharinae was published (Seevers 1978), with a checklist of the genera. This comprehensive monograph attempted to coordinate the nomenclature and classification of North American taxa with those of Eurasia. This study and Charles Seever’s earlier work on North American and European Gyrophaena and allied genera (Seevers 1951) represent “the only serious and comprehensive attempts to bring some degree of order out of chaos of numerous, superficially described genera for North American aleocharines. . .” (Ashe 1986). Prior to 1978, the fauna of North American Aleocharinae had never been treated systematically and keys to genera were non-existant and thus this became one of the first published works that allowed for the identification of aleocharines. Additionally, it provided a wealth of long-needed illustrations, although line drawings only, of important morphological characters and a greatly needed, critical synthesis of generic concepts (Klimaszewski et al. 2018). Seevers’ revision has become the standard reference for the study of North American aleocharines (Ashe 1986) and has also served as an important catalyst for later improvements to generic classification (e.g., Ashe 1984, 1992; Gusarov 2003a). Progress during the 1980s included taxonomic revisions of the tribe Falagriini (Hoebeke 1985), the tribe Myllaenini (Myllaena) (Klimaszewski 1982a), and the genus Aleochara (Klimaszewski 1984 and several supplements). Ashe (1986) erected a new athetine genus and species (Seeversiella bispinosa), only later to be revised by Gusarov (2003a) to include twenty-seven new species from the Nearctic and Neotropics, and to synonymize Ashe’s S. bispinosa with S. globicollis (Bernhauer), known to occur in British Columbia. The morphologically unique tribe Autaliini, including the single genus Autalia

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3 Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia

in North America, was reviewed by Hoebeke (1988). Additional research on aleocharines in Canada during the 1990s included a revision of the arctic members of the tribes Aleocharini and Athetini of North America (Lohse et al. 1990); several genera of the distinctive and ecologically unique tribe Liparocephalini whose members are intertidal species specialized to live in low and high tide zones [Amblopusa (Ahn and Ashe 1996), Diaulota (Ahn 1996a), Liparocephalus (Ahn 1997a), and Paramplopusa (Ahn and Ashe 1996)]; and also notable work on other marine intertidal athetine specialists of the genera Pontomalota (Ahn and Ashe 1992), and Thinusa and Tarphiota (Ahn 1996b, 1997b, 1999, respectively). During the first decade of the 2000s, most of the taxonomic literature that reinforced our understanding of the eastern Canadian aleocharine fauna also provided for a better comprehension of the shared fauna with British Columbia. A number of modern revisions, reviews, descriptions of new taxa, and new data on bionomics and distributions all resulted in new diagnostic tools for the provincial fauna of BC. The first comprehensive treatment of aleocharine rove beetles of British Columbia was by Klimaszewski and Winchester (2002), wherein 40 species in 9 tribes were recorded from the ancient Sitka spruce forest of the Carmanah Valley on Vancouver Island. Also, they erected a new genus Paraleptonia to accommodate an undescribed species (pacei Klimaszewski). Gusarov (2003c) examined a large number of North American type specimens of Aleocharinae that resulted in some valuable insights relevant to taxonomic work on Nearctic aleocharines (Klimaszewski et al. 2018). Species described by early workers (e.g., Casey) were often misidentified to genus, causing them to be often overlooked in modern taxonomic treatments. Some Aleocharinae, especially Athetini, were found to be more widespread than previously thought by workers such as Casey, underscoring the importance of checking all North American types as many species were potentially synonyms

or incorrectly classified. Also, subtle variation in external morphology had led to the proliferation of synonyms (for example, ten names are found under Strigota ambigua). Gusarov (2003c) also provided aids to reliably identify several of the most common Atheta species in Canada. In 2004, Neothetalia Klimaszewski, an oxypodine genus, was described and added to the eastern Canadian fauna (Klimaszewski and Pelletier 2004). This resulted in six described species of which five are recorded from British Columbia. In the same year, the Canadian species of Leptusa were also revised (Klimaszewski et al. 2004). Other genera of aleocharines that underwent critical revision or review in the early 2000s included Placusa (Klimaszewski et al. 2001); Earota (Gusarov 2002a); Geostiba (Gusarov 2002b); Pelioptera (as Tropimenelytron) (Gusarov 2002c); Seeversiella (Gusarov 2003a); Silusa (Klimaszewski et al. 2003); Psammostiba (Gusarov 2003d); Strigota (Gusarov 2003c), Tinotus (now a subgenus of Aleochara; Klimaszewski et al. 2002, Yamamoto and Maruyama 2016); Goniusa (Gusarov 2003b); Adota (Gusarov 2003d); Lypoglossa (Gusarov 2004); the other genera of the Ocalea group of the Oxypodini, including Alfocalea, Betocalea, Megocalea, Metocalea, Neoisoglossa, and Parocalea (Klimaszewski and Pelletier 2004); Oxypoda (Klimaszewski et al. 2006); Tachyusa (Paśnik 2006); Gnypeta (Klimaszewski et al. 2008); Calodera (Assing 2008); Ocyusa (Webster et al. 2009); and Schistoglossa (Klimaszewski et al. 2009a). Three species of adventive Palearctic Aleocharinae from the Maritime Provinces were reported by Majka and Klimaszewski (2008c)—including two species of the genus Meotica [pallens (Redtenbacher) and exilis (Knoch)], one of which (pallens) was recorded from British Columbia. Majka and Klimaszewski (2008b) provided eighty-eight new Canadian provincial records of Aleocharinae, including eleven first time records from British Columbia [Aleochara quadrata Sharp, Gnathusa eva Fenyes, Atheta strigosula Casey, A. longicornis (Gravenhorst), A. platanoffi Brundin, A. klagesi Bernhauer, A. frosti Bernhauer, Mocyta fungi

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Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia

(Gravenhorst), Philhygra botanicarum Muona, P. clemens (Casey), and P. “humivaga”]. A special issue of the journal ZooKeys, entitled ‘Biodiversity, Biosystematics, and Ecology of Canadian Coleoptera’ (Majka and Klimaszewski 2008a), triggered a measured increase in our knowledge of the composition of the eastern Canadian fauna and included taxonomic papers that newly addressed the taxonomy and biogeography of several aleocharine taxa in British Columbia. For example, a review of the genus Gnypeta from Canada, Alaska, and Greenland recorded three species from British Columbia (Klimaszewski et al. 2008). In a second issue (McLean et al. 2009a, b), two papers focused on aleocharine species collected in portions of undamaged and storm-damaged forests in Stanley Park, Vancouver, British Columbia between 2007 and 2008 (McLean et al. 2009a, b). Trapping yielded 35 species of Staphylinidae in the first survey conducted in 2007, including one species new to science, Oxypoda stanleyi Klimaszewski & McLean; three adventive aleocharine species, Dalotia coriaria (Kraatz), Mocyta fungi (Gravenhorst) and Oxypoda opaca (Gravenhorst) were recorded for the first time from British Columbia; and new distribution records for another four species (McLean et al. 2009a). An additional eighteen species of aleocharines were recorded in the second trapping survey of 2008 (McLean et al. 2009b). In another paper in this special issue, the Holarctic athetine genus Schistoglossa Kraatz was listed for the first time from Canada, including three species recorded from British Columbia (Klimaszewski et al. 2009a). New genus-level taxa were also described in the early 2000s and included species from the Pacific Northwest and British Columbia. For example, the new genus Paragoniusa Maruyama and Klimaszewski (2004) was erected for a new myrmecophilous species (myrmicae) known to occur in British Columbia, and also the new genus Paraleptonia Klimaszewski was described (in Klimaszewski and Winchester 2002). The publication of the first North American catalogue on Staphylinidae, including the Aleocharinae, by Moore and Legner (1975), was followed much

13

later by the first comprehensive catalogue of aleocharine beetles from Canada and Alaska (Gouix and Klimaszewski 2007). A worldwide checklist and a complete account on the biogeography and natural history of coastal Staphylinidae were produced by Frank and Ahn (2011). This contribution listed 392 species, in 91 genera, of Staphylinidae that are believed to be confined to coastal habitats worldwide. In this publication 4 tribes and 11 genera of aleocharines that include species adapted to saline habitats of the Pacific coast of North America were recorded. From 2011 to 2019, current aleocharine research has again focused on a number of comprehensive taxonomic reviews, revisions, and new distributional records in Canada for several little known and obscure genera—Dinaraea (Klimaszewski et al. 2013), Gnathusa, Mniusa and Ocyusa (Klimaszewski et al. 2014), Trichiusa (Klimaszewski et al. 2015a), Mocyta (Klimaszewski et al. 2015b), Clusiota (Klimaszewski et al. 2015c), Liogluta (Klimaszewski et al. 2016), and Boreophilia (Klimaszewski et al. 2019)— including the validation of species occurring in British Columbia. During the past nearly 30 years, documentation of the aleocharine fauna of British Columbia has proceeded at an accelerated rate, through the many published works mentioned above and others. Campbell and Davis (1991) checklist recorded a total of 106 species of Aleocharinae in British Columbia. By 2007, the total number of aleocharine taxa recorded in the province climbed to 153 species (in 54 genera) (Gouix and Klimaszewski 2007). In the second edition of the “Checklist of Beetles of Canada and Alaska” (Bousquet et al. 2013), 176 species were inventoried for British Columbia, a 66% increase from 1991. With the completion of the present treatment, 227 recorded species of aleocharines are now documented for the province, an additional 28% increase since 2013. Since the publication of Campbell and Davis (1991) checklist, the number of recorded aleocharines in British Columbia has more than doubled, increasing by 114%. For all Staphylinidae, Bousquet (2013) records 779 species from British Columbia alone, second in

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3 Historical Review of Research on Aleocharinae in Canada, with a Focus on British Columbia

overall species richness in Canada only to Ontario (865 spp.). With the 227 species treated herein, the Aleocharinae rove beetles constitute nearly 29% of the total number of staphylinids recorded for the province. The present work provides additional identification tools to hopefully steer and motivate others to continue the documentation of these regional faunas.

References Ahn KJ (1996a) A review of Diaulota Casey (Coleoptera: Staphylinidae: Aleocharinae), with description of a new species and known larvae. Coleopt Bull 50(3):270–290 Ahn KJ (1996b) Revision of the intertidal aleocharine genus Tarphiota (Coleoptera: Staphylinidae). Entomol News 107(4):177–185 Ahn KJ (1997a) A review of Liparocephalus Mäklin (Coleoptera: Staphylinidae: Aleocharinae). Pan-Pac Entomol 73(2):79–92 Ahn KJ (1997b) Revision and systematic position of the intertidal genus Thinusa Casey (Coleoptera: Staphylinidae: Aleocharinae). Entomol Scand 28:75–81 Ahn KJ (1999) Tarphiota densus (Moore), a new combination and key to the species of the genus Tarphiota Casey (Coleoptera: Staphylinidae: Aleocharinae). J Kansas Entomol Soc 71:191–193 Ahn KJ, Ashe JS (1992) Revision of the intertidal aleocharine genus Pontomalota (Coleoptera: Staphylinidae) with a discussion of its phylogenetic relationships. Entomol Scand 23:347–359 Ahn KJ, Ashe JS (1996) Revision of the intertidal aleocharine genus Amblopusa Casey and description of the new genus Paramblopusa (Coleoptera: Staphylinidae). J N Y Entomol Soc 103(2):138–154 Ashe JS (1984) Generic revision of the subtribe Gyrophaenina (Coleoptera: Staphylinidae: Aleocharinae) with a review of the described subgenera and major features of evolution. Quaestiones Entomol 20:129–349 Ashe JS (1986) Seeversiella bispinosa, a new genus and species of athetine Aleocharinae (Coleoptera: Staphylinidae) from North America. J N Y Entomol Soc 94(4):500–511 Ashe JS (1992) Phylogeny and revision of genera of the subtribe Bolitocharina (Coleoptera: Staphylinidae: Aleocharinae). Univ Kansas Sci Bull 54:335–406 Assing V (2008) The genus Calodera Mannerheim in Canada (Insecta, Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 2:203–208 Beaulne J-I (1919–1922) [Staphylinidae sensu lato] In: Les Coléoptères du Canada. Le Naturaliste Canadien 46: 47–48, 69–72, 94–96, 117–119, 136–143, 164– 167, 181–185, 212–216, 235–239, 260–263, 274– 284; 47:66–71, 89–95, 117–120, 138–142, 279–284;

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Aleocharine), with synonymic notes. Zootaxa 353:1–134 Gusarov VI (2003d) A revision of Nearctic species of the genera Adota Casey, 1910 and Psammostiba Yosii & Sawada, 1976 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 185:1–35 Gusarov V (2004) A revision of the genus Lypoglossa Fenyes, 1918 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 747:1–36 Hatch MH (1957) The Beetles of the Pacific Northwest. Part II: Staphyliformia. University of Washington Press, Seattle. ix + 384 pp Hoebeke ER (1976) A revision of the genus Xenodusa (Staphylinidae, Aleocharinae) for North America. Sociobiology 2(2):109–143 Hoebeke ER (1985) A revision of the rove beetle tribe Falagriini of America north of Mexico (Coleoptera: Staphylinidae: Aleocharinae). J N Y Entomol Soc 93 (2):913–1018 Hoebeke ER (1988) A new species of rove beetle, Autalia phricotrichosa (Coleoptera: Staphylinidae: Aleocharinae), from Mexico, with a key to the New World species of Autalia. Coleopt Bull 42(1):87–93 Klimaszewski J (1979) A revision of the Gymnusini and Deinopsini of the world: Coleoptera, Staphylinidae, Aleocharinae, No. 25. Agriculture Canada, Research Branch. Monograph No. 25, 169 pp Klimaszewski J (1982a) Studies of Myllaenini (Coleoptera: Staphylinidae, Aleocharinae): 1. Systematics, phylogeny, and zoogeography of nearctic Myllaena Erichson. Can Entomol 114(3):181–242 Klimaszewski J (1982b) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae). Supplementum 2. Can Entomol 114(4):317–335 Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Mem Ent Soc Can 129, [2] + 211 pp Klimaszewski J, Pelletier G (2004) Review of the Ocalea group of genera (Coleoptera, Staphylinidae, Aleocharinae) in Canada and Alaska: new taxa, bionomics, and distribution. Can Entomol 136:443–500 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc ro belge d’Entomol 40:3–126 Klimaszewski J, Pelletier G, Germain C, Hebert C (2001) Diversity of Placusa (Coleoptera: Staphylinidae: Aleocharinae) in Canada, with descriptions of two new species. Can Entomol 133(1):1–47 Klimaszewski J, Pelletier G, Sweeney J (2002) Genus Tinotus (Coleoptera: Staphylinidae, Aleocharinae) from America north of Mexico: review of the types, distribution records, and key to species. Can Entomol 134(3):281–298 Klimaszewski J, Pohl G, Pelletier G (2003) Revision of the Nearctic Silusa (Coleoptera, Staphylinidae, Aleocharinae). Can Entomol 135(2):159–186

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Klimaszewski J, Pelletier G, Majka C (2004) A revision of Canadian Leptusa Kraatz (Col., Staphylinidae, Aleocharinae): new species, new distribution records, key and taxonomic considerations. Belgian J Entomol 6(1):3–42 Klimaszewski J, Pelletier G, Germain C, Work T, Hébert C (2006) Review of Oxypoda species in Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics, and distribution. Can Entomol 138(6):737–852 Klimaszewski J, Savard K, Pelletier G, Webster R (2008) Species review of the genus Gnypeta Thomson from Canada, Alaska and Greenland (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics and distribution. ZooKeys 2:11–84 Klimaszewski J, Webster RP, Savard K (2009a) First record of the genus Schistoglossa Kraatz from Canada with descriptions of seven new species (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 22:45–79 Klimaszewski J, Webster RP, Savard K (2009b) Review of the rove beetle species of the subtribe Gyrophaenina Kraatz (Coleoptera, Staphylinidae) from New Brunswick, Canada: new species, provincial records and bionomic information. ZooKeys 22:81–170 Klimaszewski J, Webster RP, Langor DW, Bourdon C, Jacobs J (2013) Review of Canadian species of the genus Dinaraea Thomson, with descriptions of six new species (Coleoptera, Staphylinidae, Aleocharinae, Athetini). ZooKeys 327:65–101 Klimaszewski J, Webster RP, Langor DW, Bourdon C, Hammond HEJ, Pohl GR, Godin B (2014) Review of Canadian species of the genera Gnathusa Fenyes, Mniusa Mulsant & Rey and Ocyusa Kraatz (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 412:9–40 Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015a) New distributional records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Webster RP, Bourdon C, Pelletier G, Godin B, Langor DW (2015b) Review of Canadian species of the genus Mocyta Mulsant & Rey (Coleoptera, Staphylinidae, Aleocharinae), with the description of a new species and a new synonymy. ZooKeys 487:111–139 Klimaszewski J, Webster RP, Sikes D, Bourdon C, Labrecque M (2015c) A review of Canadian and Alaskan species of the genera Clusiota Casey and Atheta Thomson, subgenus Microdota Mulsant & Rey (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 524:103–136 Klimaszewski J, Webster RP, Langor DW, Sikes D, Bourdon C, Godin B, Ernst C (2016) A review of Canadian and Alaskan species of the genus Liogluta Thomson, and descriptions of three new species (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 573:217–256

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Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. xvi + 902 pp Klimaszewski J, Sikes DS, Brunke A, Bourdon C (2019) Species review of the genus Boreophilia Benick from North America (Coleoptera, Staphylinidae, Aleocharinae, Athetini): systematics, habitat, and distribution. ZooKeys 848:57–102 LeConte JL (1867) Additions to the coleopterous fauna of the United States. No. 1. Proc Acad Nat Sci Phila 19:361–394 Leng CW (1920) Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman, Jr., Mount Vernon, NY. x + 470 pp Lindroth CH (1969) The ground-beetles (Carabidae, excl. Cicindelidae) of Canada and Alaska, Part 1–3. Opusc Ent Suppl XXXV, 408 pp Lohse GA, Klimaszewski J, Smetana A (1990) Revision of Arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44:121–202 Majka CG, Klimaszewski J (eds) (2008a) Biodiversity, biosystematics, and ecology of Canadian Coleoptera, vol 2. PenSoft, Sofia Majka CG, Klimaszewski J (2008b) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). ZooKeys 2:85–114 Majka CG, Klimaszewski J (2008c) Adventive Staphylinidae (Coleoptera) of the Maritime Provinces of Canada: further contributions. ZooKeys 2:151–174 Majka CG, Sikes DS (2009) Thomas L. Casey and Rhode Island’s precinctive beetles: Taxonomic lessons and the utility of distributional checklists. ZooKeys 22:267–283 Mäklin F (1852) Descriptions of new taxa. In: Mannerheim CG (ed) Zweiter Nachtrag zur Kaefer-Fauna der nordamerikanischen Laender des Russischen Reiches. Bull Soc Imp Nat Moscou 25(2):283–387 Mäklin FG (1853) [Description of new taxa]. In: von Mannerheim CG (ed) Dritter Nachtrag zur KaeferFauna der nord-amerikanischen Laender des Russischen Reiches. Bull Soc Imp Nat Moscou 26:95–269 Mallis A (1971) American entomologists. Rutgers University Press, New Brunswick, NJ. 549 pp Mannerheim CG (1843) Beitrag zur Käfer-Fauna der Aleutischen Inseln, der Insel Sitkha und Neu-Californiens. Bull Soc Imp Nat Moscou 16:175–314 Maruyama M, Klimaszewski J (2004) A new genus and species of the myrmecophilous Athetini, Paragoniusa myrmicae (Coleoptera: Staphylinidae: Aleocharinae) from Canada. J Entomol Rev Japan 59(2):241–248 McLean JA, Klimaszewski J, Li A, Savard K (2009a) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species: Part 1. ZooKeys 22:5–17

McLean JA, Klimaszewski J, Chandler DS, Savard K, Li A (2009b) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species: Part 2. Zookeys 22:19–33 Moore I (1956) A revision of the Pacific coast Phytosi with a review of the foreign genera (Coleoptera: Staphylinidae). Trans San Diego Soc Nat Hist 12 (7):103–152 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). Division of Agricultural Science, University of California. Special Publication No 3015:514 pp Paśnik G (2006) A revision of the World species of the genus Tachyusa Erichson, 1837 (Coleoptera, Staphylinidae: Aleocharinae). Zootaxa 1146:1–152 Sahlberg CR (1831) Insecta Fennica, dissertationibus academicis, A. 1817–1834 editis. Pars I: A. Frenckelliana, Helsingforsiae, pp 361–408 Say T (1830) Descriptions of new species of North American insects, and observations on some already described. New Harmony, pp 1–41 Say T (1834) Descriptions of new North American insects and observations on some already described. Trans Am Philos Soc 4(NS):409–470 Say T (1839) Descriptions of new North American insects and observations on some already described. Trans Am Philos Soc (NS)6:155–190 Seevers CH (1951) A revision of the North American and European staphylinid beetles of the subtribe Gyrophaenae (Aleocharinae, Bolitocharini). Fieldiana Zool 32:655–762 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae). Fieldiana Zool 71:vi + 275 pp Smetana A, Herman L (2001) Brief history of taxonomic studies of the Staphylinidae including biographical sketches of the investigators, pp 17–160. In: Herman LH (ed) Catalog of the Staphylinidae (Insecta, Coleoptera): 1758 to the end of the second millennium. vol. 1, Introduction, history. Bull Am Mus Nat Hist 265:1–650 Walker F (1866) Appendix. A list of mammals, birds, insects, reptiles, fishes, shells, annelides, and Diatomaceae, collected by myself in British Columbia and Vancouver Island, with notes on their habits. [List of Coleoptera], pp 309–334. In: Lord JK (ed) The Naturalist in Vancouver Island and British Columbia, Richard Bentley, London, 375 pp Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. 1. Aleocharinae. ZooKeys 22:171–248 Yamamoto S, Maruyama M (2016) Revision of the subgenus Tinotus Sharp, stat. n., of the parasitoid rovebeetle genus Aleochara Gravenhorst (Coleoptera, Staphylinidae, Aleocharinae) from Japan, Taiwan, and the Russian Far East. ZooKeys 559:81–106

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Material and Methods

Format The classification of taxa used herein selectively follows concepts expressed by Benick and Lohse (1974), Seevers (1978), Klimaszewski (1979, 1982, 1984), Klimaszewski et al. (2018), Lohse et al. (1990), Ashe (2001), Gusarov (2003, 2011), Elven et al. (2010, 2012), Paśnik (2010), Bouchard et al. (2011), Hlaváč et al. (2011), Schülke and Smetana (2015), and several treatments of Canadian genera by Klimaszewski et al. (2018), and Yamamoto and Maruyama (2017). Distributional records and bionomic information (e.g., habitat associations, collection dates, collecting methods) are based on published records and on specimens in the collections of Laurentian Forestry Centre, the Canadian National Collection of Insects, Arachnids and Nematodes, collection of University of British Columbia, George J. Spencer Entomological Museum, and Royal British Columbia Museum. In the case of unpublished records, we provide specimen data under the respective species. Only records considered to be reliable are listed.

AB—Alberta AK—Alaska BC—British Columbia LB—Labrador MB—Manitoba NB—New Brunswick NF—Newfoundland NS—Nova Scotia NT—Northwest Territories NU—Nunavut ON—Ontario PE—Prince Edward Island QC—Quebec SK—Saskatchewan YT—Yukon Territory State abbreviations for the United States of America follow those of the United States Postal Service.

Institution Codes

AMNH

Abbreviations BC jurisdictions are boldfaced and other jurisdictions in Canada and USA are in plain text.

BGC CAS

American Museum of Natural History, New York, New York, USA Benoit Godin private collection, Whitehorse, Yukon, Canada California Academy of Sciences, San Francisco, California, USA

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_4

17

18

CNC

FMNH

LFC

MCZ

RBCM SEMUC UBC

USNM

ZMLU ZMH VAC

4 Material and Methods

Canadian National Collection of Insects, Arachnids, and Nematodes, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada. Integrative Research Center, The Field Museum of Natural History, Chicago, Illinois, USA Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, R. Martineau Insectarium, Quebec City, Quebec, Canada. Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA Royal British Columbia Museum, Victoria, British Columbia, Canada Snow Entomological Museum, University of Kansas, Kansas, USA University of British Columbia, George J. Spencer Entomological Museum, Vancouver, British Columbia, Canada United States National Museum, Washington, DC, United States of America. Zoological Collection, Lund University, Lund, Sweden Zoological Museum, Helsinki, Finland. Volker Assing collection, Hannover, Germany.

Diagnostic Features (Figs. 4.2–4.9) Most of the text below regarding diagnostic features is taken from Klimaszewski et al. (2018). It is important that BC readers have all the information accessible in one information source thus reducing unnecessary searches in other documentation. Aleocharine beetles are highly diverse taxonomically, morphologically (Figs. 4.2, 4.8, and 4.9), and ecologically. Significant challenges with species identification due to the poor state of knowledge of many groups and the lack of comprehensive diagnostic tools are the main obstacles to understanding species richness,

assemblage composition, and ecological roles of aleocharines in terrestrial ecosystems. There are many species, often forming groups of cryptic species, which are externally similar, especially in the large tribes Aleocharini (e.g., Aleochara), Athetini (e.g., Atheta), Oxypodini (e.g., Oxypoda), Homalotini (e.g., Gyrophaena). In Canada, these are often represented by pairs of sibling species that can only be distinguished with certainty by the examination of genital structures.

External Body Structures Important for Identification (Figs. 4.2–4.6) The terminology used herein follows that used by previous authors (Benick and Lohse 1974; Seevers 1978; Klimaszewski 1979, 1984; Klimaszewski et al. 2018; Lohse et al. 1990; Ashe 2001; Gusarov 2003). The body length of aleocharine beetles ranges from 1.0–13.0 mm (typically 3.0–5.0 mm) and habitus forms are diverse (Fig. 4.2a–l), reflecting different adaptations to a variety of microhabitats. They range from somewhat flat (e.g., Dinaraea, Placusa, and Xenodusa) to subcylindrical (e.g., some Leptusa), and from robust (e.g., Aleochara, Gymnusa, and Oligota) to slender (e.g., Atheta, Clusiota, Myllaena, and Meotica). The term forebody refers to head, pronotum, and elytra as a combined structure. The most important body structures used in identification of aleocharines are illustrated (Figs. 4.3–4.6). These relate to the head (Fig. 4.3): frontal suture, genae (postocular area, temples), infraorbital carina, neck; mouthparts (Fig. 4.4a–f): labrum, labium, maxillae, labial palps, and ligula; prothorax (Figs. 4.3 and 4.5): pubescence pattern, hypomeron, pro-, meso- and metaventrites and their intercoxal processes, isthmus, mesothoracic peritremes (small structures that encompass the spiracles behind the procoxae), and coxae; elytra (Fig. 4.3): size and shape, pubescence pattern, elytral suture, basal margin, and lateral emargination; abdomen (Figs. 4.3 and 4.6e) [ten segmented, position indicated by Roman numerals starting at the base; when describing features of the “first visible tergites” in the keys and

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Material and Methods

diagnoses, we are referring to the basal segments following morphological segments I and II which are reduced and largely hidden by the elytra; segments IX and X are modified]: transverse impressions of basal tergites, structures inside impressions, and shape of the apical margin of male tergite VIII, and female tergite and sternite VIII; legs (Fig. 4.3): tarsal formula and length and proportions of the tarsomeres of the hind tarsus.

Genital Structures Important for Identification (Figs. 4.7–4.9) The morphological diversification of the genitalia, particularly the shape of the median lobe of the aedeagus and its internal sac structures, including flagellum and internal sclerites (structures), morphology and macro-setal pattern on the apical lobe of parameres, and morphology, of the spermatheca, likely have played an important role in aleocharine evolution. These morphological features vary in different species and are generally the structures most important for speciesand genus-level identification. The three-lobed aedeagus of aleocharines, along with complex parameres, is the most distinctive feature of the entire subfamily, and most likely constitutes a synapomorphy for this group. Aedeagus (Fig. 4.7a, b). The male copulatory organ consists of the median lobe (penis) (Fig. 4.7a, b), and the two parameres (Fig. 4.7c, d). Median Lobe of Aedeagus (Figs. 4.7a, b and 4.8a–i). This is a tubular, sclerotized, intromittent organ with an enlarged basal portion called the bulbus (Fig. 4.7a, b), and a narrow apical portion, subcylindrical or trough-like, called the tubus (Fig. 4.7a, b). The tubus is slender, and narrow towards the apex. The lateral edges of the tubus are extended dorsally in Athetini and form a complete or incomplete arched structure called the athetine bridge (Seevers 1978). The shape of the tubus, and particularly its apical portion in lateral view, varies from species to species and is a valuable diagnostic feature. The

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subspherical bulbus has a small, ventral round opening, the foramen mediale, which is an entrance for the ejaculatory duct, the ductus ejaculatorius, and one or two anterior projections called the crista apicalis or external carina of the bulbus. The crista apicalis provides attachment sites for the muscles connecting the median lobe and the two condylites of the parameres (Fig. 4.7b). The internal space of the bulbus contains a complex of mainly dorso-ventral muscles attached internally to the base of the bulbus and to the dorsal mobile compressor plate that is oval and weakly sclerotized. Contraction or expansion of the dorso-ventral muscle complex is directly or indirectly responsible for creation of hydrostatic pressure that results in eversion or retraction of the internal sac and its associated internal structures (e.g., sclerites, membraneous structures, and flagellum). The flagellum (also known as the virga or copulatory piece) (Fig. 4.7a, b) is a short or moderately-toextremely elongate tubular structure (present in some Aleochara, Aleodorus, and Oxypoda) that can be narrow or broad, and it has a swollen basal part connected with the ejaculatory duct. The internal sac membranes frequently bear numerous spinules, denticulate plates, and other structures to aid in retention of the sac in the vulva (female copulatory tract). During eversion of the internal sac, the internal sclerites and flagellum are extended (everted) exteriorly filling the internal space of the female vulva and allowing safe transfer of sperm, or packages of sperm contained in spermatophores, with the aid of the flagellum. The internal sac is withdrawn by a set of longitudinal muscles that originate on the proximal surface of the base of the bulbus. The structures and function of the internal sac were previously explained and illustrated for Aleochara curtula by Peschke (1978). The ventral side of the median lobe of the aedeagus is considered to be the side of the bulbus containing the foramen mediale, the entrance of the ductus ejaculatorius, and the adjacent ventral side of the tubus of the median lobe with the internal sac and its structures (this part is referred to as the parameral side in some recent

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publications (e.g., Gusarov 2003); the opposite side is referred to as the dorsal part. Parameres (Fig. 4.7c, d). These are the tactile orientation organs, which enable the male to find the tip of the female abdomen just before copulation, using the setal pattern on the female abdomen as a guide (Peschke 1978; Klimaszewski 1984). There are two symmetrical parameres, one on each side of the median lobe of the aedeagus, which are attached to the crista apicalis (represented by one, two or several carinae) of the bulbus of the median lobe by a complex system of muscles. Each paramere is subdivided into a paramerite and a narrowly elongate condylite. The paramerite is the main part of the paramere, and consists of a narrow apical lobe bearing a few macrosetae arranged in a particular pattern specific to different genera. Attached to the paramerite is a semimembraneous velar sac bearing microribbings, and the posterior, slat-shaped structure delimited from the apical portion by a weakly sclerotized strip (Klimaszewski 1984). The morphology of the paramere and the setal pattern of the apical lobe of the paramerite provide diagnostic characters at the generic and higher taxonomic levels. Spermatheca (receptaculum seminis) (Figs. 4.7e and 4.9a–l). This is the female genital organ for storing and perhaps selecting sperm (Peschke 1978; Dybas and Dybas 1981). The spermatheca consists of a bulbous capsule connected to a tubular stem and a thin and membranous, weakly sclerotized seminal canal (ductus spermathecae), which connects the spermatheca to the vulva and serves to transport sperm. The seminal canal is not considered to be a part of the spermatheca (Klimaszewski 1984). The capsule is a spherical or broadly tubular structure that may be straight or bent, and bears apical invaginations of various shapes and sizes. The stem may be a simple narrow tube or more complex and subdivided into an elongate chamber and a narrower duct. The shape of the spermatheca varies from species to species and is valuable for species separation.

4 Material and Methods

In some genera (e.g., Acrotona, Amischa, some Atheta, Gnypeta, Liogluta, and Mocyta) the shape of the spermatheca may be a better diagnostic character than the shape of the median lobe of the aedeagus. Terminalia (Fig. 4.6a–e). The terminalia herein refer to the male and female tergite VIII and sternite VIII (Fig. 4.6e) and the pygidium, the latter consisting of 2–3 modified terminal segments closing the end of the abdomen (Fig. 4.6e). Tergite VIII and sternite VIII provide useful additional morphological features for species level diagnostics. These structures can be elongate, subquadrate, or transverse and can have varying pubescence patterns. The antecostal suture is located subparallel to the base of tergite VIII and sternite VIII. The antecostal suture may be separated by a wide or narrow distance from the base of the disc and may be straight or sinuate in different species. The apical margin of tergite VIII of males is highly modified in many groups of aleocharines, and particularly in athetines, and often bears teeth, denticles, and other projections or emarginations, which provide important taxonomic identification features (Fig. 4.6e). The structure of the pygidium is slightly different between males (Fig. 4.6a, b) and females (Fig. 4.6c, d). In the male, the pygidium consists of tergite X, accompanied on both sides by tergite IX (completely subdivided or connected by a narrow dorsal strip) which is extended ventrally into two posterior narrow ventral struts (Fig. 4.6a, b). Ventrally located between the subdivided tergite IX of the male is a narrowly oval and flat structure which is regarded as the modified sternite IX (Fig. 4.6a). The pygidium of the female is generally similar to that of the male but is lacking ventral struts and an elongate sternite IX (Fig. 4.6c, d). The shape and pubescence pattern of tergite X is often diagnostic for species and genera. Specimens can be sexed by examination of the ventral part of the pygidium for presence (male) or absence (female) of the apical portion of sternite IX which is present in males as a median rounded lobe (Fig. 4.6a, b).

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Material and Methods

Microdissections and Preparation of Genital Structures for Examination and Diagnostics (Figs. 4.10a–f and 4.11a–f) Identifying aleocharine species and genera can be challenging due to the small size of most taxa and minute size of many distinctive characters, coupled with a vast number of valid taxa (Ashe 2001; Hanley and Ashe 2003). There are many species, often forming groups of sibling species, which are externally similar, especially in the large tribes Athetini (e.g., Atheta), Oxypodini (e.g., Oxypoda), and Homalotini (e.g., Gyrophaena). Thus, properly dissected genital structures (median lobe of aedeagus, parameres, spermatheca, and tergite VIII and sternite VIII of both sexes) carefully mounted in Canada balsam on plastic microslides for examination, are indispensable for identification of the majority of species and is therefore essential for serious work in aleocharine systematics (Figs. 4.10a–f and 4.11a–f). The methodology of dissecting aleocharine species mouthparts is sufficiently described by Hanley and Ashe (2003) and is not discussed here because mouthparts are usually not essential for species identification. Relaxation of Specimens (Fig. 4.10a–f) Specimens stored in 70% ethanol do not need to be relaxed prior to dissection. However, dried mounted specimens need to be relaxed in 5% of ammonium hydroxide solution prior to dissections. The duration of relaxation depends on the age of the dried specimens: those from old collections need to be relaxed for several hours but more recently mounted specimens need to be relaxed for only 5–15 min. When the abdomen flexes easily it is usually ready for dissection. Extremely dry and old specimens may be quickly relaxed by immersing them in hot tap water or heating in a boiling water bath for 1–2 min. Some heavily sclerotized structures may be cleared in cold 10% potassium hydroxide for 1–3 min, washed in water, and then dehydrated in absolute ethanol prior to mounting in Canada balsam on slides.

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Dissection and Dehydration (Figs. 4.10a–f and 4.11a–f) With the aid of a stereoscopic microscope, specimens are best dissected in water placed in a microscope depression slide (commercially available slides with impressions used for blood analysis). The abdomen is held in place by forceps while a hooked or straight minuten pin, mounted in a commercially available handle, is inserted between abdominal segment VII and VIII and pulled to dislodge the terminal segments from the rest of the abdomen. The genital structures (aedeagus or spermatheca) are usually inside the dislodged apical part of the abdomen but sometimes may remain in the basal part of the abdomen and should be removed from there. The dislodged apical part of the abdomen is dissected further by removing tergite VIII and sternite VIII from the pygidium using dissecting needles and then breaking the lateral internal muscles that fix the discs in place and pulling the sclerites apart. Subsequently, the aedeagus or spermatheca are also removed and detached from other structures so that tergite VIII, sternite VIII, aedeagus (or spermatheca), and pygidium are separated from each other. Remnants of muscles attached to structures may be removed mechanically using dissecting needles while structures are immersed in water or alternatively structures may be cleared in cold 10% potassium hydroxide for 1–4 min to digest the soft tissues and then washed in water. The aedeagus needs further dissection by removing the parameres from the median lobe using dissecting needles to break the muscles connecting the condylite of the paramere with the base of the bulbus of the median lobe. All dissected structures need to be washed in clear water, transferred to 75% ethanol for about one minute and then to 100% ethanol for dehydration for about 1–2 min before mounting. Structures should then be mounted in Canada balsam on a celluloid microslide attached to the pin holding the card-mounted body of the dissected specimen. Celluloid microslides are commercially available in different sizes; the size appropriate for Aleocharinae is 4.5 mm
11 mm or 6.5 mm
14.0 mm. One source of celluloid

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slides is: Entomowinkler—insects, entomological books and supplies, Dittesgasse 11, 1180 Wien, Austria ([email protected]). For beginners it is best to practice dissection on some unimportant specimens, starting with larger specimens. The genital structures should be mounted on microslides in a consistent pattern of arrangement to facilitate easy comparison of homologous structures on different microslides. We follow the following arrangement of structures on the slide: median lobe of aedeagus in lateral view or spermatheca in right upper corner; separated parameres situated below the median lobe on the right side; tergite VIII in left upper corner, sternite VIII below tergite VIII, and pygidium below sternite VIII. For thinning Canada balsam, we use xylene or non-toxic terpineol, both commercially available. It is important to note that, at first, only a thin layer of Canada balsam is placed on a microslide to prevent floating of structures into undesirable positions. After the first layer of Canada balsam with the genital structures placed in their optimal position on a microslide is dried, a second layer of Canada balsam can be added to completely, but without excess, cover the structures. We regularly use the above procedure and we find it facilitates the microscopic examination of genital structures and their photography. The advantage of this system is that the dissected specimen and the genital structures are on the same pin in the collection instead of being stored in two separate collections which can be easily misplaced or lost. This approach is also superior to using genitalia vials which take up more room on the pins, may damage structures when taken out of the vial, and requires more handling time.

Image Production (Figs. 4.12a–f and 4.13a–f) Images of the entire body and the genital structures were taken using an image processing system (Nikon SMZ 1500 stereoscopic microscope; Nikon Digital Camera DXM 1200F, and Adobe Photoshop software). Images of the dorsal

4 Material and Methods

aspects of the head, pronotum, elytra, abdomen, legs and antennae were taken separately in the horizontal plane (except for the head image, which was taken while in a natural position) and fused together in Adobe Photoshop. Images of tergite VIII or sternite VIII that were damaged during dissection were ‘repaired’ by digitally duplicating the undamaged portion using Adobe Photoshop software.

Collection Methods and Habitats (Figs. 4.14a–f and 4.15a, b) The Aleocharinae live in almost every conceivable terrestrial habitat from alpine tundra to the intertidal zone of oceans (Klimaszewski et al. 2018). The majority of species live in forest litter and wetlands and prefer moist habitats. However, many species are more specialized and occur in fungi, carrion, and subcortical habitats. Some species are associated with ants and others live in mammal and bird nests. To fully sample the species at a given site, one must use various collection methods and sample over time as many microhabitats as possible. Among the commonly used collection methods are traps such as pitfall traps, flight intercept traps, Lindgren funnel traps, Malaise traps, and light traps. Other productive sampling methods are sifting litter, treading wet vegetation, hand collecting, and evening flight collections using an aerial net. Often the habitat will dictate the most effective collection technique(s) to use. Each collection technique will be discussed below. Pitfall Traps (Fig. 4.14a, b). Pitfall traps are cheap, simple to deploy, and collect a large number and diversity of Aleocharinae (and other epigaeic arthropods) in many habitats, e.g., forests, fields, peatlands. They do not work well on highly rocky substrates (e.g., alvars) or in very wet substrates as the higher water table forces the trap out of the substrate. Pitfall traps may have different diameters, and there are some varieties that are commercially available. However, it is

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also easy to construct them from commonly available plastic containers (e.g., yogurt, ice-cream, 16 oz beer cups). A variety of preservatives may be used (e.g., salt water, propylene glycol), and an elevated rain cover reduces dilution of the preservative and minimizes input of leaves, etc. There are many useful references about the use of pitfall traps (e.g., Martin 1977; Spence and Niemelä 1994; Skvarla et al. 2014). Pitfall traps may be baited with carrion, dung, or decaying organic materials (e.g., fruit, vegetables) to attract diverse aleocharine assemblages associated with such substrates. Baits can be placed adjacent to the trap opening, in a small screened cage above the opening, or in a slurry of the killing agent which is later strained to remove the specimens (the latter method is less “pleasant” for sample processing). Luminoc® pitfall-light traps use a small light built into the trap so that the trap has both a passive and active means of sampling insects. The cup is 12 cm in diameter, and lights are activated by a photocell with timer (Jobin and Coulombe 1992). The traps are commercially produced by Bio-Control Inc., Sainte-Foy, Quebec, Canada. These traps attract a greater diversity of species than unbaited pitfall traps and often catch rarely encountered species (Hébert et al. 2000; Klimaszewski et al. 2007), but they are more expensive to purchase and maintain. Collecting at Lights (Fig. 4.14c). Many Coleoptera are attracted to ultraviolet light at night. There are many designs of light traps that can capture a wide diversity of Coleoptera including Aleocharinae. However, these traps also collect many Lepidoptera and the small Aleocharinae (which includes most species) are often lost among the moths and moth scales, rendering this method somewhat ineffective. The most productive light-based method for capturing Aleocharinae is to use a white sheet as a background to a mercury vapour light, which is more attractive to most insects (including Aleocharinae) than ultraviolet light. In remote sites one can power the light by a portable generator. Most Staphylinidae and Aleocharinae come

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to the lights from just before sunset until about 1–2 h after sunset. Specimens can be collected off the sheet using an aspirator or collected directly into the collection vial. It is best to initially place the Staphylinidae into vials with sawdust (see below) without a killing agent (e.g. ethyl acetate), otherwise many individuals will die with their wings extended. After collecting is completed for the evening, a few drops of ethyl acetate and water should be added to the sawdust to kill the specimens. Larger Coleoptera should be placed in a separate collection vial so as to prevent them from destroying the smaller beetles. Lindgren Funnel Traps and Panel Traps (Fig. 4.15a). Lindgren funnel traps of various lengths (typically with 8–12 funnels) (Fig. 4.15a) present a vertical cylindrical profile that visually mimics tree trunks, a silhouette that is attractive to many insects associated with trees (Lindgren 1983). These traps are frequently deployed with semiochemicals (e.g., pheromones, monoterpenes, ethanol) for monitoring bark and ambrosia beetles (Curculionidae: Scolytinae), and other bark- and wood-boring beetles (e.g., Cerambycidae, Buprestidae), but they are often effective for sampling many other groups of Coleoptera that are associated with dead and dying trees (Lindgren 1983). A number of Aleocharinae species in eastern Canada were captured almost exclusively in Lindgren funnel traps (Webster et al. 2009, 2012, 2016b). As these traps sample only flying insects, they also provide useful data about flight capabilities and phenology of captured species. Traps can be suspended at any height above the ground but typically with the collecting cup at about 1.5 m. Traps deployed higher in the canopy of trees will often capture species that are rarely captured in traps near the ground (Webster et al. 2016a, c) and some may be canopy specialists. For details on Lindgren trap deployment, see Hughes et al. (2014). Black panel traps, a type of cross-vane trap, function in much the same way as Lindgren funnel traps and are less expensive. As well, the panels, made of corrugated plastic, are available in a variety of colours that may have differential attraction for different species.

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Both styles of trap are available at Synergy Semiochemicals Co., Burnaby, British Columbia, Canada. Flight-Intercept Traps (Fig. 4.14d). Flightintercept traps (or FITs) take advantage of typical beetle behaviour to drop upon hitting a barrier. FITs are typically composed of a clear plastic barrier or a tightly pulled vertical mesh barrier across a suspected flight corridor within a habitat (e.g., Peck and Davies 1980). Wide collecting trays, e.g., aluminum catering trays, are placed underneath the barrier and filled with a high concentration killing agent (see options discussed below) to prevent degradation. FITs should be serviced about once a week (more frequently in sun-exposed hot environments) to prevent the decay of fragile specimens. Aleocharines that are flight-capable can be sampled in great numbers by FITs, but it is important to take a few precautions to prevent heavily damaging specimens. First, large beetles or other insects should be removed and placed in a separate sample bag or vial. Then a fine aquarium net can be used to sieve the remaining specimens from the preservative, and these are placed into a sample container or bag already filled with ethanol. Avoid pouring the contents of the trays into the net or using a squirt bottle to wash the net as these generally create a lot of damage to aleocharine specimens. There are varieties of FITs that can be attached to tree trunks for sampling saproxylic species (e.g., Hammond 1997), and these sometimes collect species that are not collected by funnel traps or panel traps (Langor, unpublished data). Killing and Preserving Agents in Traps. When sampling with traps one must use a killing agent/preservative as traps are often sampled at intervals of one or more weeks and specimens are therefore subject to decay or destruction by larger predaceous arthropods. Commonly, propylene glycol is used alone or in combination with other chemicals as it is non-toxic to mammals and lasts a long time. However, propylene glycol

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often renders Aleocharinae specimens very stiff and difficult to dissect. One of the best killing agents and preservatives for Aleocharinae is a saturated salt solution (pickling salt) - with a few added drops of unscented dish detergent or Kodak Photo-Flo 200 (wetting agent, to prevent beetles from floating to safety)– as this allows specimens to remain flexible and this enables easier dissection later. The advantage of a saturated salt solution is that salt is readily available in most places and is non-toxic. Use of saturated salt solution is an advantage in remote areas and when traps are checked frequently; however, during long collecting intervals and in hot climates water will evaporate more readily than will propylene glycol. In dry and hot climates, one may have to periodically add water to traps to counteract dehydration, but this is usually not a problem in eastern Canada. After removing specimens from traps, they can be frozen or refrigerated. However, specimens should be rinsed in water before processing, otherwise they may become encrusted with salt after they dry. Salt on specimens is easily removed by briefly dipping the specimen in water and dabbing off the water with paper towel or tissue paper. We also recommend that specimens be frozen before processing if specimens are to be used for morphological study only. Freezing beetles before complete dehydration will cause degradation of DNA. Both propylene glycol (50%) and saturated salt solution will preserve DNA in specimens but as soon as possible, specimens to be used for molecular work should be transferred to 95% ethanol where they can be dehydrated [Note: ethanol should be changed at least once (twice or three times in the case of bulk samples) before freezer storage]. Limitations of Traps. Traps can be an effective way to sample Aleocharinae and other Coleoptera with minimal effort; however, there are some disadvantages. Using traps, one rarely obtains details on microhabitat associations of the captured species, although good macrohabitat data may be obtained. Pitfall traps baited with dung

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or carrion do provide some useful data on probable habitat associations. Many species captured in Lindgren funnel traps and black funnel traps may be associated with standing tree habitats, and data from these traps and flight intercept traps provides insights into flight capability and phenology. Another disadvantage of traps is that there are many habitats in which Aleocharinae live that cannot be effectively sampled using traps. These include various wetland habitats (marshes, fens, sphagnum bogs, ponds, vernal pools, and lake margins), and river and stream margin habitats that are often rich in Aleocharinae. Pitfall traps will likely be flooded or cannot be easily deployed in these habitats. Some specialized habitats for Aleocharinae, such as bird and mammal nests, cannot be easily sampled using traps (but see Gollkowski (2013) for an effective design). Other sampling methods such as sifting, treading, and hand collecting are very effective for collecting Aleocharinae and other Coleoptera in these kinds of habitats. Sifting (Figs. 4.14f and 4.15b). Leaf litter, moss, or other kinds of debris can be placed in a large plastic box and hand-sifted by removing excess and larger debris, which concentrates specimens. Most Coleoptera, including Aleocharinae, move or fall (inactive species) to the bottom of the container and are easily collected using an aspirator as they become active (some species may take more than 10 min to become active, especially on cool days). This method is very effective for collecting species from forest litter, mosses, grass and mosses in marshes and bogs, patches of debris in sandy habitats, flood debris, and drift material along the shores of rivers, ponds, and lakes, moist leaves along vernal pond margins, seaweed along sea beaches, compost, dung, and from rotting logs. Aleocharinae are often abundant, especially later in the summer and fall, in various kinds of fungi. These are collected by removing fungi from the substrate (forest floor, logs, standing trees) and sifting them or placing them in the plastic box, breaking them into pieces, and then aspirating the specimens as they move. It is

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important to record data on the microhabitat that the specimens were collected from, including at least the major fungal group and stage of decay (fresh, decaying, rotten) when collected from fungi. Samples can also be concentrated using sifters with the finer material placed on a plastic sheet or in a plastic box for examination and specimen aspiration. For extremely small or inactive species, this sifted material can be placed in a Berlese funnel or Winkler extractor to extract specimens passively over time. Winkler extractors are more flexible as they do not require electricity and can be hung from trees or other structures at collection sites. This may be the most effective way to collect species like Thecturota spp., which are very slow-moving (Brunke et al. 2012), but other species of the genus have been collected by car net (Klimaszewski et al. 2017). Soaking. An alternative means of extracting aleocharines from many of the same substrates noted above is to immerse the material in a pan of water. The beetles quickly make their way to the water surface where they are easily picked out with forceps. This method works especially well for extracting specimens from organic material (e.g., moss, wrack) in riparian zones (where a water supply is in close reach) but also works well for leaf litter, highly decayed wood, and dung. A white plastic pan works best as the light colour makes the specimens easier to see. A large pan that can hold water to a depth of 20–25 cm is recommended. To process material quickly, several pans can be ‘operated’ at once. After about 3–4 min of soaking the large majority of specimens will have arrived at the water surface at which point the sample can be discarded. It helps to break apart large clumps of moss when they are submerged. Treading Water Vegetation. Many Aleocharinae live in floating or emergent vegetation along edges of lakes, ponds, vernal pools, rivers and streams and amongst vegetation in wet ecosites such as marshes, bogs, and fens. Treading is the most effective method for collecting beetles in these habitats. Vegetation or moss is

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pressed into the water by walking on it. Beetles will float to the surface within 1–2 min and can be hand-picked from the surface of the water. It is best to repeat this once or twice in the same trodden spot to maximize the number and diversity of species collected as some species take more time and disturbance to reveal themselves. Hand Collecting (Figs. 4.14e and 4.15b). Many Aleocharinae species live on gravel bars and on gravel and cobblestone shorelines. These habitats are best sampled by turning over pebbles, cobblestones, or large rocks, and aspirating the specimens as they move. It is important to sample shaded and sun-exposed sites and at different distances from the stream margins as different species can be found in each habitat. Collecting under rocks and pieces of dead wood can also be effective in drier habitats. Many Aleocharinae live in subcortical habitats. These are collected by first removing bark and checking underneath for revealed specimens. Bark and pieces of rotting wood should be placed into a plastic box, broken up somewhat, and then sifted or soaked as described above. Splashing. Some species of Aleocharinae live in habitats that are best sampled by ‘splashing’ the substrates they live in. A number of species such as Gnypeta and some Philhygra live in moss or in cracks on rocks in the middle of rivers or in moss in the splash zones of waterfalls (Klimaszewski et al. 2008; Webster et al. 2016b). Splashing the mosses with water will cause the beetles living there to move to the surface where they can be collected. It often takes repeated splashing for up to ten minutes before the beetles will leave the substrate. These same substrates may also be soaked as described above. Other species live in fine sand at stream and lake margins. Lightly splashing the area will force the beetles out of the sand. Often these are very small to minute species and close observation while on hands and knees is necessary to see the specimens as they leave the sand. Splashing gravel and cobblestone habitats along streams can also be effective.

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Intertidal Habitats. Aleocharinae from the tribe Liparocephalini (Amblopusa, Diaulota, Liparocephalus, Paramblopusa), and Athetini (Tarphiota, Thinusa) live in intertidal habitats (Ahn 1996, 1997a, b; Ahn and Ashe 1995, 1996), and some species can be collected only at low tide. These species are usually found under stones and rocks with a flat surface set in a sand/ clay substrate. The beetles occur on the underside of the rocks usually in depressions that presumably trap air during high tide. Other species occur in decaying seaweed and wrack near the high tide mark and are sampled by sifting or soaking the debris. It is very likely that other undiscovered species occur in these habitats in eastern Canada. Ant Nests. Some Aleocharinae are myrmecophilous and live in ant nests (Maruyama and Klimaszewski 2004, 2006; Maruyama et al. 2008; Parker 2016); however, little information has been published on how to sample these species. Pitfall traps are sometimes placed near ant nests to collect myrmecophilous species (Maruyama et al. 2008). We suggest that ant nests can be dug up and the resulting material placed on a large plastic sheet where it can be checked for beetles. Logs with ant nests can be broken open and contents placed onto a sheet or into a plastic box for examination. In NB, species associated with mound building ant species (Formica sp.) were collected by sifting the top layer of the nests in the spring and by placing small boards on nests and checking the underside of the board several days later (Webster et al. 2009). Aleocharinae were often found on the underside of the boards intermixed with the ants. The myrmecophilous Aleocharinae fauna in Canada is poorly known and many new records are awaiting discovery. Mammal and Bird Nests. Some Aleocharinae are frequently associated with mammal and bird nests. Some species can be found in debris in the entrances to dens or tunnels of woodchucks (also known as groundhogs), foxes, and other ground nesting mammals (Webster et al. 2016b; Brunke and Buffam 2018). These species are usually

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most common in the spring as they leave the dens for their dispersal flights. Some species are associated with microhabitats created by the activities of beavers, such as the nest material inside the lodge and others, including several Liogluta species, live amongst debris in dams (Klimaszewski et al. 2016). The nests of treehole nesting birds such as barred owls also harbor a number of Aleocharinae species. To sample these, one must climb the trees and remove the nest contents and then sift it (Webster et al. 2009, 2016b). Nests of other birds such as ospreys and hawks, swallows, and waterfowl have not been properly sampled in North America and may have an interesting fauna (Brunke and Buffam 2018). Mammal and bird nests are a poorly sampled habitat in general, and new species likely remain undiscovered. Compost. One very effective substrate for attracting Aleocharinae and many other Coleoptera species is backyard composter or a pile of organic debris such as corncobs and cornhusks. Corncobs and cornhusks are placed adjacent to a composter starting in July when corn becomes available. The corncobs decay and become moldy over time, and start to attract a wide variety of species as the season progresses. Many new records including some new Canadian records were found in this substrate (Webster et al. 2009, 2012, 2016b). Evening Flight Collections. Many Coleoptera species, including Aleocharinae, fly during the late afternoon and early evenings (1500–1800 h) on warm days with very light winds from early spring to early summer (sometimes even when snow patches are still on the ground in shaded sites in the spring). Adults are sometimes abundant during these flights. Presumably these are dispersal flights from overwintering sites. The beetles are easily collected by sweeping a butterfly net with a very fine mesh. The best sites are usually along forest edges and openings, river margins or along open forest trails. One can also

27

use a net mounted on a car (car net) to sample beetles (Klimaszewski et al. 2017). Sample Containers, Killing Agents, and Processing Non-trap Collections. Before collecting it is best to prepare a number of sample containers. Good sample containers to use for collections are plastic urine collection bottles. These can be obtained from pharmacies and a number of scientific supply venders. They are a good size and have labels on which one can record detailed habitat information. The containers are one-quarter filled with sifted sawdust from Populus. The best sawdust is obtained from trees that were cut with a chainsaw as it produces coarser pieces. Conifer sawdust should be avoided as resins in the wood gum up the specimens. Sawdust from other tree species is usually too fine, making it more difficult to find the specimens. Several drops of ethyl acetate are added to the sawdust along with a few drops of water to prevent dessication of the specimens. Aspirated specimens are added to the vials which can then be stored either at room temperature or preferably in a freezer when returned to the lab. Specimens stored this way can be preserved for several years without deterioration even at room temperature. To extract specimens from the jars, the sawdust is first removed and placed it in a small box under a fume hood or in another well ventilated place to allow the ethyl acetate to evaporate (this usually takes about 30 min). The sample is then examined under a microscope, and specimens are removed and cleaned in water. Sometimes very muddy specimens are best cleaned in soapy water while subjected to sonication for 1–3 min followed by rinsing in distilled water. Specimens treated in this way are in excellent shape and easily dissected. At this point it is best to mount (and dissect if necessary) the specimens as they are most flexible. Of course, specimens can also be preserved in 70% ethanol but this reduces flexibility and makes dissections more difficult later.

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Fig. 4.2 (a–l) Example of morphological diversity of Aleocharinae in British Columbia: (a) Gymnusa grandiceps Casey; (b) Aleochara bimaculata Gravenhorst; (c) Megocalea lemieuxi Klimaszewski; (d) Oxypoda sylvia Casey; (e) Gnypeta caerulea (C.R. Sahlberg); (f) Tachyusa americanoides Paśnik; (g)

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Holobus vancouveri Klimaszewski; (h) Myllaena insomnis Casey; (i) Liparocephalus cordicollis LeConte; (j) Autalia puncticollis Sharp; (k) Pontomalota opaca (LeConte); (l) Xenodusa reflexa (Walker). Scale bar ¼ 1 mm

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Fig. 4.3 Image of Myrmecocephalus concinnus (LeConte) in dorsal view, with body parts important to identification. After Klimaszewski et al. (2018)

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Fig. 4.4 (a–f) Mouthparts: (a) labrum; (b) left mandible; (c) right mandible; (d) labium and mentum; (e) left maxilla; (f) right maxilla. After Klimaszewski et al. (2018)

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Fig. 4.5 Image of pronotum and thorax of Gnypeta nigrella (LeConte) in ventral view, with important structures to identification. After Klimaszewski et al. (2018)

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Fig. 4.6 (a–e) Morphological divisions of male and female terminal segments and cross-section of abdomen. After Klimaszewski et al. (2018)

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Fig. 4.7 (a–e) Morphological divisions of the median lobe of aedeagus, paramere, and spermatheca. After Klimaszewski et al. (2018)

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Fig. 4.8 (a–i) Examples of the median lobe of aedeagus in lateral view of different species of aleocharine beetles: (a) Gymnusa grandiceps Casey; (b) Aleochara curtula (Goeze); (c) Aleochara litoralis (Mäklin); (d) Cratarea

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suturalis (Mannerheim); (e) Oxypoda convergens Casey; (f) Brachyusa helenae (Casey); (g) Cypha crotchii (Horn); (h) Myllaena fenyesi Bernhauer; (i) Liparocephalus brevipennis Mäklin. Scale bar ¼ 0.2 mm

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Material and Methods

Fig. 4.9 (a–l) Examples of spermatheca in lateral view of different species of aleocharine beetles: (a) Aleochara suffusa (Casey); (b) Devia prospera (Erichson); (c) Oxypoda sylvia Casey; (d) Gnypeta lohsei Klimaszewski; (e) Myllaena insomnis Casey; (f) Autalia puncticollis

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Sharp; (g) Stictalia brevicornis Casey; (h) Atheta graminicola (Gravenhorst); (i) Atheta prudhoensis (Lohse); (j) Nehemitropia lividipennis (Mannerheim); (k) Trichiusa pilosa Casey; (l) Xenodusa reflexa (Walker). Scale bar ¼ 0.2 mm

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Fig. 4.10 (a–f) Microdissections and preparation of genital structures for examination and diagnostics: (a) essential chemicals (Canada balsam, 95% ethanol, distilled water, vinegar or ammonium hydroxide); (b) tools and material (dissection needles, forceps, pins, mounting cars, plastic microslides for mounting genital structures

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in Canada balsam); (c) dried specimens; (d) ceramic dish with impressions for relaxing specimens in water with vinegar or ammonium hydroxide; (e) stereoscopic microscope and dissecting person (A. Gilbert); (f) removing apical part of abdomen with genitalia

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Material and Methods

Fig. 4.11 (a–f) Microdissections and preparation of genital structures for examination and diagnostics: (a) separating median lobe of aedeagus, parameres, male tergite and sternite VIII and pygidium; (b) proposed order of mounting male genital structures on microslide; (c) separating spermatheca, female tergite and sternite VIII

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and pygidium; (d) proposed order of mounting female genital structures on microslide; (e) dissected specimens mounted on card with genital structures in Canada balsam on microslide; (f) dissected specimens with mounted genital structures in Canada balsam on microslides ready for examination

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Fig. 4.12 (a–f) Image production: (a) example of image production system consisting of Nikon SMZ 1500 stereoscopic microscope, stacking and Adobe Photoshop software and operating person (C. Bourdon); (b) taking photographs of external habitus—head, pronotum, elytra,

4 Material and Methods

abdomen, antenna, and front, middle and hind legs; (c) taking photograph of median lobe of aedeagus; (d) taking photograph of tergite and sternite VIII; (e, f) cleaning background and undesirable morphological parts from median lobe of aedeagus

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Material and Methods

Fig. 4.13 (a–f) Image production: (a) transferring color median lobe of aedeagus to black and white image; (b) fixing damaged habitus parts and taking advantage of

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symmetrical structures; (c, d, e) putting habitus parts from different planes into one image; (f) producing a final image of habitus and genital structures

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Fig. 4.14 (a–f) Different traps and tools for collecting aleocharine beetles: (a) pitfall trap; (b) Luminoc trap; (c) Light trap (photo credit N. Winchester); (d) Flight

4 Material and Methods

Interception trap (Malaise trap) (photo credit Winchester, N.); (e) aspirator; (f) sifter

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41

Fig. 4.15 (a, b) Different collecting tools and methods: (a) Lindgren funnel (after Klimaszewski et al. 2018); (b) hand collecting with aspirator or forceps from sifted material in plastic box (after Klimaszewski et al. 2018)

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References Ahn K-J (1996) Revision of the intertidal aleocharine genus Tarphiota (Coleoptera: Staphylinidae). Entomol News 107:177–185 Ahn K-J (1997a) A review of Liparocephalus Mäklin (Coleoptera: Staphylinidae: Aleocharinae) with descriptions of larvae. Pan-Pac Entomol 73(2):79–92 Ahn K-J (1997b) Revision and systematic position of the intertidal genus Thinusa Casey (Coleoptera: Staphylinidae: Aleocharinae). Entomol Scand 28:75–81 Ahn K-J, Ashe JS (1995) Revision of the intertidal aleocharine genus Amblopusa Casey and description of the new genus Paramblopusa (Coleoptera: Staphylinidae). J N Y Entomol Soc 103(2):138–154 Ahn K-J, Ashe S (1996) Phylogeny of the intertidal aleocharine tribe Liparocephalini (Coleoptera: Staphylinidae). Syst Entomol 21:99–114 Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, London, New York, Washington, DC, 443 pp [publ. 28 Dec 2000 according to CRC Press] Benick G, Lohse GA (1974) Tribus 14 (Callicerini). In: Freude H, Harde KW, Lohse A (eds) Die Käfer Mitteleuropas. Band 5: Staphylinidae II (Hypocyphtinae und Aleocharinae) Pselaphidae. Goecke & Evers, Krefeld, 381 pp Bouchard P, Bousquet Y, Davies AE, Alonso-Zarazaga MA, Lawrence JF, Lyal CHC, Newton AF, Reid CAM, Schmitt M, Ślipiński SA, Smith ABT (2011) Family-group names in Coleoptera (Insecta). ZooKeys 88:1–972 Brunke, JA, Buffam J (2018). A review of Nearctic rove beetles (Staphylinidae) specialized on the burrows and nests of vertebrates. In: Biology of rove beetles (Staphylinidae): life history, evolution, ecology and distribution. Springer, Cham, 351 pp Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:119–206 Dybas LK, Dybas HS (1981) Coadaptation and taxonomic differentiation of sperm and spermathecae in featherwings beetles. Evolution 35(1):168–174 Elven H, Bachmann L, Gusarov VI (2010) Phylogeny of the tribe Athetini (Coleoptera: Staphylinidae) inferred from mitochondrial and nuclear sequence data. Mol Phylogenet Evol 57(1):84–100 Elven H, Bachmann L, Gusarov VI (2012) Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta). Zool Scr 41(6):617–636

4 Material and Methods Gollkowski V (2013) Über Köderfallen zum Nachweis von in Tierbauen usw. Lebenden Käfern. Entomol Nachr Ber 57:163–164 Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353 (1):134 Gusarov VI (2011) Case 3537. Athetini Casey, 1910 and Geostibina Seevers, 1978 (Insecta, Coleoptera, Staphylinidae, Aleocharinae): proposed conservation. Bull Zool Nomencl 68(1):54–60 Hammond HEJ (1997) Arthropod biodiversity from Populus coarse woody material in north-central Alberta: a review of taxa and collection methods. Can Entomol 129:1009–1033 Hanley RS, Ashe JS (2003) Techniques for dissecting adult aleocharine beetles (Coleoptera: Staphylinidae). Bull Entomol Res 93(1):11–18 Hébert C, Jobin L, Fréchette M, Pelletier G (2000) An efficient pit-light trap to study diversity. J Insect Conserv 4:191–202 Hlaváč P, Newton AF, Maruyama M (2011) World catalogue of the species of the tribe Lomechusini (Staphylinidae: Aleocharinae). Zootaxa 3075:1–151 Hughes CC, Johns RC, Sweeney JD (2014) A technical guide to installing beetle traps in the upper crown of trees. J Acad Entomol Soc 10(4):12–18 Jobin LJ, Coulombe C (1992) The Luminoc® insect trap. Forestry Canada, Quebec Region, Sainte-Foy, Quebec. Information Leaflet LFC 26E, 12 pp Klimaszewski J (1979) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae: Aleocharinae). Agriculture Canada Research Branch, Monograph No. 25:1–169 Klimaszewski J (1982) Studies of Myllaenini (Coleoptera: Staphylinidae, Aleocharinae). 1. Systematics, phylogeny, and zoogeography of Nearctic Myllaena Erichson. Can Entomol 114(3):181–242 Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Memoirs of the Entomological Society of Canada No. 129 [2] + 211 pp Klimaszewski J, Langor D, Savard K, Pelletier G, Chandler DS, Sweeney J (2007) Rove beetles (Coleoptera: Staphylinidae) in yellow birch-dominated stands of southeastern Quebec, Canada: diversity, abundance, and description of a new species. Can Entomol 139:793–833 Klimaszewski J, Langor DW, Work TT, Hammond JHE, Savard K (2008) Small and more numerous harvesting gaps emulate natural forest disturbances: a biodiversity test case using rove beetles (Coleoptera, Staphylinidae). Divers Distrib 14:969–982 Klimaszewski J, Webster RP, Langor D, Sikes D, Bourdon C, Godin B, Ernst C (2016) A review of Canadian and Alaskan species of the genus Liogluta Thomson, and descriptions of three new species (Coleoptera, Staphylinidae, Aleocharinae). In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline

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biodiversity and natural history data. ZooKeys [Special Issue] 573:217–256 Klimaszewski J, Struyve T, Bourdon C, Dorval J-A (2017) First record of Thecturota tenuissima Casey from Canada (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 702:19–25 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. 902 pp Lindgren BS (1983) A multiple funnel trap for scolytid beetles (Coleoptera). Can Enomol 115(3):299–302 Lohse GA, Klimaszewski J, Smetana A (1990) Revision of Arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44(2):121–202 Martin JEH (1977) The insects and arachnids of Canada. Part 1: Collecting, preparing, and preserving insects, mites, and spiders. Research Branch, Canada Department of Agriculture. Publication 1643. Ministry of Supply and Services Canada 1978. Kromar Printing Ltd Maruyama M, Klimaszewski J (2004) A new genus and species of the myrmecophilous Athetini, Paragoniusa myrmicae (Coleoptera: Staphylinidae: Aleocharinae) from Canada. Entomol Rev Japan 59(2):241–248 Maruyama M, Klimaszewski J (2006) Notes on myrmecophilous aleocharines (Insecta, Coleoptera, Staphylinidae) from Canada, with a description of a new species of Myrmoecia. Bull Nat Sci Mus Tokyo A 32(3):125–131 Maruyama M, Patrick LB, Klimaszewski J (2008) First record of the genus Myrmedonota Cameron (Coleoptera, Staphylinidae) from North America, with descriptions of two new species. Zootaxa 1716:35–43 Parker J (2016) Myrmecophily in beetles (Coleoptera): evolutionary patterns and biological mechanisms. Myrmecol News 22:65–108 Paśnik G (2010) Phylogeny and generic classification of Tachyusini (Coleoptera, Staphylinidae: Aleocharinae). Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, 129 pp Peck SB, Davies A (1980) Collecting small beetles with large-area ‘window’ traps. Coleopt Bull 34:237–239 Peschke K (1978) Funktionsmorphologische Untersuchungen zur Kopulation von Aleochara curtula Goeze (Coleoptera, Staphylinidae). Zoomorphologie 89(2):157–184 Schülke M, Smetana A (2015) Family Staphylinidae Latreille, 1802. In: Löbl I, Löbl D (eds) Catalogue of Palearctic Coleoptera, vol 2., Revised and updated edition. Hydrophiloidea-Staphylinoidea, Brill, Leiden & Boston, pp 304–1134

43 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana: Zoology 71: vi + 289 pp Skvarla MJ, Larson JL, Dowling APG (2014) Pitfalls and preservatives: a review. J Entomol Soc Ont 145:15–43 Spence JR, Niemelä JK (1994) Sampling carabid assemblages with pitfall traps: The madness and the method. Can Entomol 126:881–894 Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:171–248 Webster RP, Klimaszewski J, Sweeney JD, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:83–118 Webster RP, Anderson RS, Webster VL, Alderson CA, Hughes CC, Sweeney JD (2016a) New Curculionoidea records from New Brunswick, Canada with an addition to the fauna of Nova Scotia. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:367–386 Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016b) Further contribution to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including description of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216 Webster RP, Webster VL, Alderson CA, Hughes CC, Sweeney JD (2016c) Further contributions to the Coleoptera fauna of New Brunswick with an addition to the fauna of Nova Scotia, Canada. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:265–338 Yamamoto S, Maruyama M (2017) Phylogeny of the rove beetle tribe Gymnusini sensu n. (Coleoptera: Staphylinidae: Aleocharinae): implications for the early branching events of the subfamily: phylogeny of Gymnusini rove beetles. Syst Entomol 43 (1):183–199. https://doi.org/10.1111/syen.12267

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British Columbia: Hotspot of Canadian Biodiversity

Staphylinidae are the most diverse family in North America (Poole and Gentili 1996) and much of this diversity is associated with habitats found in forest ecosystems (Pohl et al. 2008). Rove beetles occur in all habitat types with the exception being extremely dry habitats (McCravy and Willand 2010). At the landscape scale species composition is dictated by forest type and species have strong affinities with forest microhabitats where moisture, temperature, substrate type, and debris accumulation are variables that promote high diversity in rove beetles (Pohl et al. 2008). British Columbia (BC) is a large and ecologically diverse province that covers 11 degrees of latitude and 25 degrees on longitude and spans over 948,000 km2 (Meidinger and Pojar 1991). Proximity to the coast and features associated with mountains and islands are primary drivers of the environment and provide a habitat template where forests dominate the vegetation (Fig. 5.16a–f). Broadly speaking, BC is a cool, moist, mountainous, forested region (Meidinger and Pojar 1991) where some of the largest remaining tracts of intact ancient coniferous forests in North America occur (Fig. 5.16a–f) (Winchester 1997). Forests are the dominant vegetation but there are extensive areas of tundra, wetlands, grasslands, and rolling scrub (Figs. 5.17, 5.18a, b, and 5.19). As a result, BC has a rich biota (Cannings and Cannings 1996) and greater biodiversity than other provinces and territories in Canada.

Staphylinids found in coastal habitats have been well-documented (Jeon and Ahn 2007; Majka et al. 2008; Frank and Ahn 2011). However, the extensive coastline in BC remains largely unexplored and provides numerous microhabitats that rove beetles should occur in and include pebble beaches, rocky shores, sandy beaches, muddy flats, brown, green or red species of seaweed that have been deposited onshore (Fig. 5.20a–f) (see Davis and Browne 1996). Coastal Staphylinidae found on rocky shores are often associated with features related to the reduction in rock size (e.g., boulders, pebbles, sand) and can be found associated with a variety of microhabitats that are provided by rocky substrates (Frank and Ahn 2011), tidal pools, and intertidal plant zonation (Topp and Ring 1988a, b; Majka et al. 2008) (Fig. 5.20a–f). Nine soil orders present in Canada exist in British Columbia, a testament to the varied parent material, climate, topography, and biotic factors that have shaped soil profiles over time. Valentine et al. (1978) provided the background for the generalized physiographic regions of BC. Using this information, Meidinger and Pojar (1991) designated 14 biogeoclimatic zones used to organize ecosystems at three levels of integration: local, regional, and chronological (MacKinnon et al. 1992). This habitat classification system was originally created for the purpose of managing forestry resources but it also is important in conservation and resource management.

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_5

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In BC, the protected areas strategy (PAS) goal was to protect 12% of the land area of the province by the year 2000 (British Columbia 1993). This 12% target has been achieved, however the major ecological regions in BC lack proportional representation under the PAS (Scudder 2000, 2003). Under the umbrella of PAS, hotspot analysis has been used as one of a set of tools to help set priorities for conservation planning (Reid 1998). This technique allows for the identification of areas of high biological diversity, and areas with large numbers of endemic species or species at risk. Although the term ‘biodiversity hotspot’ was originally coined to indicate areas with a large number of endemic species, it is now most commonly used with reference to regions of high species richness (Reid 1998). Endemic species hotspot, richness hotspot, and rarity hotspot are all terms used to indicate regions in BC for hotspot areas that contain a number of rare, endemic or unique species. Structural habitat complexities at local and regional scales are major determinants of arthropod species richness and this trend likely applies to most Staphylinidae. For Staphylinidae, and in particular the Aleocharinae, a current assessment for most habitats in BC is unavailable and many hotspot areas for arthropods in general, remain undocumented. An exception to this generalization are the detailed studies of Aleocharine rove beetles from the Carmanah Valley, an ancient Sitka spruce rainforest on the West Coast of Vancouver Island (Klimaszewski and Winchester 2002). Additionally, standardized sampling and exploration of terrestrial microhabitats (e.g., accumulation of organic debris, lichens, bark and branches) associated with conifer tree species has been investigated at several study sites on Vancouver Island, British Columbia, Canada (Winchester 1997; Winchester et al. 2008). We suggest that ancient temperate rainforests in British Columbia, Canada, are repositories for aleocharine biodiversity. Spatial heterogeneity of the soil environment is one explanation for the higher diversity in soil communities and Aleocharine rove beetles follow this pattern. British Columbia is a species rich province with 227 species being documented

5 British Columbia: Hotspot of Canadian Biodiversity

(Klimaszewski and Winchester 2002; Klimaszewski 2000). Species richness increases with the structural complexity that is related to architectural complexity (e.g., soil depth and development, plant associations) and the spatial complexity that is associated with the distribution of microhabitats. In coastal temperate rainforests, habitat patches are relatively permanent and large with respect to the temporal-spatial scale of the organism; a species could persist by remaining within the patch without dispersal from external sources. Generally there is a positive relationship between species richness and abundance with patch size and moisture content for many arthropods (see Lindo and Winchester 2009). Species-specific environmental tolerances combined with the differential dispersal abilities of species contribute to the nonrandom community patterns in forest habitats. Simply stated, habitat complexity as measured by patch size (e.g., depth and area) results in a greater retention of moisture and amelioration of temperature fluctuations. This in turn acts as a niche-selection filter, eliminating species from the regional pool that cannot tolerate rapid fluctuations in environmental conditions. The discovery of several new species in ancient conifer forests is not surprising. Several of these aleocharine species are confined to microhabitats found only in the ancient forest soils/debris. In particular, representatives of the aleocharine tribes Athetini, Homalotini, and Oxypodini are usually numerically dominant in the ancient forests of the Pacific Northwest. Species descriptions coupled with samples collected at multiple study sites and multiple microhabitats over a large spatial scale are starting to accumulate evidence to support the assumption that ancient forests contain several species restricted to these sites (Klimaszewski and Winchester 2002). For example, Paraleptonia pacei represents the first record of the genus Paraleptonia in North America (Klimaszewski and Winchester 2002). These characteristic trends in Staphylinidae community composition in ancient conifer forests may be indicative of all intact conifer rainforests where soils and other microhabitats (e.g. lichens) are well developed (Fig. 5.16a–f).

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British Columbia: Hotspot of Canadian Biodiversity

Fig. 5.16 (a–f) Examples of intact ancient forests, Vancouver Is., BC: (a–c) red cedar rain forest (Pacific Rim National Park Reserve); (d, e) Douglas-fir forest

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(Cathedral Grove); (f) mixed forest (Cowichan Valley Regional District). Photo credit J. Klimaszewski

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5 British Columbia: Hotspot of Canadian Biodiversity

Fig. 5.17 Wolf Ear’s Peak near Nakusp, West Kootenay, Interior B.C. Photo credit B. Costanzo

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British Columbia: Hotspot of Canadian Biodiversity

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Fig. 5.18 (a, b) Landscape view: (a) Clayoquot Sound, West Coast, Vancouver Island; (b) Osoyoos Lake from Mt. Kobau, Okanagan Valley, BC. Photo Credits N. Winchester (a), B. Costanzo. (b)

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5 British Columbia: Hotspot of Canadian Biodiversity

Fig. 5.19 Paradise Meadows, Courtenay, Vancouver Island. Photo credits B. Costanzo

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British Columbia: Hotspot of Canadian Biodiversity

Fig. 5.20 (a–f) Coastal habitats, Vancouver Is., BC: (a) seashore landscape; (b) pebble beach (Nanoose Bay); (c) muddy flats (Nanaimo); (d, f) rocky shores (Nanoose

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Bay); (e) sandy beach with debris (Nanaimo). Photo credit J. Klimaszewski

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References British Columbia (1993) A protected areas strategy for British Columbia. Province of British Columbia, Victoria, BC Cannings R, Cannings S (1996) British Columbia: a natural history. Greystone Books, Vancouver, BC. 310 pp Davis DS, Browne S (1996) The natural history of Nova Scotia, vol 1. Topics and habitats. The Nova Scotia Museum and Nimbus Publishing, Halifax, 518 pp Frank JH, Ahn K-J (2011) Coastal Staphylinidae (Coleoptera): a worldwide checklist, bibliography and natural history. ZooKeys 103:1–98 Jeon M-J, Ahn K-J (2007) Descriptions of late instars of three littoral Cafius species (Coleoptera: Staphylinidae) by association of life stage with DNA sequences. Fla Entomol 90(3):465–474 Klimaszewski J (2000) Diversity of the rove beetles in Canada and Alaska (Coleoptera Staphylinidae). Mémoires de la Société royale belge d’Entomologie, 126 pp Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mémoire de la Societé royale belge d’Entomologie 40:3–126 Lindo Z, Winchester NN (2009) Spatial and environmental factors contributing to patterns in arboreal and terrestrial oribatid mite diversity across spatial scales. Oecologia 160:817–825 MacKinnon A, Meidinger D, Klinka K (1992) Use of the biogeoclimatic ecosystem classification system in British Columbia. Forest Chron 68:100–120 Majka CG, Klimaszewski J, Randolph RF (2008) The coastal rove beetles (Coleoptera, Staphylinidae) of Atlantic Canada: a survey and new records. ZooKeys 2:115–150 McCravy K, Willand J (2010) Habitat associations of macro-Staphylinids (Coleoptera: Staphylinidae) at Alice L. Kibbe Life Science Station, Hancock County, Illinois. Great Lakes Entomol 43:1–4

5 British Columbia: Hotspot of Canadian Biodiversity Meidinger D, Pojar J (1991) Ecosystems of British Columbia. B.C. Ministry of Forests, Victoria, BC Pohl G, Langor D, Klimaszewski J, Work T, Paquin P (2008) Rove beetles (Coleoptera: Staphylinidae) in northern Nearctic forests. Can Entomol 140:415–436 Poole RW, Gentili P (1996) Nomina Insecta Nearctica, Coleoptera, Strepsiptera, vol 1. Entomological Information Services, Rockville, MD Reid WV (1998) Biodiversity hotspots. Trends Ecol Evol 13(7):275–280 Scudder GGE (2000) Species at risk, conservation strategies, and ecological integrity. In: Darling LM (ed) Proceedings of a conference on the biology and management of species and habitats at risk. Kamloops, BC. 15–19 Feb 1999. Volume B.C. Ministry of Environment, Lands and Parks, Victoria, B.C. and University College of the Cariboo, Kamloops, BC, pp 1–12 Scudder GGE (2003) Biodiversity conservation and protected areas in British Columbia. Available from http://www.zoology.ubc.ca/biodiversity/hotspots/ hotspot_report.pdf Topp W, Ring RA (1988a) Adaptations of Coleoptera to the marine environment. I. Observations on rove beetles (Staphylinidae) from sandy beaches. Can J Zool 66:2464–2468 Topp W, Ring RA (1988b) Adaptations of Coleoptera to the marine environment. II. Observations on rove beetles (Staphylinidae) from rocky shores. Can J Zool 66:2469–2474 Valentine KWG, Sprout PN, Baker TE, Lavkulich LM (1978) The soil landscapes of British Columbia. B.C. Ministry of Environment, Resource Analysis Branch, Victoria, BC Winchester NN (1997) Arthropods of coastal old-growth Sitka spruce forests: conservation of biodiversity with special reference to the Staphylinidae. In: Watt AD, Stork NE, Hunter MD (eds) Forests and insects. Chapman and Hall, London, pp 363–376 Winchester NN, Lindo Z, Behan-Pelletier VM (2008) Oribatid mite communities in the canopy of montane Abies amabilis and Tsuga heterophylla trees on Vancouver Island, British Columbia. Environ Entomol 37:464–471

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Faunal Analysis and Discussion

Aleocharinae is a diverse subfamily of Staphylinidae with 604 species within 17 tribes recorded from Canada (present data), approximately 36% of the known rove beetle fauna (Bousquet et al. 2013). Nearly 70% of Aleocharinae species have been found in eastern Canada where the biodiversity of this group is well documented compared to western and central Canada (Fig. 6.21; Table 6.3) (Klimaszewski et al. 2018). Of the seven eastern jurisdictions, New Brunswick has been extensively studied (Webster et al. 2016a, b), with 297 species recorded (Table 6.4), 49% of the total fauna known from Canada. Aleocharinae species are underrepresented in central Canada as well as the northcentral USA because those areas have not been sufficiently sampled to date. Within western Canada, 317 species have been recorded, 52.6% of the total Aleocharinae fauna, of which 91% are native species. British Columbia (BC), a biodiversity hotspot on the western coast of Canada, has 227 recorded species, 37.6% of the total fauna documented in Canada. Forty-eight of these species have been recorded only from BC, while 22 are adventive species. New provincial records for BC are documented for 36 species. The number of Aleocharinae species found in BC is predicted to increase as additional surveys are conducted, especially in the northern region of the province. Aleocharinae species can be characterized based on their distribution patterns across Canada and the USA (Fig. 6.22; Table 6.3),

with some species documented only in the eastern, central, or western jurisdictions. Nearly 30% of species are found only in eastern Canada (Fig. 6.22b), with an additional 10.9% characterized as having an east-central distribution. Forty-two species (7.0%) are found only in central Canada (Fig. 6.22c), but this number likely is an underestimate due to low sampling efforts. Over 19% of species are recorded only from western Canada and the USA (Fig. 6.22d), including BC, Yukon, Alaska, Washington, Oregon, California, and Nevada, with another 5.5% of species having a west-central distribution. Transcontinental species (28.0%, 169 spp.) have broad distributions across eastern and western jurisdictions (Fig. 6.22a). Twenty-six species characterized as transcontinental are adventive species introduced from the Palearctic region, primarily as a consequence of trade with Europe and Asia (Klimaszewski et al. 2013). In BC, 111 of these broadly distributed species have been documented, with 18% being adventive to Canada.

Dominant Aleocharinae Tribes in British Columbia Of the 17 Aleocharinae tribes found in Canada, 14 are represented in western and west-central Canada, including BC (Table 6.5). Dominant Aleocharinae tribes in BC (based on number of species) are the Athetini (44.5%), Oxypodini

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_6

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Faunal Analysis and Discussion

Table 6.3 Large-scale patterns in the distributions of Aleocharinae tribes, genera, and species in Canada and the continental United States of America

No. of tribes No. of genera No. of species (% of total) No. of adventive species (% of total)

Transcontinental 13 58 169 (28.0%)

Eastern 15 61 178 (29.5%)

EastCentral 10 31 66 (10.9%)

Central 6 21 42 (7.0%)

WestCentral 10 22 33 (5.5%)

Western 11 44 115 (19.1%)

26 (15.4%)

15 (8.4%)

3 (4.5%)

3 (7.1%)

1 (3.0%)

2 (1.7%)

Transcontinental distribution (all Canadian Provinces and Territories), Eastern distribution (Prince Edward Island, Nova Scotia, Labrador, Newfoundland, New Brunswick, Quebec, and Ontario), East-Central distribution (Prince Edward Island, Nova Scotia, Labrador, Newfoundland, New Brunswick, Quebec, Ontario, Manitoba, Nunavut, Saskatchewan, Alberta, and Northwest Territories), Central distribution (Northwest Territories, Alberta, Saskatchewan, Nunavut, and Manitoba), West-Central distribution (Yukon, British Columbia, Northwest Territories, Alberta, Saskatchewan, Nunavut, and Manitoba), and Western distribution (Yukon and British Columbia). Numbers in parentheses indicate the percentage of the total species recorded from Canada (603 spp.) and the percentage of the total adventive species recorded by distribution type

(19.4%), Aleocharini (10.1%), Homalotini (6.6%), and Tachyusini (5.3%). The Liparocephalini are found west of the Rocky Mountains and are known only from BC in Canada. The Athetini are the largest Aleocharinae tribe (Elven et al. 2010; Newton et al. 2000) and the dominant tribe in Canada with 283 species in 41 genera, 28% of which are broadly distributed transcontinental species. Moreover, the highest number of adventive (20 spp.) and Holarctic (18 spp.) species recorded from Canada are found within the Athetini tribe. Similar to these continental patterns, Athetini is the dominant tribe in BC, comprising the greatest number of Aleocharinae genera (30) and species (101 spp.). Nearly half of these species (49 spp.) have transcontinental distributions, of which five are Holarctic species. Within the Athetini, 24 species are known only from BC, and 21 species are new provincial records. Atheta is the most speciose genus with 44 species found in BC, followed by Liogluta (7 spp.) and Philhygra (7 spp.). All species distribution types (Fig. 6.22) are represented within Atheta, including western (25 spp.; 19 spp. in BC), west-central (5 spp.; 3 spp. in BC), and transcontinental (33 spp.; 21 spp. in BC). Of the eight adventive Atheta species found in Canada, only Atheta longicornis (Gravenhorst) has been reported from BC. The Athetini genera Adota, Ousipalia, Paraleptonia,

Pontomalota, Psammostiba, Tarphiota, and Thinusa are found only in western Canada including BC, while Alevonota, Callicerus, Stethusa, and Tomoglossa are absent from BC and known only from eastern Canada. Tribe Oxypodini is represented by 89 species within 26 genera throughout Canada. Nearly 30% of species have a transcontinental distribution within this tribe, while western (25.8%) and eastern (22.5%) distributions are similarly represented among species. Within BC, the Oxypodini are a diverse tribe containing 44 species within 18 genera. Fifteen species are known only from BC, and 11 are new provincial records. Most species (56.8%) are found within the genera Oxypoda (19 spp.) and Neothetalia (6 spp.), while the other 16 genera are represented by only one or two species. Species of Oxypoda are well represented across Canada except the poorly sampled jurisdictions Nunavut and Prince Edward Island which have low total species records (Fig. 6.21). All species distribution types (Fig. 6.22) are represented within Oxypoda, including western (13 spp.; 10 spp. in BC), west-central (2 spp.; both in BC), and transcontinental (12 spp.; 7 spp. in BC). Neothetalia is primarily known from BC, but one broadly distributed species, Neothetalia canadiana Klimaszewski, is recorded from eastern Canada as well as BC and Yukon. The Oxypodini genera Betocalea, Megocalea, Metocalea, and Ocalea

YT 9 35 134 (22.2%) 6 (4.5%)

BC 14 78 227 (37.6%) 22 (9.7%)

NT 7 23 62 (10.3%) 0 (0%)

AB 12 48 116 (19.2%) 14 (12.1%)

SK 12 40 126 (20.9%) 17 (13.5%)

NU 3 7 13 (2.2%) 1 (7.7%)

MB 9 32 118 (19.5%) 10 (8.5%)

ON 13 74 245 (40.6%) 31 (12.7%)

QC 12 65 203 (33.6%) 29 (14.3%)

NB 16 84 297 (49.2%) 35 (11.8%)

NF 10 46 164 (27.2%) 26 (15.9%)

LB 10 28 97 (16.1%) 9 (9.3%)

NS 11 48 125 (20.7%) 25 (20.0%)

PE 7 16 24 (4.0%) 9 (37.5%)

YT Yukon, BC British Columbia; NT Northwest Territories, AB Alberta, SK Saskatchewan, NU Nunavut, MB Manitoba, ON Ontario, QC Quebec, NB New Brunswick, NF Newfoundland, LB Labrador, NS Nova Scotia, and PE Prince Edward Island. Numbers in parentheses indicate the percentage of the total species recorded from Canada (603 spp.) and the percentage of the total adventive species recorded by jurisdiction

No. of tribes No. of genera No. of species (% of total) No. of adventive species (% of total)

Table 6.4 Number of Aleocharinae tribes, genera, and species recorded from each Canadian jurisdiction

6 Faunal Analysis and Discussion 55

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Table 6.5 Number of Aleocharinae genera, total species, adventive species, and Holarctic species by tribe recorded in the Canadian province British Columbia compared to Tribe Aleocharini Athetini Autaliini Boreocyphini Diglottini Falagriini Gymnusini Homalotini Hoplandriini Hypocyphtini Liparocephalini Lomechusini Myllaenini Oxypodini Placusini Tachyusini Taxicerini Total

No. of genera 1 (2) 30 (41) 1 (1) 0 (1) 1 (2) 4 (7) 1 (2) 6 (18) 0 (2) 3 (3) 4 (4) 3 (6) 1 (1) 18 (26) 1 (2) 4 (4) 0 (1) 78 (123)

No. of species 23 (43) 101 (283) 3 (3) 0 (1) 1 (2) 4 (13) 3 (12) 15 (73) 0 (5) 3 (9) 5 (5) 3 (16) 4 (11) 44 (89) 6 (9) 12 (29) 0 (1) 227 (604)

are found only in western Canada including BC, while Apimela, Gennadota, Gyronycha, and Ilyobates are absent from BC and found only in eastern Canada. Tribe Aleocharini is represented by 43 species within two genera, Amarochara (4 spp.) and Aleochara (39 spp.), in Canada. Twenty-three species of Aleochara are found in BC, but species of Amarochara are known only from eastern and east-central Canada. Nearly 70% of Aleochara species from BC have transcontinental distributions, with six of those being adventive species. Although most Aleochara species are broadly distributed across Canada, 26.1% and 4.3% have western and west-central distributions, respectively. Two species, Aleochara carmanah Klimaszewski and Aleochara quadrata Sharp, are known only from BC. Tribe Homalotini is represented by 73 species within 18 genera in Canada. Over 40% of these species are found in the east with an additional 27.4% having an east-central distribution. Fewer species are distributed in the west (9.6%) and west-central (2.7%) regions of Canada. Fifteen species within six genera are found in BC,

Faunal Analysis and Discussion

the number recorded across all jurisdictions in Canada (indicated in parentheses) No. of adventive species 6 (8) 7 (20) 2 (2) 0 (0) 0 (0) 1 (3) 0 (1) 0 (2) 0 (0) 1 (4) 0 (0) 0 (1) 0 (0) 3 (6) 2 (2) 0 (0) 0 (1) 22 (50)

No. of holarctic species 1 (1) 6 (18) 0 (0) 0 (0) 0 (0) 0 (0) 1 (2) 1 (1) 0 (0) 0 (0) 0 (0) 0 (0) 0 (0) 1 (1) 0 (0) 1 (4) 0 (0) 11 (27)

20.5% of the species known from this tribe in Canada. One genus (Leptusomorpha Klimaszewski and Hoebeke) and three species, Leptusomorpha claudiae Klimaszewski, Stictalia carlottae Casey, and Stictalia kranabetteri Klimaszewski and Godin, are known only from BC. Six species, primarily in the genus Gyrophaena, are broadly distributed across Canada. The genera Agaricomorpha, Pleurotobia, and Thecturota do not occur in BC, as they are found only in eastern Canada. Tribe Tachyusini is represented by 29 species within four genera in Canada. Over 30% of these species are broadly distributed throughout Canada, and another 27.6% have west or westcentral distributions. Twelve species within Tachyusini are recorded from BC, which includes all four genera, of which the most speciose are Gnypeta (6 spp.) and Tachyusa (3 spp.). Three species, Brachyusa americana (Fenyes), Gnypeta baranowskii Klimaszewki, and Gnypeta brevicornis Casey, are known only from BC. Brachyusa helenae (Casey) is a broadly distributed species known from eastern and western jurisdictions, with BC as a new provincial record.

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Faunal Analysis and Discussion

Of the Aleocharinae tribes recorded from Canada, one tribe is known only from BC, while three tribes are reported only from eastern Canada. The Liparocephalini are known only west of the Rocky Mountains and are reported from BC and the USA states Alaska, Washington, Oregon, and California. Species within Liparocephalini are found along the Pacific coast because they are intertidal habitat specialists (Ahn et al. 2010). Tribe Liparocephalini is represented by five species within four genera, Amblopusa, Diaulota, Liparocephalus (2 spp.), and Paramblopusa. Diglottini is a new tribe record for BC, with Bryothinusa catalinae Casey being a new provincial record. Previously, Tribe Diglotiini was known in Canada from one species, Diglotta mersa (Haliday), recorded in New Brunswick (Klimaszewski et al. 2008), but the distribution of this species remains uncertain. Within the tribe Hypocyphtini, the genus Holobus and species Holobus vancouveri Klimaszewski are known only from BC. Tribes Boreocyphini, Hoplandriini, and Taxicerini are not currently known from BC and are primarily found in eastern Canada, although one transcontinental Hoplandriini species, Hoplandria lateralis (Melsheimer), is reported from California. Eleven Holarctic species are recorded in BC, for which ten have a transcontinental distribution across Canada and six are in the Tribe Athetini.

Adventive Species of Aleocharinae Fifty adventive species within 11 Aleocharinae tribes have been documented in Canada (Table 6.5). Over 50% of these species have transcontinental distributions across Canada, whereas 36% and 6% are found only in the east and west, respectively. Five adventive species are recorded from only one Canadian jurisdiction (2 spp. from Saskatchewan; 1 sp. from Manitoba; 1 sp. from Ontario; and 1 sp. from Newfoundland). Twenty-two adventive species within seven tribes are recorded from BC (Table 6.5), comprising 9.7% of the BC Aleocharinae fauna (Table 6.4). Most of these species belong to the

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Athetini (31.8%) and Aleocharini (27.3%) tribes. Two species are documented only in western Canada and the USA, Aloconota cambrica (Wollaston) and Autalia puncticollis Sharp. A third species, Atheta subsinuata (Erichson), has a west-central distribution, but is currently not known from BC. The remaining 20 adventive species found in BC have transcontinental distributions and are found across Canada and the USA. The most broadly distributed species include Mocyta fungi (Gravenhorst) (reported from 13 Canadian jurisdictions and 10 USA states), Dinaraea angustula (Gyllenhal) (11 Canadian jurisdictions and 3 USA states), Aleochara fumata Gravenhorst (10 Canadian jurisdictions and 24 USA states), Amischa analis (Gravenhorst) (10 Canadian jurisdictions and 11 USA states), and Aleochara bilineata Gyllenhal (10 Canadian jurisdictions and 9 USA states). Four adventive species, A. analis, D. angustula, Nehemitropia lividipennis (Mannerheim), and Oligota parva Kraatz, are new provincial records for BC. All adventive Aleocharinae species recorded from BC are of Palearctic origin, likely due to the similarities in climate between the Palearctic and northern Nearctic and the long history of commerce between these two regions (Lindroth 1957; Sailer 1978, 1983; Klimaszewski et al. 2007, 2013; Wheeler Jr and Hoebeke 2017; Klimaszewski and Brunke 2018). Most species were presumably introduced into Canada accidentally as a consequence of trade with Europe and Asia. Humans have inadvertently transported Aleocharinae species into regions outside of their native range through several mechanisms including: (1) the transportation of dry ballast (e.g. rocks, sand, and soil) in the holds of ships and unloading of these materials at ports of entry (Lindroth 1957); (2) importation of soil from agricultural and horticultural plants as well as tree nursery stock; and (3) importation of livestock and their associated food, bedding, and manure (Klimaszewski et al. 2013, 2010; Klimaszewski and Brunke 2018). The eastern and western coasts of Canada continue to be major points of entry for adventive beetle species, and consequently the high-traffic coastal

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jurisdictions have more adventive Aleocharinae species than the northern or central provinces and territories (Tables 6.3 and 6.4). In some cases, separate introductions have been hypothesized for adventive species on the Pacific and Atlantic coasts of Canada, for example Aleochara curtula (Goeze) and Crataraea suturalis (Mannerheim), but additional genetic analyses are needed to test these hypotheses (Klimaszewski 1984; Klimaszewski et al. 2007, 2013). Inadvertent movement of Aleocharinae species to new regions is facilitated by their high abundance in terrestrial ecosystems, small body size, and diverse habitat associations such as in litter and soil layers, on mushrooms, and among decaying organic matter, dung, and carrion. Moreover, the eurytopic and synanthropic nature of some Aleocharinae species further promote successful introduction and establishment in new regions. One species, A. bilineata, was intentionally introduced into Canada in the 1950s for the biological control of cabbage root maggot, Delia radicum (Linnaeus), but was already present in Ottawa, Ontario since the early 1900s described under the synonymic name, Baryodma ontarionis Casey (Hemachandra et al. 2005; Klimaszewski et al. 2018).

Predictions for Aleocharinae Diversity in British Columbia British Columbia is considered a biodiversity hotspot in western Canada, but the northern and coastal regions of this province remain largely unexplored. Therefore, the number of Aleocharinae species present in BC is expected to increase from 227 as sampling improves over time. Based on our characterization of species’ distributions within the surrounding Canadian jurisdictions and USA states, we predicted that an additional 50–90 Aleocharinae species may be present but currently undocumented in BC, especially in the northern region. Of the 169 transcontinental species recorded in Canada, 58 of these species are not currently

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Faunal Analysis and Discussion

documented in BC. Forty-six transcontinental species occur in the jurisdictions bordering BC (38 spp. to the north in the Yukon Territory and 6 spp. to the south in the USA state of Washington). Five of these broadly distributed species are adventive to Canada and the USA, Aleochara lata Gravenhorst (found in Yukon), Aloconota sulcifrons (Stephens), Atheta amicula (Stephens), Myrmecocephalus concinnus (Erichson), and Homalota plana (Gyllenhal) (all found in Washington). Thirty-three Aleocharinae species with western or west-central distributions are not currently recorded from BC. Ten species with west-central distributions are recorded from northern jurisdictions, including Yukon, Alaska, Northwest Territories, and/or Nunavut. Four species, Boreophilia caseyi Lohse, Mocyta amblystegii (Brundin), Gymnusa konopackii Klimaszewski, and Gnypeta ashei Klimaszewski, are recorded from all four of the aforementioned northern jurisdictions and represent the most likely westcentral species present in BC. One adventive species, A. subsinuata has been recorded from Yukon and Saskatchewan. Twenty-two species of Aleocharinae with western distributions are found in the Yukon Territory or Alaska but are not currently documented in BC. Atheta onthophaga (Lohse), Atheta sparreschneideri Munster, Boreophilia venti Lohse, Dinaraea planaris (Mäklin), Emmelostiba microptera (Lohse), and Liogluta granulosa Lohse, all within the Athetini tribe, are recorded from Yukon and Alaska. Despite the growth in knowledge of the Canadian Aleocharinae fauna in the last 20 years, we have estimated that the total number of species in BC is closer to 300, a substantial increase from the current record of 227 species. However, this prediction is conservative because it is based on the documented Aleocharinae fauna in Canada and their current described distributions, which are both likely underestimated. The total number of Aleocharinae species documented in BC and across Canada will surely increase with additional faunal surveys and research interest.

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Faunal Analysis and Discussion

Fig. 6.21 Large-scale patterns of Aleocharinae species richness recorded across Canada and those species that also occur in the United States of America. Shading indicates the current number of Aleocharinae species

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recorded from each Canadian jurisdiction, with the number of species increasing from light to dark blue. Map # Emily M. Trejo Sypolt

considered to have a transcontinental distribution if recorded from the east and west. Map # Emily M. Trejo Sypolt

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Fig. 6.22 Distributions of Aleocharinae species recorded from Canada and the United States of America. Species distributions can be characterized as transcontinental (a), eastern (b), east-central, central (c), west-central, and western (d). A species was

60 Faunal Analysis and Discussion

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Faunal Analysis and Discussion

References Ahn K-J, Jeon M-J, Branham MA (2010) Phylogeny, biogeography and the stepwise evolutionary colonization of intertidal habitat in the Liparocephalini based on morphological and molecular characters (Coleoptera: Staphylinidae: Aleocharinae). Cladistics 26:344–358. https://doi.org/10.1111/j.1096-0031. 2009.00290.x Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013) Checklist of beetles (Coleoptera) of Canada and Alaska, 2nd edn. Pensoft, Sofia Elven H, Bachmann L, Gusarov VI (2010) Phylogeny of the tribe Athetini (Coleoptera: Staphylinidae) inferred from mitochondrial and nuclear sequence data. Mol Phylogenet Evol 57:84–100. https://doi.org/10.1016/j. ympev.2010.05.023 Hemachandra KS, Holliday NJ, Klimaszewski J, Mason PG, Kuhlmann U (2005) Erroneous records of Aleochara bipustulata from North America: an assessment of the evidence. Can Entomol 137:182–187. https://doi.org/10.4039/n04-037 Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Mem Ent Soc Can 129, [2] + 211 pp. https://doi.org/10.4039/entm116129fv Klimaszewski J, Brunke AJ (2018) Canada’s adventive rove beetle (Coleoptera, Staphylinidae) fauna: a longterm case study on the detection, origin, introduction pathways, and dynamic distribution of non-native beetles. In: Betz O, Irmler U, Klimaszewski J (eds) Biology of Rove Beetles (Staphylinidae): life history, evolution, ecology and distribution. Springer, Cham, pp 65–79 Klimaszewski J, Assing V, Majka CG, Pelletier G, Webster RP, Langor D (2007) Records of adventive aleocharine beetles (Coleoptera: Staphylinidae: Aleocharinae) found in Canada. Can Entomol 139:54–79 Klimaszewski J, Webster RP, Assing V, Savard K (2008) Diglotta mersa (Haliday) and Halobrecta flavipes Thomson, two new species for the Canadian fauna (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 2:175–188 Klimaszewski J, Langor D, Majka CG, Bouchard P, Bousquet Y, LeSage L, Smetana A, Sylvestre P, Pelletier G, Davies A, DesRochers P, Goulet H,

61 Webster R, Sweeney J (2010) Review of adventive species of Coleoptera (Insecta) recorded from eastern Canada. Pensoft, Sofia-Moscow, 272 pp Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013) Synopsis of 10024 adventive species of Coleoptera (Insecta) recorded 10025 from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia-Moscow, 360 pp Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetlesof eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Lindroth CH (1957) The faunal connections between Europe and North America. Wiley, New York. 344 pp Newton AF, Thayer MK, Ashe JS, Chandler DS (2000) 22. Staphylinidae Latreille, 1802. In: Arnett RH, Thomas MC (eds) American Beetles, Archostemata, myxophaga, adephaga, polyphaga: staphyliniformia, vol 1. CRC Press, LLC, Boca Raton, FL, pp 272–418 Sailer RI (1978) Our immigrant insect fauna. Bull Entomol Soc Am 24:3–11 Sailer RI (1983) History of insect introductions. In: Wilson CL, Graham CL (eds) Exotic plant pests and North American agriculture. Academic, New York, pp 15–38. 522 pp Webster RP, Bouchard P, Klimaszewski J, Sweeney JD (2016a) History of Coleoptera collecting in New Brunswick, Canada: advancing our knowledge of the Coleoptera fauna in the early 21st century. ZooKeys 573:1–18 Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016b) Further contributions to the Aleocharinae (Coleoptera, Staphylinidae) fauna of New Brunswick and Canada including descriptions of 27 new species. ZooKeys 573:85–216 Wheeler AG Jr, Hoebeke ER (2017) Adventive (non-native) insects and the consequences for science and society of species that become invasive. In: Foottit RG, Adler PH (eds) Insect biodiversity: science and society, vol 1, 2nd edn. Wiley-Blackwell, Chichester, pp 641–712

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

[
Holarctic species, {adventive species, {species of uncertain distribution status—adventive in Canada or Holarctic, species without marks are native; numbers in brackets indicate number of species per taxon] GYMNUSINI [3]: Gymnusa (3); ALEOCHARINI [24]: Aleochara (23); OXYPODINI [44]: Alfocalea (1), Betocalea (1), Blepharhymenus (1), Calodera (1), Crataraea (1), Devia (1), Gnathusa (2), Megocalea (1), Meotica (1), Metocalea (1), Mniusa (1), Neoisoglossa (1); Neothetalia (6), Ocalea (1), Ocyusa (1), Oxypoda (19), Parocalea (1), Phloeopora (3); TACHYUSINI [12]: Brachyusa (2), Gnypeta (6), Paradilacra (1), Tachyusa (3); HYPOCYPHTINI [3]: Cypha (1), Holobus (1), Oligota (1); MYLLAENINI [4]: Myllaena (4); DIGLOTTINI [1]: Bryothinusa (1); LIPAROCEPHALINI [5]: Amblopusa (1), Diaulota (1), Liparocephalus (2), Paramblopusa (1); AUTALIINI [3]: Autalia (3); HOMALOTINI [15]: Encephalus (1), Gyrophaena (5), Leptusa (1), Leptusomorpha gen. n. (1), Silusa (3), Stictalia (4); PLACUSINI [6]: Placusa (6); ATHETINI [101]: Adota (1), Aloconota (2), Amischa (1), Atheta (44), Boreophilia (1), Boreostiba (2), Clusiota (1), Dalotia (1), Dinaraea (4), Dochmonota (1); Earota (1), Geostiba (1), Goniusa (1), Liogluta (7), Lypoglossa (2), Mocyta (3), Nehemitropia (1), Ousipalia (1), Paragoniusa (1), Paraleptonia (1), Pelioptera (1), Philhygra (7), Pontomalota

(1), Psammostiba (2), Schistoglossa Seeversiella (1), Strigota (2), Tarphiota Thinusa (2), Trichiusa (2); FALAGRIINI Bryobiota (1); Cordalia (1), Falagria Myrmecocephalus (1); LOMECHUSINI Myrmoecia (1), Xenodusa (1), Zyras (1).

(4), (2); [4]: (1), [3]:

Family STAPHYLINIDAE Latreille, 1802 (Rove beetles) Subfamily ALEOCHARINAE Fleming, 1821 Tribe Gymnusini Heer, 1839 Genus GYMNUSA Gravenhorst, 1806 Brevicollis Species Group 1. G. atra Casey, 1911
Grandiceps Species Group 2. G. grandiceps Casey, 1915 Variegata Species Group 3. G. pseudovariegata Klimaszewski, 1979 Tribe Aleocharini Fleming, 1821 Subtribe Aleocharina Fleming, 1821 Genus ALEOCHARA Gravenhorst, 1802 (Subgenus ALEOCHARA Gravenhorst, 1802)

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_7

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

4. A. assiniboin Klimaszewski, 1985 5. A. curtula (Goeze, 1777){ 6. A. gracilicornis Bernhauer, 1901b
7. A. tahoensis Casey, 1906

Subtribe Microglottina Fenyes, 1918

(Subgenus CALOCHARA Casey, 1906) 8. A. laramiensis (Casey, 1906) 9. A. rubricalis (Casey, 1911) 10. A. villosa Mannerheim, 1830{

Subtribe Oxypodina C.G. Thomson, 1859

(Subgenus COPROCHARA Mulsant and Rey, 1874) 11. A. bilineata Gyllenhal, 1810{ 12. A. bimaculata Gravenhorst, 1802 13. A. suffusa (Casey, 1906)
14. A. sulcicollis Mannerheim, 1843 15. A. verna Say, 1833

Genus DEVIA Blackwelder, 1952 31. D. prospera (Erichson, 1839)


(Subgenus EMPLENOTA Casey, 1884) 16. A. curtidens Klimaszewski, 1984 17. A. litoralis (Mäklin, 1853) 18. A. pacifica (Casey 1893)

Genus MNIUSA Mulsant and Rey, 1875 34. M. yukonensis (Klimaszewski and Godin, in Klimaszewski et al. 2012)

(Subgenus TINOTUS Sharp, 1883) 19. A. acerba (Casey, 1911). New generic combination, Lectotype designation 20. A. morion Gravenhorst, 1802{ (Subgenus XENOCHARA Mulsant and Rey, 1874) 21. A. carmanah Klimaszewski, in Klimaszewski and Winchester 2002 22. A. castaneipennis Mannerheim, 1843 23. A. quadrata Sharp, 1883 24. A. fumata Gravenhorst, 1802{ 25. A. lacertina Sharp, 1883 26. A. lanuginosa Gravenhorst, 1802{ Tribe Oxypodini C.G. Thomson, 1859 Subtribe Dinardina Mulsant and Rey, 1873b Genus BLEPHARHYMENUS Solier, 1849 27. B. illectus (Casey, 1911) Subtribe Meoticina Seevers, 1978 Genus MEOTICA Mulsant and Rey, 1873a 28. M. pallens (Redtenbacher, 1849){

Genus CRATARAEA C.G. Thomson, 1858 29. C. suturalis (Mannerheim, 1830){

Genus CALODERA MANNERHEIM, 1830 30. C. bennetti Klimaszewski, sp. n.

Genus GNATHUSA Fenyes, 1910 32. G. eva Fenyes, 1910 33. G. tenuicornis Fenyes, 1921

Genus OCYUSA Kraatz, 1856 35. O. canadensis Lohse, in Lohse et al. 1990 Genus OXYPODA Mannerheim, 1830 Convergens Species Group 36. O. canadensis Klimaszewski, in Klimaszewski et al. 2006 37. O. convergens Casey, 1893 38. O. manitobae Casey, 1911 39. O. opaca (Gravenhorst, 1802){ Gatosensis Species Group 40. O. gatosensis Bernhauer, 1905 41. O. longicarinata Klimaszewski, in Klimaszewski et al. 2006 Irrasa Species Group 42. O. irrasa Mäklin, 1853 Lacustris Species Group 43. O. famula Casey, 1911 44. O. lacustris Casey, 1906 45. O. nimbata Casey, 1911

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

Orbicollis Species Group 46. O. frigida Bernhauer, 1907 47. O. orbicollis Casey, 1911 Smithi Species Group 48. O. smithi Klimaszewski, in Klimaszewski et al. 2006 Stanleyi Species Group 49. O. stanleyi Klimaszewski and McLean, in McLean et al. 2009 Sylvia Species Group 50. O. sylvia Casey, 1906 Vancouveri Species Group 51. O. vancouveri Klimaszewski, in Klimaszewski and Winchester 2002 Species incertae sedis 52. O. glenorae Casey, 1893 53. O. impressa Casey, 1893 54. O. regressa Casey, 1911 OCALEA Group of Genera Klimaszewski and Pelletier, 2004 Genus ALFOCALEA Klimaszewski, in Klimaszewski and Pelletier 2004 55. A. montana Klimaszewski, in Klimaszewski and Pelletier 2004 Genus BETOCALEA Klimaszewski, in Klimaszewski and Pelletier (2004) 56. B. pacifica Klimaszewski, in Klimaszewski and Pelletier 2004 Genus MEGOCALEA Klimaszewski, in Klimaszewski and Pelletier (2004) 57. M. lemieuxi Klimaszewski, in Klimaszewski and Pelletier 2004 Genus METOCALEA Klimaszewski, in Klimaszewski and Pelletier (2004) 58. M. lindgreni Klimaszewski, in Klimaszewski and Pelletier 2004

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Genus NEOISOGLOSSA Casey, 1893 59. N. agnita (Casey, 1911) Genus NEOTHETALIA Klimaszewski, in Klimaszewski and Pelletier 2004 60. N. canadiana Klimaszewski, in Klimaszewski and Pelletier 2004 61. N. columbiana (Klimaszewski and Winchester 2002) 62. N. nimia (Casey, 1910) 63. N. pallitarsis (Kirby, 1837) 64. N. robergei Klimaszewski, sp.n. 65. N. smetanai Klimaszewski, in Klimaszewski and Pelletier 2004 Genus OCALEA Erichson, 1837 66. O. vancouveri Casey, 1910 Genus PAROCALEA Bernhauer, 1901a 67. P. pseudobaicalica Lohse, 1990 Subtribe Phloeoporina C.G. Thomson, 1859 Genus PHLOEOPORA C.G. Thomson, 1859 68. P. arctica Lohse, in Lohse et al. 1990 69. P. canadensis Klimaszewski and Langor, in Klimaszewski et al. 2011 70. P. oregona Casey, 1906 Tribe Tachyusini C.G. Thomson, 1859 Genus BRACHYUSA Mulsant and Rey, 1873c 71. B. americana (Fenyes, 1921) 72. B. helenae (Casey, 1911) Genus GNYPETA C.G. Thomson, 1858 Caerulea Species Group 73. G. caerulea (C.R. Sahlberg, 1831)
74. G. lohsei Klimaszewski, in Klimaszewski et al. 2008a Crebrepunctata Species Group 75. G. crebrepunctata Casey, 1886

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

Helenae Species Group 76. G. helenae Casey, 1906 Nigrella Species Group 77. G. baranowskii Klimaszewski, sp. n. Sellmani Species Group 78. G. brevicornis Casey, 1906 Genus PARADILACRA Bernhauer, 1909 79. P. densissima (Bernhauer, 1909) Genus TACHYUSA Erichson, 1837 (Subgenus TACHYUSA Erichson, 1837) 80. T. americanoides Paśnik, 2006 81. T. cavicollis LeConte, 1863 82. T. obsoleta Casey, 1906 Tribe Hypocyphtini Laporte, 1835 Genus CYPHA Leach, 1819 83. C. crotchii (Horn 1877) Genus HOLOBUS Solier, 1849 84. H. vancouveri Klimaszewski, in Klimaszewski and Winchester (2002) Genus OLIGOTA Mannerheim, 1830 85. O. parva Kraatz, 1862{ Tribe Myllaenini Ganglbauer, 1895 Genus MYLLAENA Erichson, 1837 Insomnis Species Group 86. M. audax Casey, 1911 87. M. decreta Casey, 1911 88. M. insomnis Casey, 1911 Potawatomi Species Group 89. M. fenyesi Bernhauer, 1907 Tribe Diglottini Jakobson, 1909 Genus BRYOTHINUSA Casey, 1904 90. B. catalinae Casey, 1904

91. A. brevipes Casey, 1893 Genus PARAMBLOPUSA Ahn and Ashe, 1996 92. P. borealis (Casey, 1906) Genus DIAULOTA Casey, 1893 93. D. densissima Casey, 1893 Genus LIPAROCEPHALUS Mäklin, 1853 94. L. brevipennis Mäklin, 1853 95. L. cordicollis LeConte, 1880 Tribe Autaliini C.G. Thomson, 1859 Genus AUTALIA Leach, 1819 96. A. puncticollis Sharp, 1864{ 97. A. rivularis (Gravenhorst, 1802){ 98. A. truncatula Casey, 1911 Tribe Homalotini Heer, 1839 Subtribe Bolitocharina C.G. Thomson, 1859 Genus LEPTUSA Kraatz, 1856 (Subgenus Gatineauleptusa Klimaszewski, in Klimaszewski et al. 2018a) 99. L. gatineauensis Klimaszewski and Pelletier, in Klimaszewski et al. 2004 Genus LEPTUSOMORPHA Klimaszewski and Hoebeke, gen. n 100. L. claudiae Klimaszewski, sp. n. Genus STICTALIA Casey, 1906 101. S. brevicornis Casey, 1906 S. arcuata Casey, 1906. New synonym, lectotype designation. 102. S. californica (Casey, 1885) S. densicollis Casey, 1906. New synonym, lectotype designation. 103. S. carlottae Casey, 1911. Lectotype designation. 104. S. kranabetteri Klimaszewski and Godin, sp. n. Subtribe Gyrophaenina Kraatz, 1856

Tribe Liparocephalini Fenyes, 1918 Genus AMBLOPUSA Casey, 1893

Genus ENCEPHALUS Stephens, 1832

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

105. E. americanus Seevers, 1951

67

Genus ADOTA Casey, 1910 120. A. maritima (Mannerheim, 1843)

Genus GYROPHAENA Mannerheim, 1830 Genus ALOCONOTA C.G. Thomson, 1858 121. A. cambrica (Wollaston, 1855){ 122. A. carmanah Klimaszewski, in Klimaszewski and Winchester 2002

Affinis Species Group 106. G. affinis Mannerheim, 1830{ Bihamata Species Group 107. G. uteana Casey, 1906

Genus AMISCHA C.G. THOMSON, 1858 123. Amischa analis (Gravenhorst, 1802){

Keeni Species Group 108. G. keeni Casey, 1911

Genus ATHETA C.G. Thomson, 1858

Nana Species Group 109. G. nana (Paykull, 1800)


(Subgenus ALAOBIA C.G. Thomson, 1858) 124. A. ventricosa Bernhauer, 1907

Strictula Species Group 110. G. californica Casey, 1906

(Subgenus ATHETA C.G. Thomson, 1858) 125. A. graminicola (Gravenhorst, 1806)
126. A. ringi Klimaszewski, in Klimaszewski and Winchester (2002)

Subtribe Silusina Fenyes, 1918

(Subgenus BESSOBIA C.G. Thomson, 1858) 127. A. cryptica (Lohse, in Lohse et al. 1990)

Genus SILUSA Erichson, 1837 Californica Species Group 111. S. californica (Bernhauer, 1905) 112. S. opaca Fenyes, 1909

(Subgenus CHAETIDA Mulsant and Rey, 1873c) 128. A. longicornis (Gravenhorst, 1802){ (Subgenus DATOMICRA Mulsant and Rey, 1873c) 129. A. celata (Erichson, 1837)
Datomicra wrangleri Casey, 1910. New synonym, lectotype designation. Pseudota nanulina Casey, 1911. New synonym, lectotype designation. 130. A. dadopora C.G. Thomson, 1867


Vesperis Species Group 113. S. vesperis Casey, 1893 Tribe Placusini Mulsant and Rey, 1871 Genus PLACUSA Erichson, 1837 Canadensis Species Group 114. P. petulans Casey, 1911 115. P. tacomae Casey, 1893 116. P. vaga Casey, 1911 Tachyporoides Species Group 117. P. incompleta Sjöberg, 1934{ 118. P. pseudosuecica Klimaszewski, Klimaszewski et al. 2001 119. P. tachyporoides (Waltl, 1838){ Tribe Athetini Casey, 1910 Subtribe Athetina Casey, 1910

(Subgenus DIMETROTA Mulsant and Rey, 1873a)

in

Alesi Species Group 131. A. alesi Klimaszewski and Brunke, in Brunke et al. 2012 Campbelli Species Group 132. A. remulsa Casey, 1910 Columbica Species Group

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

133. A. stercoris Fenyes, 1920 A. columbica Bernhauer 134. A. winchesteri Klimaszewski, sp. n. Crassicornis (¼Modesta) Species Group 135. A. godini Klimaszewski, sp. n. 136. A. metlakatlana Bernhauer, 1909 137. A. pseudometlakatlana Klimaszewski and Godin, in Klimaszewski et al. (2008b) 138. A. wheelerae Klimaszewski, sp. n. Districta Species Group 139. A. districta Casey, 1911 Fanatica Species Group 140. A. fanatica Casey, 1910 A. irrita Casey 141. A. munsteri Bernhauer, 1902
. Lectotype designation Hampshirensis Species Group 142. A. hampshirensis Bernhauer, 1909 Picipennis Species Group 143. A. picipennis (Mannerheim, 1843) A. cursor (Mäklin) A. carlottae (Casey) Prudhoensis Species Group 144. A. brumalis Casey, 1910 145. A. capsularis Klimaszewski, in Klimaszewski et al. (2005) 146. A. prudhoensis (Lohse, in Lohse et al. 1990) 147. A. rurigena Casey, 1911 148. A. strigosula Casey, 1910 149. A. terranovae Klimaszewski and Langor, in Klimaszewski et al. (2011) (Subgenus HYDROSMECTOMORPHA Klimaszewski and Webster, in Klimaszewski et al. 2018b) 150. A. meduxnekeagensis Webster and Klimaszewski, in Klimaszewski et al. 2018b (Subgenus LAMIOTA Casey, 1910) 151. A. keeni Casey, 1910

152. A. relicta (Casey, 1911). New subgeneric combination, lectotype designation. (Subgenus METADIMETROTA Klimaszewski and Winchester, 2002) 153. A. cheersae Klimaszewski, in Klimaszewski and Winchester 2002 (Subgenus MICRODOTA Mulsant and Rey, 1873a) 154. A. copleyi Klimaszewski, sp. n. 155. A. holmbergi Bernhauer, 1907. Lectotype designation 156. A. lucifera Bernhauer, 1906 157. A. platonoffi Brundin, 1948
158. A. ripariides Newton (2017: 11) 159. A. surgens (Casey, 1910). New subgeneric combination, lectotype designation. (Subgenus PANCOTA Casey, 1910) 160. A. nescia Casey, 1910 (Subgenus PSEUDOTA Casey, 1910) 161. A. concessa Casey, 1911 162. A. pseudoklagesi Klimaszewski and Webster, in Webster et al. 2016 (Subgenus TETROPLA Mulsant and Rey, 1873c) 163. A. frosti Bernhauer, 1909 Atheta Incertae Sedis 164. A. brunswickensis Klimaszewski, 2005 165. A. formalis (Casey, 1911) 166. A. irrupta (Casey, 1910) 167. A. pseudoschistoglossa Klimaszewski and Webster, in Webster et al. (2016) ATHETA Species Excluded from Valid Species List in BC A. basicornis (Mulsant and Rey, 1852){ [not confirmed in BC, removed from BC list] A. subrugosa (Märkel and Kiessenwetter, 1848){ [not confirmed in BC, removed from BC list] Genus BOREOPHILIA Benick, 1973

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

168. B. nomensis (Casey, 1910) Genus BOREOSTIBA Lohse, in Lohse et al. (1990) Parvipennis Species Group 169. B. pseudolaticollis Klimaszewski and Godin, sp. n. 170. B. parvipennis (Bernhauer, 1907) Genus CLUSIOTA Casey, 1910 171. C. impressicollis (Bernhauer, 1907)

69

Terminalis Species Group 183. L. quadricollis (Casey, 1893) 184. L. scudderi Klimaszewski, sp. n. 185. L. terminalis (Casey, 1906) 186. L. trapezicollis Lohse, in Lohse et al. (1990) 187. L. wickhami (Casey, 1893) Genus LYPOGLOSSA Fenyes, 1918 188. L. angularis (Mäklin, 1853) 189. L. franclemonti Hoebeke, 1992 Genus MOCYTA Mulsant and Rey, 1873c

Genus DALOTIA Casey, 1910 172. D. coriaria (Kraatz, 1856){ Genus DINARAEA C.G. Thomson, 1858 173. D. angustula (Gyllenhal, 1810){ 174. D. inexpectata Klimaszewski, sp. n. 175. D. pacei Klimaszewski and Langor, in Klimaszewski et al. 2011 176. D. subdepressa (Bernhauer, 1907) Genus DOCHMONOTA C.G. Thomson, 1859 177. D. rudiventris (Eppelsheim, 1886){ Genus EAROTA Mulsant and Rey, 1873c 178. E. dentata (Bernhauer, 1906) Genus GEOSTIBA C.G. Thomson, 1858 (Subgenus Geostiba C.G. Thomson, 1858) 179. G. horwoodae Klimaszewski and Godin, sp. n. Genus GONIUSA Casey, 1906 180. G. caseyi Gusarov, 2003a Genus LIOGLUTA C.G. Thomson, 1858 Microgranulosa Species Group 181. L. atriventris (Casey, 1906) Nigropolita Species Group 182. L. nitens (Mäklin, 1852)

Amblystegii Species Group 190. M. breviuscula (Mäklin, 1852) 191. M. fungi (Gravenhorst, 1806){ 192. M. discreta (Casey) Genus NEHEMITROPIA Lohse, 1971 193. N. lividipennis (Mannerheim, 1830){ Genus OUSIPALIA Gozis, 1886 194. O. pacifica Casey, 1910. Lectotype designation Genus PARALEPTONIA Klimaszewski, in Klimaszewski and Winchester (2002) 195. P. pacei Klimaszewski, in Klimaszewski and Winchester (2002) Genus PELIOPTERA Kraatz, 1857 (Subgenus Tropimenelytron Pace, 1983) 196. P. robusta (Gusarov, 2002) Genus PHILHYGRA Mulsant and Rey, 1873a 197. P. angusticauda (Bernhauer, 1909) 198. P. botanicarum (Muona, 1983)
199. P. charlottae Klimaszewski, sp. n. 200. P. clemens (Casey, 1910) 201. P. laevicollis (Mäklin, 1852) 202. P. terrestris Klimaszewski and Godin, in Klimaszewski et al. 2012 203. P. terrivaga Klimaszewski, sp. n.

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

Genus PONTOMALOTA Casey, 1885 204. P. opaca (LeConte, 1863) Genus PSAMMOSTIBA Yosii and Sawada (1976) 205. P. comparabilis (Mäklin, 1853) 206. P. kenaii Gusarov Genus SCHISTOGLOSSA Kraatz, 1856 (Subgenus Boreomorpha Klimaszewski and Webster, in Klimaszewski et al. 2009) 207. S. carexiana Klimaszewski, in Klimaszewski et al. 2009 (Subgenus Schistoglossa Kraatz, 1856) 208. S. charlottae Klimaszewski, in Klimaszewski et al. 2009 209. S. campbelli Klimaszewski, in Klimaszewski et al. 2009 210. S. hampshirensis Klimaszewski, in Klimaszewski et al. 2009 Genus SEEVERSIELLA Ashe, 1986 211. S. globicollis (Bernhauer, 1907) Genus STRIGOTA Casey, 1910 212. S. ambigua (Erichson, 1839) 213. S. obscurata Klimaszewski and Brunke, in Brunke et al. 2012 Genus TARPHIOTA Casey, 1893 214. T. fucicola (Mäklin, 1852) 215. T. geniculata (Mäklin, 1852) Genus THINUSA Casey, 1893 216. T. fletcheri Casey, 1906 217. T. maritima (Casey, 1885) Genus TRICHIUSA Casey, 1893

218. T. columbica Casey, 1911. Lectotype designation. 219. T. pilosa Casey, 1893 Athetini with Subtribe Affiliation Uncertain Genus PARAGONIUSA Maruyama and Klimaszewski, 2004 220. P. myrmicae Maruyama and Klimaszewski, 2004 Tribe Falagriini Mulsant and Rey, 1873c Genus BRYOBIOTA Casey, 1893. Revised concept. 221. B. bicolor (Casey, 1885) Genus CORDALIA Jacobs, 1925 222. C. obscura (Gravenhorst, 1802){ Genus FALAGRIA Leach, 1819 223. F. dissecta Erichson, 1839 Genus MYRMECOCEPHALUS MacLeay, 1871 224. Myrmecocephalus arizonicus (Casey, 1906) Tribe Lomechusini Fleming, 1821 Subtribe Lomechusina Fleming, 1821 Genus XENODUSA Wasmann, 1894 225. X. reflexa (Walker, 1866) Subtribe Myrmedonina C.G. Thomson, 1867 Genus MYRMOECIA Mulsant and Rey, 1873c 226. M. lugubris (Casey, 1893) Genus ZYRAS Stephens, 1835 227. Z. obliquus (Casey, 1893)

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

References Ahn K-J, Ashe S (1996) Phylogeny of the intertidal aleocharine tribe Liparocephalini (Coleoptera: Staphylinidae). Syst Entomol 21(2):99–114 Ashe JS (1986) Seeversiella bispinosa, a new genus and species of athetine Aleocharinae (Coleoptera: Staphylinidae) from North America. J N Y Entomol Soc 94(4):500–511 Benick G (1973) Die von Prof. Dr. Kaszab in der Mongolei gesammelten Arten der Subfamilie Athetae (Col. Staphyl.). Nouvelle Rev d’Entomol 3 (4):211–217 Bernhauer M (1901a) Die Staphyliniden der paläarktischen Fauna. Verh Kaiserli-Königli ZoolBota Gesell Wien 51(6): 430–466 51(7):467–506 Bernhauer M (1901b) Neue exotische Arten der Gattung Aleochara Gravenhorst. Steitti Entomol Ztg 62 (3–4):366–373 Bernhauer M (1902) Die Staphyliniden der paläarktischen Fauna. 1. Tribus: Aleocharini (part II). Verh KaiserliKönigli Zool-Bota Gesell Wien 52:87–284 Bernhauer M (1905) Neue Aleocharinen aus Nordamerika. Deutsche Entomologische Zeitschrift 1905 (2):249–256 Bernhauer M (1906) Neue Aleocharinen aus Nordamerika. (II. Teil). Deutsche Entomologische Zeitschrift 1906 (2):337–348 Bernhauer M (1907) Neue Aleocharini aus Nordamerika. (Col.) (3. Stück.). Deutsche Entomologische Zeitschrift 1907(4):381–405 Bernhauer M (1909) Neue Aleocharini aus Nordamerika. (Col.) (4. Stück.). Deutsche Entomologische Zeitschrift 1909(4):515–528 Blackwelder RE (1952) The generic names of the beetle family Staphylinidae , with an essay on genotypy. US Nat Museum Bull 200:(I–IV) + 1–483 Brundin L (1948) Microdota-Studien. (Col. Staphylinidae). Entomologisk Tidskrift 69(1–2):8–66 Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and Ecology of Canadian Staphylinidae (Coleoptera) II, Zookeys [Special Issue], vol 186, pp 119–206 Casey TL (1884) Contributions to the descriptive and systematic Coleopterology of North America. Collins Printing House, Philadelphia. Part I, pp 1–60, pl. 1; Part II, pp 61–124 Casey TL (1885) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1(4):283–336 Casey TL (1893) Coleopterological notices. V. Annals of the New York Academy of Sciences 7[1894]:281–606. [these fascicles (Oct–Dec 1893) of the Annals were published in Mar 1894 (Trans. New York Acad. Sci.

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13:152; Ent. News 5 (6):187); however, it was listed by Bertkau in his annual “Bericht über die wissenschaftlichen Leistungen im Gebiete der Entomologie während des Jahres 1893”, which was published in Archiv für Naturgeschichte, 60. Jahrg., II. Band, Zweites Heft, which was issued in December 1893 (see Inhalt p. V and p. 198); it was also recorded on 24 Jan 1894, as a separate of 325 pp, 1893 (Ann. Soc. Ent. France 63: Bull. Ent.: xxxv), and in Jan 1894 (Naturae Novitates 16 1894 (2):72)] Casey TL (1904) On some new Coleoptera including four new genera. Can Entomol 36:312–324 Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St Louis 16(6):125–434 Casey TL (1910) New species of the staphylinid tribe Myrmedoniini [pp 1–183]. In: Memoirs on the Coleoptera. I. New Era Printing Co., Lancaster, Pennsylvania, 205 pp Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, Pennsylvania, 259 pp Casey TL (1915) Studies in some staphylinid genera of North America [pp 395–450]. In: Memoirs on the Coleoptera. VI. The New Era Printing Co., Lancaster, Pennsylvania, 460 pp Eppelsheim E (1886) Neue Staphylinen vom Amur. Deutsche Entomologische Zeitschrift 30(1):33–46 Erichson W F (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Erichson WF (1839) Genera et species Staphylinorum insectorum coleopterorum familiae. Erster Band. F. H. Morin, Berlin, viii + 400 pp Fenyes A (1910) Two new species of Aleocharinae from California. Proc Entomol Soc Wash 11:197–199 Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera Insectorum, vol XXVII, Fascicule 173A. M. Nijhoff, Den Haag. 453 pp + 7 pls Fenyes A (1920) Aleocharinae. Genera Insectorum, fasc. 173B, pp 111–414 Fenyes A (1921) New genera and species of Aleocharinae with a polytomic synopsis of the tribes. Bull Mus Comp Zool 65(2):17–36 Fleming J (1821) Insecta [pp 41–56]. In: Supplement to the fourth, fifth and sixth editions of the Encyclopaedia Britannica, with preliminary dissertations on the history of sciences, vol 5. A. Constable and Company, Edinburgh, 584 pp Ganglbauer L (1895) Die Käfer von Mitteleuropa. Die Käfer der österreichisch-ungarischen Monarchie, Deutschlands, der Schweiz, sowie des französischen und italienischen Alpengebietes. Zweiter Band. Familienreihe Staphylinoidea, I. Theil: Staphylinidae, Pselaphidae. Carl Gerold’s Sohn, Vienna. vi + 881 pp

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Goeze JAE (1777) Entomologische Beyträge zu des Ritter Linné zwölften Ausgabe des Natursystems. Erster Theil. Weidmanns, Erben & Reich, Leipzig, xvi + 736 pp Gozis M (1886) Récherches de l’espéce typique de quelque anciens genres: Rectifications synonymiques et notes diverse. Herbin, Montluçon. 36 pp Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Gravenhorst JLC (1806) Monographia Coleopterorum Micropterorum. Henricus Dieterich, Göttingen: xvi + 236 + [12] pp Gusarov VI (2002) A revision of Nearctic species of the genus Tropimenelytron Pace, 1983 (Coleoptera: Staphylinidae: Aleocharinae), a new genus for North America. Zootaxa 114:1–24 Gusarov VI (2003a) A revision of the genus Goniusa Casey, 1906 (Coleptera: Staphylinidae, Aleocharinae). Zootaxa 164:1–20 Gusarov VI (2003b) A revision of the genus Seeversiella Ashe, 1986 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 142:1–102 Gyllenhal L (1810) Insecta Svecica. Classis I: Coleoptera sive Eleuterata. Tomi I pars II. F.J. Leverentz, Scaris. xix + [1] + 660 pp Heer O (1839) (1838–1841) Fasciculus secundus [pp 145–360] In: Fauna Coleopterorum Helvetica. Pars I. Orellii, Fuesslini et Sociorum: xii + 652 pp Hoebeke ER (1992) Taxonomy and distribution of the athetine genus Lypoglossa Fenyes (Coleoptera: Staphylinidae: Aleocharinae) in North America, with description of a new species. J N Y Entomol Soc 100 (2):381–398 Horn GH (1877) Synopsis of the genera and species of the Staphylinid ae [sic] tribe Tachyporini of the United States. Tras Amer Ent Soc 6:81–128 + p 11 Jacobs W (1925) Über den Gattungsnamen Cardiola Mulsant et Rey (Col.). Entomol Zeitschrift (Frankfurt am Main) 38:82 Kirby W (1837) The Insects. Coleoptera. In: Richardson, Fauna Boreali-Americana; or the zoology of the northern parts of British America, [etc.] Part the fourth and Last. J. Fletcher, Norwich Klimaszewski J (1979) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae: Aleocharinae). Research Branch Agriculture Canada Monograph No. 25:1–169 Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Memoirs of the Entomological Society of Canada No. 129, [2] + 211 pp Klimaszewski J, Génier F (1985) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae, Aleocharinae). Supplement 3. Coleopt Bull 39(1):60–66

Klimaszewski J, Pelletier G (2004) Review of the Ocalea group of genera (Coleoptera, Staphylinidae, Aleocharinae) in Canada and Alaska: new taxa, bionomics, and distribution. Can Entomol 136 (4):443–500 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc belge d’Entomol 40:3–126 Klimaszewski J, Pelletier G, Germain C, Hébert C, Humble LM, Winchester NN (2001) Diversity of Placusa (Coleoptera: Staphylinidae, Aleocharinae) in Canada, with descriptions of two new species. Can Entomol 133(1):1–47 Klimaszewski J, Pelletier G, Majka C (2004) A revision of Canadian Leptusa Kraatz (Col., Staphylinidae, Aleocharinae): new species, new distribution records, key and taxonomic considerations. Belg J Entomol 6 (1):3–42 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48 Klimaszewski J, Pelletier G, Germain C, Work T, Hébert C (2006) Review of Oxypoda species in Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics, and distribution. Can Entomol 138(6):737–852 Klimaszewski J, Savard K, Pelletier G, Webster R (2008a) Species review of the genus Gnypeta Thomson from Canada, Alaska and Greenland (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics and distribution. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:11–84 Klimaszewski J, Godin B, Pelletier G, Savard K (2008b) Six new species and records of aleocharine beetles from the Yukon and Alaska (Coleoptera: Staphylinidae: Aleocharinae). Can Entomol 140 (3):265–291 Klimaszewski J, Webster RP, Savard K (2009) First record of the genus Schistoglossa Kraatz from Canada with descriptions of seven new species (Coleoptera, Staphylinidae, Aleocharinae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:45–79 Klimaszewski J, Langor D, Pelletier G, Bourdon C Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Bourdon C (2012) Further contributions to the aleocharine fauna of the Yukon Territory, Canada (Coleoptera, Staphylinidae). In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II, ZooKeys [Special Issue], vol 186, pp 207–237

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List of Recorded Species of British Columbian Aleocharinae and Their Composition

Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018a) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, Switzerland, xvi + 902 pp Klimaszewski J, Webster RP, Davies A, Bourdon C (2018b) Description of Hydrosmectomorpha Klimaszewski and Webster, a new subgenus of Atheta C.G. Thomson, with three new Canadian species (Coleoptera: Staphylinidae: Aleocharinae). Insecta Mundi 0648:1–12 Kraatz G (1856) Staphylinii. Naturgeschichte der Insecten Deutschlands. Erste Abtheilung Coleoptera. Zweiter Band. Erste, zweite Lieferung. Bogen 1–24.Nicolai, Berlin, viii + 376 pp Kraatz G (1857). Genera Aleocharinorum illustrata. Linnaea Entomol 11:1–43, 2 pls Kraatz G (1862) Synonymische Bemerkungen. Ueber Coleopteren. Berliner Entomol Zeits 6(1/2):298–300 Laporte FL (1835) Études entomologiques, ou description d'insectes nouveaux: et observations sur leur synonymie. Premiére Partie, Méquignon-Marvis Père et Fils, Paris: 159 pp + 4 pls Latreille PA (1802) Histoire naturelle, générale et particulière, des crustacés et des insectes. Familles naturelles des genres. Tome neuvième. F. Dufart, Paris, 416 pp + pls. 74–80 Leach W E (1819) New genera. In: Samouelle (ed) The entomologist’s useful compendium; or an introduction to the knowledge of British insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; etc. Thomas Boys, London, 496 pp + 12 pls LeConte JL (1863) New species of North American Coleoptera. Prepared for the Smithsonian Institution. Part I. Smiths Misc Collect 6(167):1–86 LeConte JL (1867) Additions to the coleopterous fauna of the United States. No. 1. Proc Acad Natl Sci Phila 18 (5):361–394 LeConte JL (1880) Short studies of North American Coleoptera. Trans Am Entomol Soc 12:163–218 Lohse GA (1971) Über gattungsfremde Arten und Artemkreise innerhalb der “Grossgattung” Atheta Thomson. Verh Ver Naturwiss Heimatfors Hamburg 38:67–83 Lohse GA, Klimaszewski J, Smetana A (1990) Revision of arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44(2):121–202 MacLeay W (1871) Notes on a collection of insects from Gayndah. Trans Entomol Soc New South Wales 2 [1869–1873] (2):79–158; (3): 159–205 Mäklin FW (1852) [New species and notes]. In: Mannerheim, C.G.: Zweiter Nachtrag zur KaeferFauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soci Imp Naturali Moscou 25(2):283–372 Mäklin FW (1853) [New species and notes]. In: Mannerheim CG: Dritter Nachtrag zur Kaefer-

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Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Imp Naturali Moscou 26(3):95–273 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp [separate, re-issued in Mém Prés Acad Impé Sci St.-Pétersbourg 1:415–501, in 1831] Mannerheim CG (1843) Beitrag zur Kaefer-fauna der Aleutischen Inseln, der Insel Sitkha und Neu-Californiens. Bull Soci Imp Naturali Moscou 16 (2):175–314 Märkel F, Kiessenwetter EAH (1848) Bericht über eine entomologische excursion in die Käarthner Alpen im Jahre 1847. Entomol Ztg (Stettin) 9(11):321–329 Maruyama M, Klimaszewski J (2004) A new genus and species of the myrmecophilous Athetini, Paragoniusa myrmicae (Coleoptera: Staphylinidae: Aleocharinae) from Canada. Entomol Rev Japan 59(2):241–248 McLean JA, Klimaszewski J, Li A, Savard K (2009) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species. Part 1. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:5–17 Mulsant E, Rey C (1852) Description de quelques coléoptères nouveaux ou peu connus de la tribu des Brachelytres. Opuscules Entomologiques 1:15–46 Mulsant E, Rey C (1871) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens [Huitième Branche, Bolitocharaires]. Deyrolle Fils, Paris. [4] + 321 pp + [5, explic.] + [4, table, erratum], pls. 1–5 Mulsant E, Rey C (1873a) Description de divers coléoptères brévipennes nouveaux ou peu connus. Opusc Entomol 15:147–189 Mulsant E, Rey C (1873b) Histoire naturelle des Coléoptères de France: Brévipennes Aléochariens [Première branche dinardaires – cinquième branche oligotaires]. Deyrolle Fils, Paris. [4] + 155 pp, pls. 1–2 Mulsant E, Rey C (1873c) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens (Suite) [Tableau... des Aléochariens + Septième Branche, Myrmédoniaires, part 1]. Deyrolle, Paris, [2] + 695 pp, pls. 1–5 Mulsant E, Rey C (1874) [Histoire naturelle des Coléoptères de France]. Tribu des brévipennes: Famille des aléochariens (suite): Sixième branche: Aléocharaires. Anna Soci Linn Lyon (NS) 20 [1873]:285–447 Mulsant E, Rey C (1875) Histoire naturelle des Coléoptères de France: Brévipennes Aléochariens (suite). Myrmédoniaires (2eme partie). Deyrolle, Paris: [2] + 470 pp + 4 pls Muona J (1983) Two new Palearctic Atheta species (Coleoptera, Staphylinidae). Ann Entomol Fenn 49 (2):57–58

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Pace R (1983) Il genere Tropimenelytron Scheerpeltz (Coleoptera, Staphylinidae) (XXXIX contributo alla conoscenza delle Aleocharinae). Nouvelle Rev d’Entomol 13(2):185–190 Paśnik G (2006) A revision of the World species of the genus Tachyusa Erichson, 1837 (Coleoptera, Staphylinidae: Aleocharinae). Zootaxa 1146:1–152 Paykull G (1800) Fauna Suecica. Insecta. Tomus III. J.F. Edman, Upsala: 459 pp Redtenbacher L (1849) Fauna Austriaca. Die Käfer. Nach der analytischen Methode bearbeitet. C. Gerold, Wien: pp I–xxvii + 641–883 Sahlberg CR (1831) [Pp 361–408]. In: Insecta Fennica, dissertationibus academicis, A. 1817–1834 editis. Pars I: A. J.C. Frenckel, Helsingfors, VIII + 519 pp Say T (1833) Descriptions of new North American insects and observations on some of the already described. T. Say, New Harmony, Indiana, pp. 58–65 Seevers CH (1951) A revision of the North American and European staphylinid beetles of the subtribe Gyrophaenae (Aleocharinae, Bolitocharini). Fieldiana Zool 32(10):657–762 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana Zool 71:(i–vi), 1–289 Sharp D (1864) New taxon. J Proc Entomol Soc Lond 1864:45 Sharp D (1883) Fam. Staphylinidae [part]. In: Godman FD, Salvin O (eds) Biologia Centrali-A312, pls. 5–7 Sjöberg O (1934) Zwei neue Bolitocharinen aus Schweden. Entomologisk Tidskrift 55:281–285 Solier AJJ (1849) Zoología. Tomo cuarto. Insectos. Orden III. Coleópteros. [pp 105–380, 414–511, pls. 1–12]. In: Gay C (ed) Historia física y política de Chile. [etc.]. C. Gay, Paris. 511 pp Stephens JF (1832) (1832–1835) Illustrations of British Entomology. [etc.]. Mandibulata, V. Baldwin & Cradock, London. pp 1–240, pls. 24–26 Stephens JF (1833) The nomenclature of British insects, together with their synonymies: being a compendious list of such species as are contained in the Systematic

Catalogue of British Insects. 2nd edn, Baldwin and Cradock, London. iv pp, 135 columns Thomson CG (1858) Försök till uppställning af Sveriges Staphyliner. Öfversigt af Kongliga VetenskapsAkademiens Förhandlingar 15(1):27–40 Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom. Berlingska Boktryckeriet, Lund, [3] + 290 pp Thomson CG (1867) Skandinaviens Coleoptera, synoptiskt bearbetade. Supplementum. Tom IX. Lund: Lundbergska Boktryckeriet, 407 pp Walker F (1866) List of Coleoptera [and descriptions]. [Appendix pp 309–334]. In: Lord JK: The naturalist in Vancouver Island and British Columbia, vol II. R. Bentley, London, vii + [2] + 375 pp Waltl J (1838) Verzeichniss der um Passau vorkommenden seltnern Käfer [cols. 263–267]; Um Passau vorkommende neue Arten von Käfern [cols. 267–273]. In: Beyträge zur nähern naturhistorischen Kenntniss des Unterdonaukreises in Bayern. Isis von Oken 1838(4):250–273 Wasmann E (1894) Kritisches Verzeichniss der myrmekophilen und termitophilen Arthropoden. Mit Angabe der Lebensweise und mit Beschreibung neuer Arten. F. L. Dames, Berlin. xiii + 231 pp Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016) Further contributions to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including descriptions of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216 Wollaston TV (1855) Descriptions of two Coleopterous insects new to the British fauna. Zoologist 13: (Appendix) Yosii R, Sawada K (1976) Studies on the genus Atheta Thomson and its allies (Coleoptera, Staphylinidae). II. Diagnostic characters of genera and subgenera with descriptions of representative species. Contr Biol Lab Kyoto Univ 25:11–140

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Key to Tribes of Aleocharine Occurring in British Columbia

Diagnosis. Aleocharinae can be recognized by the following combination of characters: anterior tentorial pits located at inner corner of antennal fossae which may be connected by frontal suture (e.g. Oxypoda); antennal insertions exposed, usually located between eyes and posterior to a line drawn between anterior edges of eyes (except in species with reduced eyes); elytra transverse to slightly elongate; posterior coxae expanded laterally under the femora; male tergite IX usually subdivided (except in Gymnusa), and extends lateroposteriad forming two ventral struts; unique tergal defensive gland with associated structures present in higher Aleocharinae at base of segment VII in adults and at apex of segment VIII in larvae; aedeagus trilobite, with large and complex parameres (lateral lobes). The form of the parameres is unique to Aleocharinae (consisting of paramerite and condylite, with paramerite subdivided into several parts as illustrated in Chap. 3, it is a shared derived feature that provides strong evidence for monophyly of this subfamily (Klimaszewski et al. 2018). Key to Tribes (Figs. 8.23–8.31) (Modified from Klimaszewski et al. 2018) 1. Body form distinctive (Figs. 8.24a–f and 8.25a), head at least as wide as pronotum at front and abdomen at base in most (exception Tachyusini, head about as wide as pronotum at

front), elytra wide and at least one-fourth wider than base of pronotum, punctation of forebody fine, first three visible abdominal tergites with deep transverse impression often with coarse punctation or ridges............................................ 2 – Body form and other characters not as above ............................................................................4 2. Head with narrow neck, less than one-third width of head (Figs. 8.24d–f and 8.25a); pronotum with distinct median sulcus or median elongate depression (Fig. 8.24d, e, f), and/or with basal foveae (Fig. 8.25a)....................................3 – Head with neck more than one-third as wide as head or without distinct neck (Fig. 8.24a–c); pronotum without median sulcus, median elongate depression, or basal foveae (Fig. 8.24a–c) ................................. Tachyusini C.G. Thomson 3. Pronotum widest subapically, narrowed basally to about three fourths of maximum width of pronotum (Fig. 8.24a–f) ........................ ...............................Falagriini Mulsant and Rey – Pronotum as wide basally as subapically or only slightly wider subapically (Fig. 8.25a) .....................................Autaliini C.G. Thomson 4. Posterior margin of first two first visible abdominal tergites each with ctenidium of short comb-like cuticular teeth (projections) consisting of one (Gymnusa, Fig. 8.23f), or two types of teeth (Deinopsis); tarsal formula 5-5-5

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_8

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(Gymnusa), or 3-3-3 (Deinopsis); body as illustrated (Fig. 8.25b).......................................... ................................. Gymnusini C.G. Thomson – Posterior margin of basal abdominal tergites without ctenidium; body not as above ................ 5 5. Body robust (Aleochara) (Fig. 8.25c, d) to moderately slender (Amarochara, not recorded in BC); maxillary and labial palpi with pseudosegment on last palpomere, appearing as five and four palpomeres, respectively (Aleochara, Fig. 8.23d, e), or pseudosegment not apparent (Amarochara); antennae moderately to distinctly robust (Fig. 8.25c, d) or swollen; pronotal hypomera not visible or only narrowly visible in lateral view; tarsal formula 5-5-5 or 4-5-5 (subgenus Tinotus) ..................................... ........................................... Aleocharini Fleming – Body of a different shape; other characters in different combination; tarsal formula in other combinations and rarely 5-5-5 (e.g. Oxypoda).... 6 6. Body wide and flattened (Fig. 8.25e); pronotum with anterolateral angles obtuse, pubescence short, dense and evenly distributed, directed straight posteriad on pronotum and elytra (Fig. 8.25e); tarsal formula 4-4-5......................... ................................ Placusini Mulsant and Rey – Body and other characters different; tarsal formula variable......................................................7 7. Body fusiform (Fig. 8.25f), labial palps very long and stylate (Fig. 8.23b), or body linear or robust with very short elytra (Fig. 8.26a–d); riparian or intertidal species ......................................8 – Body shape not as above and other characters different ............................................................10 8. Body fusiform (Fig. 8.25f); pronotum widely rounded apically in dorsal view; hypomera not visible in lateral view; elytra at suture at least as long as pronotum; abdomen strongly tapering and bearing numerous robust macrosetae; tarsal claws slender without spatulate setae; riparian habitats ......................... Myllaenini Ganglbauer – Body shape different (Fig. 8.26a–d); other characters not as above; intertidal zone of sea beaches ...............................................................9

Key to Tribes of Aleocharine Occurring in British Columbia

9. Body form distinctive, with large head, pronotum strongly narrowed posteriorly, elytra slightly shorter than pronotum, abdomen broadly oval (Fig. 8.26a); antennomere III subquadrate and much shorter than antennomere II (Fig. 8.26a); tarsal formula 4-4-5, tarsal claws scythe-like ...........................Diglottini Jakobson – Body form diverse, robust in most (except Amblopusa, Paramblopusa) (Fig. 8.26b–d); antennomere III elongate, almost as long as antennomere II or shorter but not subquadrate (Amblopusa, Paramblopusa); tarsal formula 4-4-5 or 4-4-4 (some Diaulota), tarsal claws robust with spatulate setae........ Liparocephalini Fenyes 10. Body minute and compact (Fig. 8.26e,f), length 0.8–1.6 mm; antennae with 10 antennomeres, antennomeres VIII–X enlarged and forming loose club; hind coxa with large ventral lamella covering base of femur; tarsal formula 4-4-4 ................. Hypocyphtini Laporte – Body elongate, small to large-sized (1.0–4.5 mm); antennae with 11 antennomeres and without club; hind coxae without ventral lamella covering base of femur; tarsal formula variable.............................................................11 11. Body narrowly elongate or ovoid, body compact or loosely articulated, moderately to distinctly flattened, usually compact, often well sclerotized and with coarse punctation (Fig. 8.27a–f); head with broad neck (not always well visible because head is contracted and covered by pronotum); mandibles with minute denticles in ventral molar region; prementum with bases of medial setae very close together, medial pseudopore field very narrow; ligula broadly rounded, completely split (divided from apex to base) or bifid (divided only apically); mesocoxae broadly or narrowly separated (Fig. 8.23c); tarsal formula 4-4-5; most species associated with fungi............... Homalotini Heer – Body shape different; other characters different or in different combinations ....................... 12 12. Body shape diverse (Figs. 8.28a–f and 8.29a–f); head with frontal suture present in most species (Fig. 8.23a); mesocoxae narrowly

8

Key to Tribes of Aleocharine Occurring in British Columbia

separated in most species; abdominal tergite IX narrowly subdivided at base; median lobe of aedeagus without athetine bridge (transverse sclerotized strip extending from subapical lateral edges of median lobe in front of compressor plate, Fig. 8.23g); tarsal formula predominantly 5-5-5, rarely 4-5-5 or 4-4-4 (in Meotica)........................ .................................. Oxypodini C.G. Thomson – Body shape not as above; median lobe of aedeagus with athetine bridge; other characters different or in different combinations; tarsal formula 4-5-5........................................................13 13. Body rather robust (Fig. 8.30a–c); maxillae with galea and lacinia moderately to highly elongate; mesocoxae widely separated by mesoventral processes; which is broad and short; compressor

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plate of median lobe of aedeagus covered by plate which extends from base of bulbus; most taxa associated with ants......... Lomechusini Fleming – Body elongate, loosely articulated and very flexible, only some species robust (Fig. 8.30d–f and 8.31a–f); maxillae with galea and lacinia moderately elongate; mesocoxae narrowly separated by mesoventral processes, which is narrow and moderate in length; compressor plate of median lobe of aedeagus exposed and not covered by a plate which extends from base of bulbus; “athetine bridge” of median lobe of aedeagus present (Fig. 8.23g); some species associated with fungi but most found in decaying organic matter ..........................................Athetini Casey

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8

Fig. 8.23 (a–g) Some morphological characters useful in identification to tribal level: (a) frontal suture of head (Oxypoda); (b) long and stylate labial palps (Myllaena); (c) mesocoxae narrowly separated and mesoventral process, isthmus and metaventral process; (d, e) maxillary palpus with terminal pseudosegment (Aleochara); (f)

Key to Tribes of Aleocharine Occurring in British Columbia

ctenidium of short comb-like cuticular teeth (projections) of first two visible abdominal tergites (Gymnusa); (g) “athetine bridge” of median lobe of aedeagus (Athetini). (a, b, c) Modified from Ashe 2001; (d, e) after Klimaszewski 2004

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Key to Tribes of Aleocharine Occurring in British Columbia

Fig. 8.24 (a–f) Habitus in dorsal view: (a) Brachyusa helenae (Casey); (b) Gnypeta caerulea (C.R. Sahlberg); (c) Tachyusa americanoides Paśnik; (d) Cordalia obscura

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(Gravenhorst); (e) Falagria dissecta (Erichson); (f) Myrmecocephalus arizonicus (Casey). Scale bar ¼ 1 mm

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Fig. 8.25 (a–f) Habitus in dorsal view: (a) Autalia rivularis (Gravenhorst); (b) Gymnusa grandiceps Casey; (c) Aleochara gracilicornis Bernhauer; (d) Aleochara

Key to Tribes of Aleocharine Occurring in British Columbia

lanuginosa Gravenhorst; (e) Placusa tacomae Casey; (f) Myllaena insomnis Casey. Scale bar ¼ 1 mm

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Key to Tribes of Aleocharine Occurring in British Columbia

Fig. 8.26 (a–f) Habitus in dorsal view: (a) Bryothinusa catalinae Casey; (b) Diaulota densissima Casey; (c) Liparocephalus cordicollis LeConte; (d) Amblopusa

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brevipes Casey; (e) Cypha crotchii (Horn); (f) Holobus vancouveri Klimaszewski. Scale bar ¼ 1 mm

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Fig. 8.27 (a–f) Habitus in dorsal view: (a) Encephalus americanus Seevers; (b) Gyrophaena californica Casey; (c) Leptusa gatineauensis Klimaszewski and Pelletier (apical part of abdomen removed); (d) Leptusomorpha

Key to Tribes of Aleocharine Occurring in British Columbia

claudiae Klimaszewski; (e) Silusa versperis Casey; (f) Stictalia kranabetteri Klimaszewski and Godin (apical part of abdomen removed). Scale bar ¼ 1 mm

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Key to Tribes of Aleocharine Occurring in British Columbia

Fig. 8.28 (a–f) Habitus in dorsal view: (a) Alfocalea montana Klimaszewski; (b) Calodera bennetti Klimaszewski; (c) Cratarea suturalis (Mannerheim); (d)

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Devia prospera (Erichson) (apical part of abdomen removed); (e) Gnathusa eva Fenyes; (f) Megocalea lemieuxi Klimaszewski. Scale bar ¼ 1 mm

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Fig. 8.29 (a–f) Habitus in dorsal view: (a) Meotica pallens (Redtenbacher); (b) Mniusa yukonensis (Klimaszewski and Godin); (c) Neothetalia nimia

Key to Tribes of Aleocharine Occurring in British Columbia

(Casey); (d) Ocyusa canadensis Lohse; (e) Oxypoda convergens Casey; (f) Oxypoda sylvia Casey. (a–d) apical part of abdomen removed. Scale bar ¼ 1 mm

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Key to Tribes of Aleocharine Occurring in British Columbia

Fig. 8.30 (a–f) Habitus in dorsal view: (a) Myrmoecia lugubris (Casey); (b) Zyras obliquus (Casey); (c) Xenodusa reflexa (Walker); (d) Aloconota cambrica

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(Wollaston); (e) Amischa analis (Gravenhorst); (f) Atheta ventricosa Bernhauer. (a, e) apical part of abdomen removed. Scale bar ¼ 1 mm

86

8

Fig. 8.31 (a–f) Habitus in dorsal view: (a) Atheta graminicola (Gravenhorst); (b) Atheta longicornis (Gravenhorst); (c) Boreophilia nomensis (Casey); (d)

Key to Tribes of Aleocharine Occurring in British Columbia

Boreostiba parvipennis (Bernhauer); (e) Mocyta discreta (Casey); (f) Pontomalota opaca (LeConte). Scale bar ¼ 1 mm

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Key to Tribes of Aleocharine Occurring in British Columbia

Reference Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF,

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Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. XVI + 902 pp

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Tribe Gymnusini Heer, 1839

Tribe Gymnusini Heer, 1839 (Figs. 9.32–9.34) (Klimaszewski, 1979; Ashe 2000, 2002, 2005; Ashe and Chatzimanolis 2003a, b; Wolf-Schwenninger 2004; Cai and Huang 2015; Yamamoto and Maruyama 2018 (revised definition of the tribe, Deinopsini merged with Gymnusini); Klimaszewski et al. 2018) Diagnosis. Body small to medium-sized (3.0–7.0 mm), sublimuloid to tear-drop shaped; densely pubescent throughout; uniformly blackish-brown to dark reddish-brown; head slightly hypognathous, moderately deflexed; vertex with (Gymnusa) or without (Deinopsis) six setigerous pores; antennae filiform, with 11 antennomeres, antennomere XI without papilla; mouthparts distinctive (see Klimaszewski 1979; Ashe 2000, 2002, 2005; Ashe and Chatzimanolis 2003a, b; Wolf-Schwenninger 2004; Cai and Huang 2015); mandibles with preapical tooth (Gymnusa) or two teeth (Deinopsis); maxillary palpi markedly long, slender, stylate to substylate; pronotum usually transverse, at least as long as wide, posterior margin truncate to bisinuate, narrowing apicad; elytra relatively short; posterolateral margins of elytra deeply emarginate; tarsal formula 3-3-3 (Deinopsis), or 5-5-5 (Gymnusa), terminal tarsomeres without two long setae surrounding the claws (Gymnusa); empodia with two setae; tibiae with irregularly scattered outer spines, front tibiae with preapical longitudinal ctenidium of

setulae in shallow or deep depression (Gymnusa); metacoxae with medium to large ventral lamella; abdominal segments III–VI (i.e., first four visible segments) with tergites, sternites and paratergites each with a dense apical ctenidium of short comblike cuticular projections, consisting of one (Gymnusa) or two (Deinopsis) types of teeth; abdominal segments III–VII each with two pairs of paratergites; tergite VII lacking tergal gland opening on anterior margin, at least for the extant genera; male tergite IX entire (Gymnusa) or divided (Deinopsis). The genus Deinopsis was not recorded from BC.

Genus Gymnusa Gravenhorst, 1806 (Figs. 9.32–9.34) (Klimaszewski Klimaszewski et al. 2018)

1979,

Diagnosis. Body fusiform, length 4.2–7.0 mm; colour predominantly black, pubescence piceous or sometimes with patches of yellowish pubescence on elytra and abdomen; head strongly deflexed, almost as wide as pronotum at anterior margin; frons obtusely and triangularly protruding; vertex glabrous with three setigerous pores on each side of disc (one above antennal insertion, one at anteromedial margin of eye, and one behind eye); clypeus trapezoidal in shape; labrum transverse, broadly rounded; eye oviform; antennal insertion near anteromesal margin of eye, antennae almost filiform, widened toward apex;

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_9

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90

maxillary palpi with four palpomeres, palpomeres I and IV inconspicuous, II and III elongate and each twice as long as length of palpomere I; mandibles with one subapical tooth; labial palpi with three palpomeres; proventrite reduced to narrow transverse plate in front of procoxae; procoxae conical and prominent with cavities open behind; mesocoxae oblique; metacoxae flattened, longer than wide; abdominal tergites III– VII (basal five visible tergites) each with two laterosternites; tergites III–VII each with dense apical comb of teeth with rounded apices; tarsal formula 5-5-5, tarsomere I of all legs at least as long as tarsomere II and III combined; male tergite IX annular, with two lateral lobes and two ventral struts; median lobe of aedeagus with two ventral flaps serving as attachments to parameres; female tergite IX annular, without ventral struts; sclerotized spermatheca absent. Species associated with forested swamps, marshes, and sphagnum bogs. Key to Species of Gymnusa (modified from Klimaszewski 1979, and Klimaszewski et al. 2018) 1. Pubescence on elytra and often on abdomen bicoloured, mostly piceous with a few, small, irregularly distributed, yellowish spots on elytra and abdomen (Fig. 9.34a); body length 4.2–6.0 mm; genitalia and terminalia as illustrated (Fig. 9.34b–h)....................................................... .......... Gymnusa pseudovariegata Klimaszewski – Pubescence on elytra and abdomen uniformly piceous; body length 5.0–6.5 mm; genitalia and terminalia not as above ......................................2 2. Antennomeres I and II approximately subequal in length; body convex (Fig. 9.32a); genitalia and terminalia as illustrated (Fig. 9.32b–h) [females cannot be determined to species unless associated with males] .................. ........................................... Gymnusa atra Casey – Antennomere I longer than II; body flattened (Fig. 9.33a); genitalia and terminalia as illustrated (Fig. 9.33b–i) ......... Gymnusa grandiceps Casey

Brevicollis 1979)

9

Tribe Gymnusini Heer, 1839

Species

Group (Klimaszewski

Diagnosis. Body fusiform, length 5.0–6.5 mm; pubescence uniformly piceous; punctation fine, coarser on elytra and abdomen. 1. Gymnusa atra Casey
(Fig. 9.32a–h) Gymnusa atra Casey, 1911. Klimaszewski 1979, 1982, Klimaszewski and Génier 1985, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2018. Diagnosis. Body convex, length 5.0–6.5 mm, colour black to dark brown, pubescence on elytra and abdomen uniformly piceous, punctation fine, coarser on elytra and abdomen, antennomeres I and II reddish-brown (Fig. 9.32a). MALE. Tergite and sternite VIII as illustrated (Fig. 9.32d, e); median lobe of aedeagus arcuate subapically in lateral view (Fig. 9.32b), with apex sharp in dorsal view (Fig. 9.32c). FEMALE. Tergite VIII with deep apical emargination (Fig. 9.32f); sternite VIII and pygidium as illustrated (Fig. 9.32g, h). This species is not distinguishable externally from G. brevicollis but in G. atra the subapical part of the median lobe of the aedeagus is arcuate in lateral view (Fig. 9.32b) (truncate in G. brevicollis), and the apex of the median lobe is sharp in dorsal view (Fig. 9.32c). Females are not distinguishable and have similar terminalia. DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT. USA: AK, IL, IA, MA, ME, MI, MN, NH, NY. Eastern Russia, northeastern China. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: tamarack (Larix laricina (Du Roi)) and black spruce (Picea mariana (Mill.) BSP)) bogs, marshy pond and lake margins, cattail/Carex marshes, and marshy margins of slow flowing rivers.

9

Tribe Gymnusini Heer, 1839

Adults usually found in Carex hummocks, moist to wet sphagnum with Carex, and in saturated marsh vegetation. Collecting period: V–XI. Collecting method: sifting or treading vegetation (Carex), Carex hummocks, and wet sphagnum into water. Grandiceps 1979)

Species

Group (Klimaszewski

Diagnosis. Body broadly oval, length 5.7–6.5 mm; integument with distinct meshed microsculpture; pubescence piceous; punctation shallow. 2. Gymnusa grandiceps Casey (Fig. 9.33a–i) Gymnusa grandiceps Casey, 1915. Klimaszewski 1979, 1982, Klimaszewski and Génier 1985, Gouix and Klimaszewski 2007, Webster et al. 2009, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2018. Diagnosis. Body widely oval, flattened, length 5.7–6.5 mm, colour dark brown to almost black, microsculpture distinct, pubescence on elytra and abdomen uniformly piceous, punctation shallow (Fig. 9.33a); antennal article I distinctly longer than II. MALE. Tergite and sternite VIII as illustrated (Fig. 9.33e, f); sternite IX deeply emarginate apically (an exception in Gymnusa) (Fig. 9.33d); median lobe of aedeagus in lateral view narrow, broadly arched (Fig. 9.33b); in dorsal view narrowly elongate and produced apically (Fig. 9.33c). FEMALE. Tergite VIII truncate apically (Fig. 9.33g); sternite VIII broadly produced medially at apex (Fig. 9.33h); valvulae of pygidium flat and emarginate apically (Fig. 9.33i). DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], LB, MB, NB, NF, NS, ON, QC. USA: IL, MA, MD, ME, MI, NH, NY, RI. We report this species from BC for the first time here based on the following: BC, Smithers perimeter trail wetland S of Hwy 16, 54.46.22 N,

91

127.09.13 W, 490 m, 31.VII.2012, in moss in swamp, A. Davies (CNC) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: wet forested biotypes such as black spruce bogs (usually in lag zone), alder swamps, spruce and eastern white cedar (Thuja occidentalis L.) forests, eastern white cedar swamps, vernal pond margins, and margins of small brooks under alders. Adults in moist leaf litter and moss in these habitats, and in moist leaf litter in seepage areas in cedar forests. Collecting period: III–IX. Collecting method: Sifting leaf litter and moss. Variegata Species Group (Klimaszewski 1979) Diagnosis. Body fusiform, length 4.2–6.0 mm; pubescence piceous with yellowish spots; interspaces of punctures on elytra and abdomen rugose. 3. Gymnusa pseudovariegata Klimaszewski (Fig. 9.34a–h) Gymnusa pseudovariegata Klimaszewski, 1979. Klimaszewski 1982, Klimaszewski and Génier 1985, Caterino and Chatzimanolis 2007, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2018. Diagnosis. Body length 4.8–6.0 mm, colour black to dark brown, pubescence on elytra and abdomen piceous with small yellow spots, basal antennomeres uniformly dark brown to black; elytra and abdomen with interspaces of punctures rugose (Fig. 9.34a). MALE. Tergite VIII sternite VIII as illustrated (Fig. 9.34d, e); tubus of median lobe of aedeagus in lateral view with small tooth on ventral margin (Fig. 9.34b), in dorsal view apex narrowly produced (Fig. 9.34c). FEMALE. Tergite VIII emarginate apically, without or with minute sclerotized lobe of variable shape and size in middle of apical emargination (Fig. 9.34f); sternite VIII as illustrated (Fig. 9.34g); pygydium

92

with tergite IX having short and acute basal tooth (Fig. 9.34h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT. USA: AK, CA, MA, OR, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: wet marshy habitats and bogs; sedge marshes, cattail/Carex

9

Tribe Gymnusini Heer, 1839

marshes, boggy marshes and fens, tamarack bogs, pool margins in sphagnum bogs, vegetated edges of swamps, lakes, and slow flowing rivers, and small ponds on coastal barrens. Usually among emergent Carex, floating vegetation mats, and wet to saturated marsh vegetation. Also under litter and stones on a sandy lakeshore. Collecting period: IV–XI (easily found in spring and early summer and then declines). Collecting method: treading vegetation into water.

9

Tribe Gymnusini Heer, 1839

Fig. 9.32 (a–h) Images of Gymnusa atra Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

93

tergite VIII; (g) female sternite VIII; (h) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

94

Fig. 9.33 (a–i) Images of Gymnusa grandiceps Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male sternite IX; (e) male tergite VIII; (f) male sternite

9

Tribe Gymnusini Heer, 1839

VIII; (g) female tergite VIII; (h) female sternite VIII; (i) female pygidium. (c) after Klimaszewski (1979). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

9

Tribe Gymnusini Heer, 1839

Fig. 9.34 (a–h) Images of Gymnusa pseudovariegata Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

95

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) female pygidium. (b, c) after Klimaszewski (1979). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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References Ashe JS (2000) Mouthpart structure of Stylogymnusa subantarctica Hammond, 1975 (Coleoptera: Staphylinidae: Aleocharinae) with a reanalysis of the phylogenetic position of the genus. Zool J Linnean Soc 130(4):471–498 Ashe JS (2002) Allodinopsis, new genus of deinopsine aleocharine from Central America, and a new species of Metadeinopsis Klimaszewski 1979 (Staphylinidae: Aleocharinae: Deinopsini). J Kansas Entomol Soc 75 (2):61–72 Ashe JS (2005) Phylogeny of the tachyporine group subfamilies and ‘basal’ lineages of the Aleocharinae (Coleoptera: Staphylinidae) based on larval and adult characteristics. Syst Entomol 30(1):3–37 Ashe JS, Chatzimanolis S (2003a) Gymnusini. Version 06 November 2003. Tree of life web project. http:// tolweb.org/Gymnusini/9814/2003.11.06. Accessed 22 Sept 2016 Ashe JS, Chatzimanolis S (2003b) Deinopsini. Version 06 November 2003. http://tolweb.org/Deinopsini/ 9815/2003.11.06. Accessed 22 Sept 2016 Cai C, Huang D (2015) The oldest aleocharine rove beetle (Coleoptera, Staphylinidae) in Cretaceous Burmese amber and its implications for the early evolution of the basal group of hyper-diverse Aleocharinae. Gondwana Res 28(4):1579–1584 Casey TL (1911) New American species of Aleocharinae and Myllaeninae. In: Casey TL (ed) Memoirs on the Coleoptera, vol II. The New Era Printing Co., Lancaster, PA, pp 1–245. 259 pp Casey TL (1915) Studies in some staphylinid genera of North America. In: Memoirs on the Coleoptera, vol VI. The New Era Printing Co., Lancaster, PA, pp 395–450. 460 pp Caterino MS, Chatzimanolis S (2007) Newly recorded and noteworthy California Staphylinidae. Coleopts Bull 61 (3):398–407 Gouix N, Klimaszewski J (2007) Catalogue of aleocharine rove beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae). Pensoft, Sofia. 165 pp Gravenhorst JLC (1806) Monographia Coleopterorum Micropterorum. Henricus Dieterich, Göttingen, xvi + 236 + [12] pp Heer O (1839) (1838–1841) Fasciculus secundus [pp. 145–360]. In: Fauna Coleopterorum Helvetica.

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Tribe Gymnusini Heer, 1839

Pars I. Turici [Zurich], Orellii, Fuesslini et Sociorum: xii + 652 pp Klimaszewski J (1979) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae: Aleocharinae). Research Branch Agriculture Canada Monograph No. 25:1–169 Klimaszewski J (1982) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae). Supplementum 2. Can Entomol 114(4):317–335 Klimaszewski J, Génier F (1985) A revision of the Gymnusini and Deinopsini of the world (Coleoptera: Staphylinidae, Aleocharinae). Supplement 3. Coleopts Bull 39(1):60–66 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sophia, 313 pp Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon, C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018). Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. , 902 pp Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39 Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys 22:171–248 Wolf-Schwenninger K (2004) A new fossil staphylinid genus and species from Baltic amber (Coleoptera, Staphylinidae, Aleocharinae, Gymnusini). Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie) 345:1–6 Yamamoto S, Maruyama M (2018) Phylogeny of the rove beetle tribe Gymnusini sensu n. (Coleoptera: Staphylinidae: Aleocharinae): implications for the early branching events of the subfamily: phylogeny of Gymnusini rove beetles. Syst Entomol 43 (1):183–199. https://doi.org/10.1111/syen.12267

Tribe Aleocharini Fleming, 1821

Tribe Aleocharini Fleming, 1821 (Figs. 10.35–10.59) (Klimaszewski 1984, Osswald et al. 2013, Klimaszewski et al. 2018) Diagnosis. Body robust to moderately slender (Figs. 10.37a–10.59a), length 2.0–13.0 mm; maxillary and labial palpi with a pseudosegment on last palpomere, appearing five- and foursegmented respectively (Aleochara), or without apparent pseudosegment (Amarochara, not recorded in BC); pronotal hypomera not visible or narrowly visible in lateral view; flagellum of median lobe of aedeagus long, longer than median lobe in most species; some species with velum of paramere reticulated; tarsal formula 5-5-5 in most species, 4-5-5 in the subgenus Tinotus (Klimaszewski et al. 2018). Comments. The genus Tinotus was recently placed in the tribe Aleocharini on the basis of molecular data (Osswald et al. 2013) and is now treated as subgenus of Aleochara. Tinotus is probably most closely related to Coprochara (Yamamoto and Maruyama 2016).

Genus Aleochara Gravenhorst, 1802 (Figs. 10.35–10.59) (Klimaszewski Klimaszewski et al. 2018)

1984,

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Diagnosis. Body length very variable within same species, length within genus 1.8–13.0 mm; body shape broadly oval or subparallel, robust; tarsal formula 5-5-5 except 4-5-5 in Tinotus; last palpomere of maxillary and labial palpi with pseudosegment; laciniae with tuft of apical setae; ligula small, bilobed apically; mesoventrite varies from non-carinate to completely carinate, depending on subgenus; male tergite IX divided by tergite X, each half narrowly elongate anteriad and posteriad and bearing ventral strut; median lobe of aedeagus usually consisting of narrowly elongate tubus and moderate-sized bulbus (except in some species in subgenus Aleochara); internal sac with complex sclerites; spermatheca consisting of capsule (head and neck), chamber and duct (stem), variably modified in different subgenera. Adults are predaceous on larvae, eggs, and pupae of cyclorhaphous Diptera, and larvae are ectoparasitoids of dipteran pupae (Fig. 10.35). Key to Subgenera of Aleochara (Figs. 10.36–10.40) 1. Tarsal formula 4-5-5, body as illustrated (Figs. 10.52a and 10.53a) ......... Tinotus Sharp – Tarsal formula 5-5-5, body as illustrated (Figs. 10.37a–10.51a and 10.54a–10.59a) ......... 2

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_10

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2. Pronotum with two longitudinal, subparallel rows of more-or-less confluent setigerous punctures near midline of disc (Figs. 10.36c and 10.44a–10.48a); mesoventrite with complete longitudinal carina (Fig. 10.36d); spermatheca with multiple, small coils roughly regular in shape and size (Figs. 10.44h, 10.45h, 10.46h, and 10.47i)... .............................Coprochara Mulsant and Rey – Pronotum evenly pubescent, without two longitudinal rows of setigerous punctures near midline of disc; mesoventrite either not carinate, with partial or complete longitudinal carina; spermatheca with or without irregular-shaped posterior coils .. 3 3. Body robust, broad and compact (Figs. 10.36e and 10.37a–10.40a); antennomeres V to X moderately to distinctly transverse and forming an elongate club, each antennomere at least 2.2 times as broad as long (Figs. 10.36e and 10.37a–10.40a); mesoventrite wide and not carinate (Fig. 10.36f); spermatheca often inconspicuous, narrowly elongate with small, spherical or sac-shaped capsule and thin stem (Figs. 10.37h– 10.40h) ................................................................. ...................................... Aleochara Gravenhorst – Body spindle-shaped or subparallel (Figs. 10.36a, g, i and 10.37a–10.40a); antennomeres V to X gradually broadening apically, slightly to moderately transverse, each antennomere 1.2–2.2 times as broad as long; mesoventrite either not carinate or partially to completely longitudinally carinate (Fig. 10.36b, j, h) ........................... 4 4. Mesoventrite completely longitudinally carinate; body moderately to highly convex, spindle-shaped to broadly subparallel (Fig. 10.36b); pronotum with pubescence on disc directed lateroposteriad (Figs. 10.36a and 10.54a–10.59a); elytra at suture at most as long as pronotum......... Xenochara Mulsant and Rey – Mesoventrite not carinate or with short carina no longer than half its length (Fig. 10.36h, j); body moderately flattened, subparallel (Figs. 10.54a–10.59a); pronotum with pubescence on disc directed laterad or obliquely or straight posteriad from midline (Figs. 10.36i, g and 10.54a–10.59a); elytra at suture at least as long as pronotum ...............................................5

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Tribe Aleocharini Fleming, 1821

5. Forebody with strongly pronounced hexagonal microsculpture consisting of convex sculpticells (Figs. 10.49a–10.51a); elytra with pubescence on disc forming wavy pattern (Figs. 10.49a–10.51a); mesoventrite not carinate (Fig. 10.36h); tubus of median lobe of aedeagus with subapical ventral process/es in lateral view (Figs. 10.49b, c, 10.50b and 10.51b); seashore species....Emplenota Casey – Forebody with moderately pronounced hexagonal microsculpture consisting of flattened or moderately convex sculpticells (Figs. 10.41a–10.43a); elytra with pubescence on disc directed obliquely toward outer posterior angles and not forming wavy pattern (Figs. 10.41a–10.43a); mesoventrite partially carinate, carina no longer than half its length (Fig. 10.36j); terrestrial species, with most recorded in association with animal burrows.................................... Calochara Casey Subgenus Aleochara Gravenhorst, 1802 (Figs. 10.36e, f and 10.37a–10.40a) (Klimaszewski 1984, Klimaszewski et al. 2018) Diagnosis. Body robust, length 3.0–9.0 mm; dorsal surface of body moderately to highly glossy, without distinct microsculpture (Figs. 10.36e and 10.37a–10.40a); antennomeres V–X moderately to distinctly transverse; pronotum evenly pubescent; mesoventrite wide and not carinate (Fig. 10.36f); first three visible abdominal tergites with deep transverse depressions; maxillary palpi with apical palpomere at least two-thirds as long as penultimate palpomere; species predominantly associated with carrion. Key to Species of Subgenus Aleochara 1. Antennomere IV moderately transverse and V–X strongly transverse, more than twice as wide as long (Fig. 10.38a); male tergum VIII without clearly serrate apical margin (Fig. 10.38d); abdomen more densely pubescent (Fig. 10.38a); aedeagus and spermatheca as in Fig. 10.38b, c, h....................................... ...........Aleochara (Aleochara) curtula (Goeze) – Antennomere IV subquadrate or slightly elongate (Figs. 10.37a, 10.39a and 10.40a), and V–X

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Tribe Aleocharini Fleming, 1821

subquadrate to moderately transverse (Figs. 10.37a, 10.39a and 10.40a); genitalic characters not as above........................................................................ 2 2. Pronotum red-brown, sparsely pubescent and highly glossy (Fig. 10.37a); sclerites of internal sac of aedeagus as in Fig. 10.37b, c; shape of spermatheca as in Fig. 10.37h................................ Aleochara (Aleochara) assiniboin Klimaszewski – Pronotum brown or black, densely pubescent and moderately to highly glossy (Figs. 10.39a and 10.40a); genitalia not as above ..........................3 3. Body broad and moderately glossy (Fig. 10.40a); internal sac of aedeagus with reversed V-shaped median structure in dorsal view (Fig. 10.40c); spermatheca with elongate chamber (Fig. 10.40h)........ ................ Aleochara (Aleochara) tahoensis Casey – Body narrow and highly glossy (Fig. 10.39a); internal sac of aedeagus with U-shaped median structure in dorsal view (Fig. 10.39c); spermatheca with short chamber (Fig. 10.39h).... Aleochara (Aleochara) gracilicornis Bernhauer 4. Aleochara (Aleochara) assiniboin Klimaszewski (Fig. 10.37a–h) Aleochara assiniboin Klimaszewski, 1985 [replacement name for preoccupied name Aleochara rufonigra Klimaszewski, 1984]. Gouix and Klimaszewski 2007, Klimaszewski et al. 2018. Diagnosis. Body length 4.0–5.0 mm; colour dark brown, pronotum and elytra red-brown, highly glossy, or elytra dark with with two lighter spots, appendages and two basal antennomeres reddishbrown; antennomere IV subquadrate or slightly transverse (Fig. 10.37a). MALE. Tergite VIII truncate apically (Fig. 10.37d); sternite VIII broadly arcuate apically (Fig. 10.37e); tubus of median lobe of aedeagus with ventral margin arcuate in lateral view (Fig. 10.37b); internal sac with three pairs of elongate scelerites (Fig. 10.37c). FEMALE. Tergite VIII truncate apically, apical margin not serrate (Fig. 10.37f); sternite VIII broadly arcuate apically (Fig. 10.37g); spermatheca as illustrated (Fig. 10.37h).

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DISTRIBUTION. Origin: Nearctic. CANADA: BC, MB, ON, SK, YT. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: associated with grasses, Carex, and mosses alongside a small lake; some found under a moose (Alces alces (L.) carcass. Collecting period: V–VIII. Collecting method: treading vegetation and collecting from moose carcass. 5. Aleochara (Aleochara) curtula (Goeze) { (Fig. 10.38a–h) Staphylinus curtulus Goeze, 1777. As Aleochara: Klimaszewski 1984, Klimaszewski and Frank 1992, Sikes 2004, Gouix and Klimaszewski 2007, Klimaszewski et al. 2011, 2018, Majka and Klimaszewski 2010, Majka et al. 2011. Diagnosis. Body widely oval, length 4.0–7.5 mm; colour dark brown to black, with elytra reddish- to rusty-brown except for darker margins, tarsi, palpi and often basal antennomere rusty-brown; antennomeres V–X distinctly transverse; abdomen moderately glossy (Fig. 10.38a). MALE. Tergite VIII truncate, with shallow median emargination (Fig. 10.38d); sternite VIII rounded apically (Fig. 10.38e); median lobe of aedeagus in dorsal view with bulbus large, oval, tubus short, triangular (Fig. 10.38c), in lateral view tubus hooked apically (Fig. 10.38b); internal sac structures complex (Fig. 10.38b, c). FEMALE. Tergite VIII truncate and with shallow median emargination (Fig. 10.38f); sternite VIII rounded apically (Fig. 10.38g); spermatheca C-shaped, capsule spherical with apical invagination deep, elongate (Fig. 10.38h). DISTRIBUTION. Origin: Palearctic: Europe, North Africa, Asia; adventive in North America; likely introduced separately on the east coast and in BC. CANADA: BC, NB, NF, NS, ON, PE, QC. USA: IL, MA, ME, NH, NJ, NY, OH, PA, RI, VT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: open

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habitats such as fields, sea beaches, and river margins. Found on carrion (dead fish, eel, bat, muskrat, and squirrel). Also in other kinds of decaying organic matter such dog scat, sea wrack on sea beaches, and well rotted mushrooms. Collecting period: VI–XI. Collecting method: examining carrion, sifting organic debris, pitfall traps, and dung traps. 6. Aleochara (Aleochara) gracilicornis Bernhauer (Fig. 10.39a–h) Aleochara gracilicornis Bernhauer, 1901. Klimaszewski 1984, Klimaszewski and Génier, 1987, Klimaszewski et al. 2005, 2018, Gouix and Klimaszewski 2007. Diagnosis. Body narrowly oval, length 4.0–6.0 mm, integument highly glossy, colour dark brown to black, with elytra red-brown except for darker margins and scutellar region, legs, palpi and 2–3 basal antennomeres yellow to reddishbrown; antennomere IV slightly elongate, and V– X moderately transverse; abdomen glossy (Fig. 10.39a). MALE. Tergite VIII truncate apically (Fig. 10.39d); sternite VIII arcuate apically (Fig. 10.39e); median lobe of aedeagus in dorsal view with small oval bulbus and long subparallel tubus (Fig. 10.39c), in lateral view with apex narrow and slightly bent ventrad (Fig. 10.39b); internal sac structures as illustrated (Fig. 10.39b, c). FEMALE. Tergite VIII truncate, with shallow median emargination (Fig. 10.39f); sternite VIII rounded apically (Fig. 10.39g); spermatheca sinuate, capsule small, sack-shaped and slightly curved, stem narrow, long and sinuate (Fig. 10.39h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, MB, NB, NF, NS, NT, ON, QC, SK. USA: AR, AZ, CO, FL, IL, IN, KS, LA, MA, MD, ME, MI, MN, MO, MT, NH, NJ, NM, NY, OH, PA, RI, SD, UT, WI. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: river margins in flood debris, drift material and decaying grass and litter, temporary creek margins in debris, lake margins in drift material,

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Tribe Aleocharini Fleming, 1821

hardwood forests and swampy areas in leaves and debris, Carex marsh in leaf litter and moss. Also in an old American beaver (Castor canadensis Kuhl) lodge, in debris on beaver dam (common), and in carrion. Others from pitfall traps in mature red spruce forest. Collecting period: III–X. Collecting method: sifting organic debris. 7. Aleochara (Aleochara) tahoensis Casey (Fig. 10.40a–h) Aleochara tahoensis Casey, 1906. Hatch 1957, Moore and Legner 1975, Klimaszewski 1984, Klimaszewski and Génier 1987, Klimaszewski et al. 2005, 2015, 2018, Gouix and Klimaszewski 2007. Diagnosis. Body broadly oval, length 4.5–7.0 mm; colour dark brown to black with elytra reddish-brown except antero-laterad margins dark brown, legs, maxillary palps, and often basal antennomere red-brown; antennomere IV slightly elongate and V–X moderately transverse (Fig. 10.40a). MALE. Tergite VIII with apical margin slightly emarginate medially (Fig. 10.40d); sternite VIII with apical margin broadly rounded and slightly pointed at apex (Fig. 10.40e); median lobe of aedeagus in dorsal view with bulbus moderately large, oval, tubus narrowly triangular (Fig. 10.40c); in lateral view tubus with ventral margin straight with apex narrow, slightly produced ventrad (Fig. 10.40b); internal sac structures complex, as illustrated (Fig. 10.40b, c). FEMALE. Tergite VIII truncate and slightly emarginate medially (Fig. 10.40f); sternite VIII arcuate apically (Fig. 10.40g); spermatheca thin and sinuate (Fig. 10.40h). This species is similar externally to A. gracilicornis, but in A. tahoensis the body is broader, antennomeres V–X are more transverse, the distance between the eyes is equal to 2.5 times the maximum diameter of the eye, and the genitalic features are distinctive. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NF, NS, NT, ON, SK, YT. USA: AR, CA, CO, MT, NH, NM, NV, OR, SD, WA, WI.

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Tribe Aleocharini Fleming, 1821

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: forests and swampy areas in flood debris, debris around dead elm, under moose carcass, in leaf litter on a beaver dam, and along a lake shore in drift material. Some collected from pitfall traps in mature red spruce (Picea rubens Sarg.) forest. Collecting period: V–IX. Collecting method: sifting litter, pitfall traps. Subgenus Calochara Casey, 1906 (Figs. 10.36i, j and 10.41–10.43) (Klimaszewski 1984, Klimaszewski et al. 2018) Diagnosis. Forebody with moderately pronounced hexagonal microsculpture consisting of flattened or moderately convex sculpticells (Figs. 10.41a–10.43a); pronotum evenly pubescent (Figs. 10.41a–10.43a); mesoventrite narrow, wedge-shaped with carina no longer than half its length (Fig. 10.36j); elytra with pubescence on disc directed obliquely toward outer posterior angles and not forming wavy pattern; male sternite VIII triangularly produced apically (Figs. 10.41e–10.43e); median lobe of aedeagus narrow, strongly bent ventrally and with two pairs of elongate sclerites (Figs. 10.41b, 10.42b and 10.43b); spermatheca with capsule more-or-less spherical, stem usually thin and simple (Figs. 10.41h, 10.42h, and 10.43h); terrestrial species associated with animal nests and burrows. Key to Species of Subgenus Calochara 1. Elytra rusty-brown, at suture shorter than maximum length of pronotum (Fig. 10.41a); tubus of median lobe of aedeagus with ventral margin strongly curved and apically pointed ventrad in lateral view (Fig. 10.41b); internal sac of median lobe of aedeagus with elongate structures, as illustrated (Fig. 10.41b, c); spermatheca as illustrated (Fig. 10.41h)........................................ ...... Aleochara (Calochara) laramiensis (Casey) – Elytra uniformly black or black with traces of reddish tinge, at suture at least as long as maximum length of pronotum (Figs. 10.42a and 10.43a); genitalia shaped differently.............2

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2. Body moderately narrow; integument highly glossy, elytra black with reddish tinge, microsculpture of forebody weak (Fig. 10.42a); tubus of median lobe of aedeagus with ventral margin strongly curved in lateral view (Fig. 10.42b), internal sac structures of median lobe of aedeagus narrowly elongate with apex hooked (Fig. 10.42b, c); spermatheca with stem evenly narrow (Fig. 10.42h) ................................ ......... Aleochara (Calochara) rubricalis (Casey) – Body broad; integument moderately glossy, elytra usually uniformly black, microsculpture of forebody distinct (Fig. 10.43a); median lobe of aedeagus with tubus less curved, median part straight, in lateral view (Fig. 10.43b), internal sac with falciform and apically rounded structures (Fig. 10.43b, c); spermatheca with spherical capsule and swollen chamber (Fig. 10.43h)...................................................... ...Aleochara (Calochara) villosa Mannerheim 8. Aleochara (Calochara) laramiensis (Casey) (Fig. 10.41a–h) Oreochara laramiensis Casey, 1906. Klimaszewski 1984, Gouix and Klimaszewski 2007. Diagnosis. Body length 5.0–8.0 mm; brown to black with elytra and legs rusty-brown (Fig. 10.41a); pubscence sparse and yellowish brown; antennomeres V–X moderately transverse (Fig. 10.41a); maxillary palpi with apical palpomere conical; elytra shorter than pronotum (Fig. 10.41a). MALE. Tergite VIII with apical margin truncate (Fig. 10.41d); sternite VIII with apical part triangularly produced (Fig. 10.41e); median lobe of aedeagus in dorsal view with bulbus narrowly oval, tubus narrowly subparallel, slightly broadening apically (Fig. 10.41c); in lateral view tubus with ventral margin strongly arcuate, with apex bent ventrad and pointed (Fig. 10.41b); internal sac structures as illustrated (Fig. 10.41b, c). FEMALE. Tergite VIII truncate, slightly emarginate medially (Fig. 10.41f); sternite VIII rounded apically (Fig. 10.41g); spermatheca with hemispherical capsule and thin, sinuate stem, as illustrtrated (Fig. 10.41h).

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DISTRIBUTION. Origin: Nearctic. CANADA: BC, SK. USA: CO, WY. COLLECTION AND HABITAT DATA. Habitat: not recorded, may be associated with animal burrows. Collecting period: not recorded. Collecting method: not recorded. 9. Aleochara (Calochara) rubricalis (Casey) (Fig. 10.42a–h) Baryodma rubricalis Casey, 1911. Klimaszewski 1984, Gouix and Klimaszewski 2007, Klimaszewski et al. 2016b, 2018, Webster et al. 2016. Diagnosis. Body length 2.5–4.0 mm; colour dark brown to black, with tarsi, labial and maxillary palps, posterior or sutural part of elytra and apex of abdomen reddish-brown; sparsely pubescent (Fig. 10.42a); antennomeres V–X transverse (Fig. 10.42a); maxillary palpi with apical palpomere more than half as long as penultimate one; elytra at suture at least as long as pronotum (Fig. 10.42a). MALE. Tergite VIII with apical margin arcuate (Fig. 10.42d); sternite VIII with apical part triangularly produced (Fig. 10.42e); median lobe of aedeagus in dorsal view with bulbus small, round, tubus narrowly subparallel (Fig. 10.42c); in lateral view tubus with ventral margin strongly arcuate, with apex bent ventrad (Fig. 10.42b); internal sac structures as illustrated (Fig. 10.42b, c). FEMALE. Tergite VIII truncate, slightly emarginate medially (Fig. 10.42f); sternite VIII rounded apically (Fig. 10.42g); spermatheca L-shaped (Fig. 10.42h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, ON [cited as doubtful in ON, Klimaszewski 1984], SK. USA: AZ, CA, OR, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: collected from Lindgren funnel traps in an old black spruce (Picea mariana (Mill.) BSP) forest, an old-growth white spruce (Picea glauca (Moench)

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Tribe Aleocharini Fleming, 1821

Voss) and balsam fir (Abies balsamea (L.) Mill.) forest, and an old balsam poplar stand (Populus balsamifera L.) near a river. One specimen was collected from a mouse nest and another from decaying mushrooms in a regenerating (8 yearsold) mixed forest. Collecting period: II–XI, in Canada V, VI, X. Collecting method: Lindgren funnel traps, sifting litter. 10. Aleochara (Calochara) villosa Mannerheim { (Fig. 10.43a–h) Aleochara villosa Mannerheim, 1830. Klimaszewski 1984, Klimaszewski and Frank 1992; Gouix and Klimaszewski 2007, Webster et al. 2009, Klimaszewski et al. 2018. Diagnosis. Body moderately broad, length 3.0–5.0 mm; colour dark brown to black, with tarsi, and last palpomere of maxillary palps often reddish-brown (Fig. 10.43a); pubescence sparse; forebody with meshed microsculpture; antennomeres V–X moderately transverse (Fig. 10.43a); apical palpomere of maxillary palpus about as long as half of penultimate one. MALE. Tergite VIII sinuate apically, with distinct pattern of macrosetae (Fig. 10.43d); sternite VIII with apical margin broadly triangularly produced, sinuately emarginate on either side, with apex narrowly rounded (Fig. 10.43e); median lobe of aedeagus in dorsal view with bulbus elongate, oval, evenly coalescent with triangular tubus (Fig. 10.43c); in lateral view tubus with ventral margin straight, broadly curved ventrad at base, with apex narrow, slightly curved ventrad (Fig. 10.43b); internal sac with dorsal part sclerotized in lateral view (Fig. 10.43b), with structures as illustrated (Fig. 10.43b, c). FEMALE. Tergite VIII truncate apically (Fig. 10.43f); sternite VIII rounded apically (Fig. 10.43g); spermatheca U-shaped, similar in shape to that of A. speculicollis (Fig. 10.43h). DISTRIBUTION. Origin: Palearctic or Holarctic, if the former then adventive in North America. CANADA: AB, BC, NB, ON, QC, SK. USA: AK, CA, OK, OR, WA.

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Tribe Aleocharini Fleming, 1821

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: nest contents of barred owls (many sites in NB), a great horned owl, Bubo virginianus (Gmelin) and probably occurs in other tree hole nesting birds and mammals. Adults in moist smelly (urine smell) organic material consisting of small animal and bird remains, fur, feathers, and owl pellets, and many fly larvae. Nests were in tree holes in large trees in old mixed forests, old hardwood forests, an old silver maple forest (Acer saccharinum L.), and a black spruce forest. Other individuals from old pile of hay and in carrion. Collecting period: II–VIII. Collecting method: sifting litter. Subgenus Coprochara Mulsant and Rey, 1874 (Figs. 10.36c, d and 10.44–10.48) (Klimaszewski 1984, Klimaszewski et al. 2018) Diagnosis. Pronotal disc with two longitudinal, subparallel rows of setigerous punctures laterad to median line (Figs. 10.36c and 10.44–10.48), punctures fine or coarse, sometimes confluent, with interspace along midline glabrous or with fine punctation; mesoventrite completely carinate longitudinally (Fig. 10.36d); posterior stem of spermatheca with multiple coils (Figs. 10.44h– 10.48h).

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with two small yellow to reddish spots (Figs. 10.45a, 10.46a, and 10.48a) ....................... 3 3. Body narrowly subparallel (Fig. 10.48a); elytral spots small, orange to rust-brown; pronotal punctation coarse (Fig. 10.48a); genitalia as illustrated (Fig. 10.48b, c, h)................................ ....................Aleochara (Coprochara) verna Say – Body widely oval (Figs. 10.45a and 10.46a); elytral spots large, yellowish-brown (Fig. 10.45a), or entire elytra yellow red or yellow brown (Fig. 10.46a) ......................................................... 4 4. Pronotal punctation moderately coarse, surface moderately glossy (Fig. 10.45a); genitalia as illustrated (Fig. 10.45b, c, h)................................ .................. Aleochara (Coprochara) bimaculata Gravenhorst – Pronotal punctation weak, surface strongly glossy (Fig. 10.46a); genitalia as illustrated (Fig. 10.46b, c, h)................................................. .......... Aleochara (Coprochara) suffusa (Casey) 11. Aleochara (Coprochara) bilineata Gyllenhal { (Fig. 10.44a–h) Aleochara bilineata Gyllenhal, 1810. Klimaszewski 1984, Hemachandra et al. 2005, Klimaszewski et al. 2005, 2013, 2018, Gouix and Klimaszewski 2007.

Key to Species of Subgenus Coprochara 1. Pronotal subparallel rows of punctures situated in two prominent grooves (Fig. 10.47a); genital structures as illustrated (Fig. 10.47b–d, i)............ .................... Aleochara (Coprochara) sulcicollis Mannerheim – Pronotal subparallel rows of punctures not in deep grooves (Figs. 10.44a–10.46a); genital structures not as above ......................................... 2 2. Body unicoloured, dark brown to black (Fig. 10.44a), sometimes elytra slightly paler than rest of body; genitalia as illustrated (Fig. 10.44b, c, h)................................................. .... Aleochara (Coprochara) bilineata Gyllenhal – Body bicoloured, elytra reddish-brown with basal-sutural area darker, or dark brown to black

Diagnosis. Body moderately broad, subparallel, length 2.0–6.0 mm; colour piceous black, elytra entirely black or with posteromedial angle reddish; tarsi, palps, and sometimes basal antennomere red-brown (Fig. 10.44a); antennomeres V–X moderately transverse (Fig. 10.44a). MALE. Tergite VIII truncate apically with row of short setae (Fig. 10.44d); sternite VIII with apical margin broadly triangularly produced, with apex broadly rounded (Fig. 10.44e); median lobe of aedeagus and internal sac structures as illustrated (Fig. 10.44b, c). FEMALE. Tergite VIII with apical margin truncate, slightly emarginate medially (Fig. 10.44f); sternite VIII rounded apically (Fig. 10.44g); spermatheca with capsule spherical, stem with 5–7 tight posterior coils (Fig. 10.44h).

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10

DISTRIBUTION. Origin: Palearctic; adventive in North America CANADA: AB, BC, MB, NF, NS, NB, ON, QC, PE, SK. USA: ID, IL, MA, ME, NY, OR, RI, WA, WI. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: open habitats such as agricultural fields (cabbage and turnip), lake and forest margins, including jack pine (Pinus banksiana Lamb.) forests, mixed forests, and spruce-fir forests. Many adults from Canada reared from puparia from agricultural fields. Adults also found in coyote dung, bear dung, and rotten Boletus mushrooms. Others collected in yellow pan traps in potato fields. One from pitfall trap in mature red spruce forest. Collecting period: VI–IX. Collecting method: Sifting dung and mushrooms, yellow pan traps, pitfall traps, and collecting puparia of Anthomyiidae, Calliphoridae and Muscidae from rotting cabbage and turnips and rearing them in laboratory. Comments (Klimaszewski 1984, Klimaszewski et al. 2018). Most adults examined in Canada emerged in the laboratory from the puparia of Anthomyiidae, Calliphoridae and Muscidae. Adults were collected during the summer months. This species is used in Canada for biological control of the cabbage root maggot, Delia radicum (L.). The beetle larvae feed on D. radicum puparia and the adults feed on dipteran larvae. It was intentionally introduced into Canada in 1950 by the Belleville laboratory, Canadian Department of Agriculture (Hemachandra et al. 2005). However, it was described in 1916 under the synonymic name of Baryodma ontarionis Casey based on specimens reared from D. radicum at Ottawa in 1910 (Gibson and Treherne 1916), and was apparently reported from Ottawa as early as 1901 (Fletcher 1902, as Aleochara nitidum). This species now lives in natural habitats.

Tribe Aleocharini Fleming, 1821

1992, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2018. Diagnosis. Body broadly oval, length 4.0–8.0 mm; colour piceous black, with tarsi, palps, and basal antennomere rusty-brown, each elytron with a large yellowish-brown spot in apical median part of disc (Fig. 10.45a); pronotum moderately glossy; setigerous punctures on head and pronotum slightly impressed; antennomeres V–X distinctly transverse (Fig. 10.45a). MALE. Tergite VIII with apical margin arcuate, with row of long setae (Fig. 10.45d); sternite VIII with apical margin arcuate (Fig. 10.45e); median lobe of aedeagus and internal sac structures as illustrated, including very long flagellum coiled in bulbus (Fig. 10.45b, c). FEMALE. Tergite VIII truncate apically (Fig. 10.45f); sternite VIII rounded apically (Fig. 10.45g); spermatheca with capsule spherical, with deep, broad apical invagination, stem with at least eight tight posterior coils (Fig. 10.45h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK. USA: AL, AZ, CA, CO, CT, DC, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY. MEXICO: Mexico City, Oaxaca, Puebla. BRAZIL. CHILE. COLOMBIA. ARGENTINA. ECUADOR.

bimaculata

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: diverse open and treed habitats; adults found in human feces and other animal droppings such as cow, horse, deer, dog; carcasses of dead animals such as rabbit, fish; decaying fungi, moss and leaf litter along streams, and plant debris in beaver lodges. Collecting period: I–XI (mainly summer months). Collecting method: sifting decaying organic matter, dung- and carrion-baited pitfall traps.

Aleochara bimaculata Gravenhorst, 1802. Klimaszewski 1984, Klimaszewski and Frank

Comments. This species may be used for biological control of the horn and face fly (Klimaszewski 1984).

12. Aleochara (Coprochara) Gravenhorst (Fig. 10.45a–h)

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Tribe Aleocharini Fleming, 1821

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13. Aleochara (Coprochara) suffusa (Casey)
(Fig. 10.46a–h) Baryodma suffusa Casey, 1906. Klimaszewski 1984, Klimaszewski and Cervenka 1986, Klimaszewski and Frank 1992, Gouix and Klimaszewski 2007, Klimaszewski et al. 2016b, 2018. Diagnosis. Body broadly oval, length 2.8–4.0 mm; colour dark brown, with elytra, tarsi, palps, basal antennomeres yellow-brown or reddish-brown (Fig. 10.46a); setigerous punctures on head and pronotum fine; pronotum sparsely pubescent and highly glossy, two subparellel rows of punctures slightly impressed; antennomeres V–X moderately transverse (Fig. 10.46a). MALE. Tergite VIII with apical margin truncate, with row of long setae (Fig. 10.46d); sternite VIII with apical margin slightly produced medially, rounded at apex (Fig. 10.46e); median lobe of aedeagus similar to that of A. verna, in lateral view with ventral margin of tubus broadly curved ventrad at middle, with apex narrow, slightly bent ventrad (Fig. 10.46b), sclerites of internal sac as illustrated (Fig. 10.46b, c). FEMALE. Tergite VIII truncate apically (Fig. 10.46f); sternite VIII slightly produced apically (Fig. 10.46g); spermatheca with capsule club-shaped, stem widely U-shaped, with up to 18 tight posterior coils (Fig. 10.46h). DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, MB, QC, SK. USA: AK, AZ, CO, ID, NM, WA, WY. COLLECTION AND HABITAT DATA. Habitat: open habitats and woodlands. Adults found under rocks in a meadow. Collecting period: VII–VIII. Collecting method: hand picking from under rocks and rearing in laboratory from dipteran pupae collected in fields. 14. Aleochara (Coprochara) Mannerheim
(Fig. 10.47a–i)

sulcicollis

Aleochara sulcicollis Mannerheim, 1843. Klimaszewski 1984, Gouix and Klimaszewski 2007.

Diagnosis. Body length 3.0–7.0 mm; colour piceous to black, with tarsi and antennae redish brown; setigerous punctures on head and pronotum coarse and deeply impressed, those along midline of pronotum in grooves (Fig. 10.47a); antennae short, antennomeres V– X distinctly transverse (Fig. 10.47a). MALE. Tergite VIII with apical margin truncate, with row of very short setae (Fig. 10.47e); sternite VIII with apical margin slightly produced medially, rounded at apex (Fig. 10.47f); median lobe of aedeagus in lateral view with ventral margin of tubus broadly curved ventrad from base with apex narrowly triangular, slightly bent ventrad at tip in lateral view (Fig. 10.47b, c), sclerites of internal sac as illustrated (Fig. 10.47b–d). FEMALE. Tergite VIII truncate apically (Fig. 10.47g); sternite VIII slightly produced apically (Fig. 10.47h); spermatheca with capsule spherical, stem sinuate, with numerous (up to 80) tight posterior coils (Fig. 10.47i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, OR, WA. COLLECTION AND HABITAT DATA. Habitat: seaweed on sea beaches. Collecting period: II–X. Collecting method: hand picking from seaweed, sifting seaweed and aspirating specimens. 15. Aleochara (Coprochara) verna Say (Fig. 10.48a–h) Aleochara verna Say, 1836. Klimaszewski 1984, Hemachandra et al. 2005, Klimaszewski et al. 2005, 2011, 2018, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010. Diagnosis. Body narrowly subparallel, length 2.0–4.0 mm; colour piceous to black, with tarsi, palps, and basal antennomere rusty-brown, each elytron with yellowish to rusty-brown spot in apico-median part of disc (Fig. 10.48a); setigerous punctures on head and pronotum coarse; two subparellel pronotal punctures not in grooves; antennomeres V–X distinctly transverse (Fig. 10.48a). MALE. Tergite VIII with apical

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margin truncate, with row of short setae (Fig. 10.48d); sternite VIII with apical margin slightly produced medially, rounded at apex (Fig. 10.48e); median lobe of aedeagus in lateral view with ventral margin of tubus broadly curved ventrad from base with apex narrowly triangular, slightly bent ventrad at tip (Fig. 10.48b), sclerites of internal sac as illustrated (Fig. 10.48b, c). FEMALE. Tergite VIII truncate apically (Fig. 10.48f); sternite VIII slightly produced apically (Fig. 10.48g); spermatheca with capsule club-shaped, stem widely U-shaped, with 9–11 tight posterior coils (Fig. 10.48h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT. USA: AK, AZ, CA, CO, DC, FL, GA, IA,ID, IL, IN, KS, MA, ME, MI, MN, MO, MT, NH, NC, NJ, NM, NY, NV, OK, OR, PA, SC, SD, TN, TX, UT, VT, WA, WI, WY. MEXICO: Guerrero. GUATEMALA. COLLECTION AND HABITAT DATA. Habitat: agricultural fields, pastures, forests, coastal limestone barrens, sea beaches, and on a sandy area near the ocean. Adults in human feces and other animal droppings such as cows, horse, dog, woodchuck, moose, coyote; in carcasses of dead animals such as deer, squirrel, and coyote; under beach drift material and decaying sea wrack; in rotting fungi, in moss and decaying vegetable matter; some specimens emerged from cabbage and radish maggots (Diptera, Anthomyiidae). Many specimens from pitfall traps in a mature red spruce forest in NB. Collecting period: VI–IX. Collecting method: pitfall traps, malt traps, light trap, sifting decaying organic matter. Comments. Recent records of this species by Schülke and Smetana (2015) from Europe, North Africa and Asia need confirmation. Subgenus Emplenota Casey, 1884 (Figs. 10.36g, h and 10.49–10.51) (Klimaszewski 1984, Klimaszewski et al. 2018)

10

Tribe Aleocharini Fleming, 1821

Diagnosis. Body subparallel; forebody with strongly pronounced hexagonal microsculpture consisting of convex sculpticells (Figs. 10.49a– 10.51a); elytra with pubescence on disc forming wavy pattern; mesoventrite not carinate (Fig. 10.36h); tubus of median lobe of aedeagus with subapical ventral process/es in lateral view; seashore species. Key to Species of Emplenota 1. Antennae as long as pronotum or only slightly longer (Fig. 10.50a); elytra red-brown, less contrasting with dark brown or black remainder of body (Fig. 10.50a) ........................................... ..........Aleochara (Emplenota) litoralis (Mäklin) – Antennae as long as pronotum and half length of elytra (Figs. 10.49a and 10.51a); elytra yellowor redbrown, strongly contrasting with dark brown remainder of body (Figs. 10.49a and 10.51a) ...............................................................2 2. Tubus of median lobe of aedeagus with one median projection in lateral view (Fig. 10.49b, c); female sternite VIII rounded apically (Fig. 10.49h); spermatheca as illustrated (Fig. 10.49i, j)............... Aleochara (Emplenota) curtidens Klimaszewski – Tubus of median lobe of aedeagus with two median projections in lateral view (Fig. 10.51b); female sternite VIII triangularly produced apically and pointed medially (Fig. 10.51g); spermatheca as illustrated (Fig. 10.51h).......................... ............Aleochara (Emplenota) pacifica (Casey) 16. Aleochara (Emplenota) Klimaszewski (Fig. 10.49a–j)

curtidens

Aleochara (Emplenota) curtidens Klimaszewski 1984. Diagnosis. Body subparallel, length 3.5–7.0 mm; colour black, with tarsi and palps rusty-yellowish brown, and elytra red-brown with scutellar and sutural areas of disc darker, sometimes entire body dark brown or black (Fig. 10.49a);

10

Tribe Aleocharini Fleming, 1821

integument with coarse meshed microsculpture, sculpticells flat; antennae as long as pronotum and half length of elytra, antennomeres V–X moderately transverse, less than twice as wide as long (Fig. 10.49a). MALE. Tergite VIII with apical margin shallowly emarginated medially (Fig. 10.49e); sternite VIII with apical margin broadly rounded (Fig. 10.49f); median lobe of aedeagus in lateral view with bulbus mediumsized, elongate, crista apicalis narrowly elongate, tubus with ventral margin broadly curved from base, with one projection, as illustrated (Fig. 10.49b, c), in dorsal view as illustrated (Fig. 10.49d). FEMALE. Tergite VIII with apical margin truncate, slightly emarginate medially (Fig. 10.49g); sternite VIII broadly rounded apically (Fig. 10.49h); spermatheca tubular, of complex shape, as illustrated (Fig. 10.49i, j). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA. COLLECTION AND HABITAT DATA. Habitat: seashores, especially sandy beaches. In rotting seaweed and organic debris. Collecting period: I, VI–VIII. Collecting method: sifting rotting seaweed and organic debris. 17. Aleochara (Emplenota) litoralis (Mäklin) (Fig. 10.50a–h) Homalota litoralis Mäklin, 1853. Klimaszewski 1984, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2018. Diagnosis. Body subparallel, length 3.5–7.0 mm; colour black, with tarsi and palps rusty-brown, or black with elytra red-brown with scutellar area and sides of disc darker; integument with coarse meshed microsculpture, sculpticells flat (Fig. 10.50a); antennae short, as long as pronotum or only slightly longer, antennomeres V–X twice as wide as long (Fig. 10.50a). MALE. Tergite VIII with apical margin truncate, finely crenulate

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(Fig. 10.50d); sternite VIII with apical margin very broadly triangularly produced medially, sinuate laterally (Fig. 10.50e); median lobe of aedeagus in lateral view with bulbus medium-sized, elongate, crista apicalis narrowly elongate, tubus with ventral margin broadly curved from base, with two projections, apex narrow, elongate (Fig. 10.50b), in dorsal view as illustrated (Fig. 10.50c). FEMALE. Tergite VIII with apical margin truncate, slightly emarginate medially (Fig. 10.50f); sternite VIII slightly produced apically (Fig. 10.50g); spermatheca of a complex shape, as illustrated (Fig. 10.50h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, NF, NS, QC. USA: AK, CA, FL, MA, NJ, NY, RI, VA. COLLECTION AND HABITAT DATA. Habitat: seashores, especially sandy beaches and nearby dunes. In rotting seaweed and organic debris. Collecting period: II–VIII. Collecting method: sifting rotting seaweed and organic debris. 18. Aleochara (Emplenota) pacifica (Casey) (Fig. 10.51a–h) Polistoma pacifica Casey, 1893. Klimaszewski 1984. Diagnosis. Body subparallel, length 4.0–8.0 mm; colour black, with tarsi and palps rusty-brown, elytra red-brown with scutellar area and base of disc darker; integument with coarse meshed microsculpture, sculpticells flat (Fig. 10.51a); antennae as long as pronotum and half length of elytra antennomeres V–X one and a half as wide as long (Fig. 10.51a). MALE. Tergite VIII with apical margin truncate, finely crenulate (Fig. 10.51d); sternite VIII with apical margin broadly rounded (Fig. 10.51e); median lobe of aedeagus in lateral view with bulbus medium-sized, elongate, crista apicalis narrowly elongate, tubus with ventral margin broadly curved from base, with two parallel projections and bump-shaped small projection

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posterior to two processes, apex narrow, elongate (Fig. 10.51b), in dorsal view median lobe as illustrated (Fig. 10.51c). FEMALE. Tergite VIII with apical margin truncate (Fig. 10.51f); sternite VIII slightly produced apically, apex pointed (Fig. 10.51g); spermatheca of complex shape, as illustrated (Fig. 10.51h). Differing externally form similar A. litoralis by the less transverse antennomeres V–X, weaker body microsculpture, broadly rounded apex of male sternite VIII, and the genital characters. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, WA. MEXICO: Baja California Norte (Klimaszewski 1984). COLLECTION AND HABITAT DATA. Habitat: seashores, especially sandy beaches. In rotting seaweed and organic debris. Collecting period: III–X. Collecting method: sifting rotting seaweed and organic debris. Subgenus Tinotus Sharp, 1883 (Figs. 10.52 and 10.53) (Klimaszewski et al. 2002, 2018, Yamamoto and Maruyama 2016) Diagnosis. Species of this subgenus differ from all other Aleochara species in having a 4-5-5 tarsal formula (5-5-5 in all other Aleochara). Key to Species of Subgenus Tinotus 1. Body evenly dark brown to black (Fig. 10.53a); maximum width of elytra greater than that of pronotum, elytra at suture shorter than pronotum along midline (Fig. 10.53a); tubus of median lobe of aedeagus with large rounded basal projection in lateral view (Fig. 10.53b); spermatheca with 1–2 coils (Fig. 10.53g)......................................... ......... Aleochara (Tinotus) morion Gravenhorst – Body evenly red brown (Fig. 10.52a); maximum width of elytra equal to that of pronotum, elytra at suture as long as pronotum along midline (Fig. 10.52a); tubus of median lobe of aedeagus straight, without large rounded projection in lateral view (Fig. 10.52b); female

10

Tribe Aleocharini Fleming, 1821

undescribed........................................................ ................ Aleochara (Tinotus) acerba (Casey) 19. Aleochara (Tinotus) acerba (Casey), New Generic Combination (Fig. 10.52a–d) Acrimea acerba Casey, 1911. Bousquet et al. 2013 [as Tinotus]. Lectotype (male): USA, Wash. [ington]; N. Yakima; acerba Csy.; Type USNM 39737; Casey bequest 1925; Lectotypus Acrimea acerba Casey, Gusarov 2003; our lectotype designation label (USNM). Present designation. Diagnosis. Body narrowly oval, robust, length 2.8 mm; colour red brown, head, apical parts of antennae and bases of abdominal tergites dark brown (Fig. 10.52a); surface of forebody densely pubescent, moderately glossy, integument with meshed microsculpture; head small, deflexed, almost half as wide as maximum width of pronotum (Fig. 10.52a); antennomeres IV–X slightly to highly transverse, incrassate; pronotum distinctly wider than head and about as wide as elytra; maximum width of elytra equal to that of pronotum, elytra at suture as long as pronotum along midline; abdominal impressions with deep, round or longitudinally elongate pores. MALE. Tergite VIII with apical margin slightly emarginate medially, margin finely serrate (Fig. 10.52c); sternite VIII parabolic apically (Fig. 10.52d); median lobe of aedeagus in lateral view with tubus straight and pointed apically (Fig. 10.52b). FEMALE. Undescribed. DISTRIBUTION. Origin: Neractic. CANADA: BC. USA: ID, WA. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. We have designated a lectotype for this species because the holotype was not originally designated and Gusarov’s designation was never published.

10

Tribe Aleocharini Fleming, 1821

20. Aleochara (Tinotus) morion Gravenhorst { (Fig. 10.53a–g) Aleochara morion Gravenhorst, 1802. Klimaszewski et al. 2002, 2005, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2008, Bousquet et al. 2013 [as Tinotus], Klimaszewski et al. 2011, 2015. Diagnosis. Body narrowly oval, robust, length 1.8–3.0 mm; colour entirely black, rarely brown with appendages and sometimes elytra paler, reddish, surface densely pubescent, moderately glossy, integument with meshed microsculpture (Fig. 10.53a); head strongly deflexed; antennomeres IV–X slightly to highly transverse, about twice as wide as long; maximum width of elytra greater than that of pronotum, elytra at suture shorter than pronotum along midline; abdominal impressions with deep, round or longitudinally elongate pores. MALE. Tergite VIII with apical margin crenulate, slightly emarginate medially (Fig. 10.53c); sternite VIII rounded apically and slightly pointed medially (Fig. 10.53d); median lobe of aedeagus in lateral view with tubus weakly produced apically, bearing large rounded basal projection (Fig. 10.53b). FEMALE. Tergite VIII with apical margin truncate, slightly emarginate (Fig. 10.53e); sternite VIII broadly rounded apically (Fig. 10.53f); spermatheca L-shaped, capsule sac-shaped and bent laterally, stem with 1–2 posterior coils (Fig. 10.53g). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, NB, NF, NS, ON, QC, SK. USA: CA, CT, NV. COLLECTION AND HABITAT DATA. Habitat: open habitats and various forest types, in decaying organic matter, fungi, animal droppings, human feces, and carrion. Collecting period: II– IX. Collecting method: sifting organic debris, carrion baited pitfall traps.

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Comments. This species arrived in North America some considerable time ago, judging from its widespread distribution in Canada and the USA (Klimaszewski et al. 2002). It was first captured and reported in Canada in 1929, in Knowlton, Quebec (Klimaszewski et al. 2002), and in the USA in Nevada before 1889 (Fauvel 1889; Klimaszewski et al. 2013, 2018). Subgenus Xenochara Mulsant and Rey, 1874 (Figs. 10.36a, b and 10.54–10.59) (Klimaszewski 1984, Klimaszewski et al. 2017, 2018) Diagnosis. Body moderately to highly convex, spindle-shaped to broadly subparallel; antennomeres V–X gradually broadening apically, slightly to moderately transverse, each antennomere 1.2–2.2 times as broad as long; pronotum evenly pubescent, pubescence on disc directed lateroposteriad (Figs. 10.36a and 10.54a–10.59a); mesoventrite with complete or almost complete longitudinal median carina (Fig. 10.36b); maxillary palpi with apical palpomere one-third to three-quarters as long as penultimate one; median lobe of aedeagus in dorsal view with bulbus moderate-sized, tubus narrowly elongate, subparallel basally and at middle (Figs. 10.55d, 10.56d, 10.57d, 10.58c, and 10.59c), spermatheca diverse in shape but capsule usually spherical (Figs. 10.54d, e, 10.55i, 10.56i, j, 10.57i, 10.58h, and 10.59h). Comments (Klimaszewski et al. 2018). There are many different evolutionary linages in this subgenus that were formerly recognized as species groups (see Klimaszewski 1984). Habitats are diverse, including various decaying organic matter, rotting mushrooms, moss, animal droppings, and forest litter. Key to Species of Subgenus Xenochara 1. Abdominal tergites with distinctive imbricate microsculpture (Fig. 10.58a); genitalia as illustrated

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(Fig. 10.58b, c, h) ..................................................... ................Aleochara (Xenochara) lacertina Sharp – Abdominal tergites without imbricate microsculpture....................................................2 2. Body broad, oval and robust; posteromedial angle of each elytron with reddish-spot or reddish margin (Fig. 10.59a); genitalia as illustrated (Fig. 10.59b, c, h)................................................. ....................Aleochara (Xenochara) lanuginosa Gravenhorst – Body narrowly elongate, subparallel, moderately robust; posteromedial angle of each elytron without reddish spot, sometimes with paler, yellowish-brown area (Figs. 10.54a–10.57a).....3 3. Pronotum distinctly transverse, sides distinctly converging apically (Fig. 10.57a); pronotal microsculpture weak or absent (Fig. 10.57b); spaces betweeen setae smooth and highly glossy (Fig. 10.57a); genitalia as illustrated (Fig. 10.57c, d, i) ....................................................................... ....Aleochara (Xenochara) fumata Gravenhorst – Pronotum moderately transverse, with sides moderately converging apically (Figs. 10.54a– 10.56a); pronotal microsculpture strong, spaces between setae covered with microsculpture and appearing less glossy (Figs. 10.54a, 10.55a, b, and 10.56a); genitalia of different shape (Figs. 10.54d, e, 10.55c, d, i, and 10.56b, c, d, i, j) ................................................................... 4 4. Median lobe of aedeagus strongly sinuate ventrally in lateral view (Fig. 10.56b, c); spermatheca with large, evenly spherical capsule amd large apical invagination, posterior stem strongly twisted posteriorly (Fig. 10.56i, j) ....................... Aleochara (Xenochara) quadrata Klimaszewski – Median lobe of aedeagus broadly arched ventrally in lateral view (Fig. 10.55c); spermatheca with less evenly spherical capsule, apical invagination smaller, stem not twisted posteriorly (Figs. 10.54d, e and 10.55i).............5 5. Antecostal suture of female sternite VIII slightly sinuate and not pointed medially (Fig. 10.55h);

10

Tribe Aleocharini Fleming, 1821

spermatheca with apical invagination of capsule broad, stem with chamber connected to duct at 90 in vertical plane (Fig. 10.55i).......................... ................Aleochara (Xenochara) castaneipennis Mannerheim – Antecostal suture of female sternite VIII sharply pointed medially (Fig. 10.54c); spermatheca with apical invagination of capsule narrow, stem with chamber connected to duct at 90 in horizontal plane (Fig. 10.54d, e) ........................................... ..................... Aleochara (Xenochara) carmanah Klimaszewski 21. Aleochara (Xenochara) Klimaszewski (Fig. 10.54a–e)

carmanah

Aleochara (Xenochara) carmanah Klimaszewski, in Klimaszewski and Winchester 2002. Klimaszewski et al. 2017. Diagnosis (based on female). Body narrowly elongate, subparallel, moderately robust, length 3.8 mm; colour dark brown, with paler elytra, posterior margins of tergites and legs, strongly glossy (Fig. 10.54a); forebody with coarse meshed microsculpture, sculpticells flat, punctation on pronotum strong and coarse, antennomeres V–X distinctly transverse (Fig. 10.54a). MALE. Unknown. FEMALE. Tergite VIII with apical margin truncate, shallowly emarginate medially (Fig. 10.54b); sternite VIII with apical margin rounded, with short pubescence, antecostal suture sharply pointed medially (Fig. 10.54c); spermatheca with apical invagination of capsule narrow, stem with chamber connected to duct at 90 in horizontal plain (Fig. 10.54d, e). This species can be misidentified as the externally similar sibling species A. castaneipennis, but in A. carmanah the body is generally broader and more evenly coloured, and the capsule of the spermatheca is larger and more evenly spherical. It can also be confused with A. fumata from which it differs in having coarser pronotal punctation, strong microsculpture, a less glossy pronotum, often a larger body, and different shape of spermatheca.

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Tribe Aleocharini Fleming, 1821

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: transition zone in Sitka spruce forest. Collecting period: IX. Collecting method: pitfall trap. 22. Aleochara (Xenochara) castaneipennis Mannerheim (Fig. 10.55a–i) Aleochara castaneipennis Mannerheim, 1843. Klimaszewski 1984, Klimaszewski and Peck 1986, Gouix and Klimaszewski 2007, Klimaszewski et al. 2005, 2011, 2017, 2018, Majka and Klimaszewski 2010, Majka et al. 2011. Diagnosis. Body subparallel, length 2.8–6.0 mm; colour brown to dark brown, with tarsi and palps rusty-brown, elytra often paler than remainder of the body (Fig. 10.55a); forebody with coarse meshed microsculpture, sculpticells flat (Fig. 10.55b); antennomeres V–X distinctly transverse (Fig. 10.55a). MALE. Tergite VIII with apical margin truncate and slightly emarginate (Fig. 10.55e); sternite VIII triangularly produced apically (Fig. 10.55f); median lobe of aedeagus in lateral view with ventral margin of tubus evenly arcuate (Fig. 10.55c), apical sclerites of internal sac directed anteriad (Fig. 10.55c), in dorsal view as illustrated (Fig. 10.55d). FEMALE. Tergite VIII with apical margin truncate, shallowly emarginate medially (Fig. 10.55g); sternite VIII with apical margin rounded, with short pubescence (Fig. 10.55h); spermatheca L-shaped, capsule slightly elongate sac-shaped (Fig. 10.55i). This species is often misidentified as the externally similar A. fumata but in A. castaneipennis the body is often larger, the pronotal punctation slightly coarser with the interstices between punctures less glossy, and the genital features are different. It also may be confused with A. carmanah, from which it differs in having the antecostal suture of female sternite VIII feebly sinuate (sharply pointed medially in A. carmanah), and spermatheca with apical

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invagination of capsule broad, stem with chamber connected to duct at 90 in vertical plane in most specimens examined. DISTRIBUTION (verified localities). Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, NT, ON, QC, YT. USA: AK, AR, AZ, CO, WA, WY. Based on a re-examination of the material originally examined by Klimaszewski (1984) and new material available to us, only the above records are confirmed here. Many of the state records from the eastern U.S. actually refer to A. castaneimarmotae. All specimens from Oregon and the available males from California listed by Klimaszewski (1984) refer to A. quadrata. All 134 specimens re-examined from Alaska (UAM), and identified by one of us (JK) as A. castaneipennis, belong to this species. COLLECTION AND HABITAT DATA. Habitat: coniferous, deciduous and mixed wood forest types, coastal limestone barrens. Adults often found in rotting mushrooms, carrion, forest leaf litter, moss, and human feces. In NB, this species was often abundant in decaying mushrooms but was seldom found in other microhabitats. Collecting period: IV–XI. Collecting method: sifting decaying mushrooms, forest litter, using unbaited and carrion-baited pitfall traps, flight intercept traps. 23. Aleochara (Xenochara) Klimaszewski (Fig. 10.56a–j)

quadrata

Aleochara quadrata Klimaszewski, 1984. Klimaszewski et al. 2017. Diagnosis. Body subparallel, length 3.8–6.5 mm; colour dark brown, with reddish sutural part of elytra, and posterior margins of tergites and legs (Fig. 10.56a); forebody with microsculpture not apparent or weakly defined; antennomeres V–X distinctly transverse (Fig. 10.56a). MALE. Tergite VIII with apical margin truncate and sinuate laterally (Fig. 10.56e); sternite VIII triangularly produced apically (Fig. 10.56f); median lobe of aedeagus with apical half of tubus sinuate in lateral

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view, apex of tubus bent ventrad, subapical sclerites of internal sac of about equal width throughout its length and with apices not upturned (Fig. 10.56b, c). FEMALE. Tergite VIII with apical margin truncate, shallowly emarginate medially and sinuate laterally (Fig. 10.56g); sternite VIII with apical margin rounded, with short pubescence, antecostal suture sinuate, slightly pointed medially (Fig. 10.56h); spermatheca with large, evenly spherical capsule and broad and deep apical invagination, posterior stem strongly twisted (Fig. 10.56i, j). Generally, A. quadrata is much darker and the pronotum is glossier than in A. castaneipennis. Males of A. quadrata can be reliably distinguished from A. castaneipennis by the sinuate apical half of the tubus and subapical sclerites of the internal sac, which are not upturned. Aleochara quadrata is extremely similar to the Palaearctic species A. moerens Gyllenhal but can be distinguished by its subapical sclerite which does not gradually narrow to a point in A. moerens. The latter species is not yet known from the Nearctic region (Klimaszewski et al. 2017). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, NV. MEXICO: locality unspecified (Klimaszewski 1984). COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. 24. Aleochara (Xenochara) fumata Gravenhorst{ (Fig. 10.57a–i) Aleochara fumata Gravenhorst, 1802. Klimaszewski 1984, Klimaszewski and Peck 1986, Klimaszewski and Génier 1987, Gouix and Klimaszewski 2007, Klimaszewski et al. 2005, 2011, Majka and Klimaszewski 2010. Diagnosis. Body subparallel, length 2.8–5.0 mm; colour brown to dark brown, with basal 1–3 antennomeres, legs/tarsi and palps rusty-brown

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Tribe Aleocharini Fleming, 1821

(Fig. 10.57a); forebody with meshed microsculpture fine or absent (Fig. 10.57b); antennomeres V–X distinctly transverse (Fig. 10.57a). MALE. Tergite VIII with apical margin minutely crenulate, truncate, slightly broadly emarginate (Fig. 10.57e); sternite VIII slightly triangularly produced apically (Fig. 10.57f); median lobe of aedeagus in lateral view with ventral margin of tubus evenly arcuate (Fig. 10.57c), apical sclerites of internal sac directed ventrad (Fig. 10.57c), in dorsal view median lobe as illustrated (Fig. 10.57d). FEMALE. Tergite VIII arcuate apically (Fig. 10.57g); sternite VIII rounded apically (Fig. 10.57h); spermatheca L-shaped, capsule club-shaped and angular, stem short and straight (Fig. 10.57i). This species is externally very similar to A. castaneipennis and A. castaneimarmotae (reported from eastern Canada), but in A. fumata the body is often smaller, the pronotal punctation finer, the microsculpture absent or fine, the elytra shorter, the pronotum highly glossy, and the genital features are different. DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, MB, NB, NF, NS, ON, QC, PE, YT. USA: AL, AR, AZ, CA, GA, MA, MD, ME, MN, MO, NC, NH, NJ, NM, NY, OK, OR, PA, TN, VT, WA, WI, WV, WY. COLLECTION AND HABITAT DATA. Habitat: hardwood, mixed, and coniferous forests. In rotting fungi (often very common), organic debris, carrion, and wet moss. Collecting period: IV–XI. Collecting method: sifting mushrooms and forest debris, pitfall traps, and flight intercept traps. 25. Aleochara (Xenochara) lacertina Sharp (Fig. 10.58a–h) Aleochara lacertina Sharp, 1883. Klimaszewski 1984, Downie and Arnett 1996, Gouix and Klimaszewski 2007, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2016a.

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Tribe Aleocharini Fleming, 1821

113

Diagnosis. Body subparallel, length 3.0–6.0 mm; colour piceous to black, with tarsi and palps rustybrown, elytra unicolourous or with two yellow spots in apico-median part of disc; abdomen with imbricate microsculpture (Fig. 10.58a); antennomeres V–X subquadrate to slightly transverse (Fig. 10.58a). MALE. Tergite VIII with apical margin truncate, crenulate (Fig. 10.58d); sternite VIII rounded apically (Fig. 10.58e); median lobe of aedeagus with ventral margin of tubus broadly arcuate in lateral view (Fig. 10.58b); apical sclerites of internal sac as illustrated (Fig. 10.58b, c). FEMALE. Tergite VIII with apical margin truncate, slightly emarginate (Fig. 10.58f); sternite VIII rounded apically (Fig. 10.58g); spermatheca very small, shape as illustrated (Fig. 10.58h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NF, NS, ON, QC, SK. USA: AZ, CA, CO, FL, MA, ME, MN, MT, ND, NH, NM, NV, NY, OK, OR, PA, RI, TN, TX, UT, WA, WI. MEXICO: Tabasco. COLLECTION AND HABITAT DATA. Habitat: often in organic substrates such as animal droppings including horse, cow, deer, jay, human feces, as well as in carrion, fungi, decaying seaweed and sea wrack along sea beaches and salt marshes, and in flood debris along shores of stream and lakes, and litter in marshy areas. Some were collected from funnel extracts of water hyacith plants. Six adults emerged in laboratory from puparia of Lispe uliginosa (Fallen) (Muscidae), collected under stones. Collecting period: IV–XI. Collecting method: sifting leaf litter, sea wrack, seaweed, rotting wood with primitive mushrooms, Malaise trap, UV trap. 26. Aleochara (Xenochara) Gravenhorst{ (Fig. 10.59a–h)

lanuginosa

Aleochara (Xenochara) lanuginosa Gravenhorst, 1802. Klimaszewski 1984, Cervenka and Moon

1991, Klimaszewski and Frank 1992, Gouix and Klimaszewski 2007, Webster et al. 2009, Majka and Klimaszewski 2010, Klimaszewski et al. 2011. Diagnosis. Body broad, oval, length 3.0–7.0 mm; colour piceous to black, with tarsi and palpi rustybrown, elytra black or with posterior margin pale reddish (Fig. 10.59a); antennomeres V–X slightly transverse (Fig. 10.59a). MALE. Tergite VIII with apical margin serrate, broadly emarginate medially, rounded laterally (Fig. 10.59d); sternite VIII slightly triangularly produced apically (Fig. 10.59e); median lobe of aedeagus with ventral margin of tubus in lateral view with angular elevation, apex thin, elongate (Fig. 10.59b); median lobe in dorsal view as illustrated (Fig. 10.59c); apical sclerites of internal sac apically broadly falciform (Fig. 10.59b, c). FEMALE. Tergite VIII with apical margin arcuate, slightly emarginate (Fig. 10.59f); sternite VIII with apical margin slightly produced, rounded at apex (Fig. 10.59g); spermatheca as illustrated, similar to that of A. inexspectata (Fig. 10.59h). This species is externally very similar to A. inexspectata, but in A. lanuginosa the pronotum is narrower, with the margins more strongly converging apically, the apical margin of male tergite VIII is serrate, and the internal sac of the aedeagus has sclerites which are broad and apically falciform (narrow apically in A. inexspectata). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, NB, NF, NS, ON, QC. USA: CA, CO, ID, MA, MN, NC, NH, NJ, NY, OR, PA, RI, WA, WI. COLLECTION AND HABITAT DATA. Habitat: woodlands and open habitats including beaches and pastures. Usually in animal droppings, including cow and horse dung, human feces, carrion including dead Limulus sp. on a sea beach, oak duff, and compost (decaying vegetables). Collecting period: V–VIII or II–XI in southern localities. Collecting method: sifting, berlese litter, dung-baited pitfall traps.

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Fig. 10.35 (a–h) Schematic representaion of life cycle in the genus Aleochara: (a) adult female; (b) egg; (c) first instar larva of Aleochara is actively searching for fly puparium; (d) second instar larva of Aleochara remains

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Tribe Aleocharini Fleming, 1821

in fly puparium; (e) third instar larva of Aleochara leaving fly puparium and pupates in soil; (f) pupa, (h) fly adult and egg; (g) maggott and fly pupa. After Klimaszewski and Jansen (1993)

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Tribe Aleocharini Fleming, 1821

Fig. 10.36 (a–j) Representatives of subgenera of the genus Aleochara with important diagnostic features indicated by arrows (Modified from Klimaszewski and Jansen 1993): (a) habitus of Xenochara; (b) almost completely or completely carinate mesosternum of Xenochara; (c) habitus of Coprochara; (d) completely

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carinated mesosternum of Coprochara; (e) habitus of Aleochara; (f) not carinate mesosternum of Aleochara; (g) habitus of Emplenota; (h) not carinate mesosternum of Emplenota; (i) habitus of Calochara; (j) partialy carinate mesosternum of Calochara

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Fig. 10.37 (a–h) Images of Aleochara (Aleochara) assiniboin Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

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Tribe Aleocharini Fleming, 1821

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.38 (a–h) Images of Aleochara (Aleochara) curtula (Goeze): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c, h) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

118

Fig. 10.39 (a–h) Images of Aleochara (Aleochara) gracilicornis Bernhauer: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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Tribe Aleocharini Fleming, 1821

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.40 (a–h) Images of Aleochara (Aleochara) tahoensis Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 10.41 (a–h) Images of Aleochara (Calochara) laramiensis (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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Tribe Aleocharini Fleming, 1821

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.42 (a–h) Images of Aleochara (Calochara) rubricalis (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c, h) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

122

Fig. 10.43 (a–h) Images of Aleochara (Calochara) villosa Mannerheim: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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Tribe Aleocharini Fleming, 1821

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.44 (a–h) Images of Aleochara (Coprochara) bilineata Gyllenhal: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 10.45 (a–h) Images of Aleochara (Coprochara) bimaculata Gravenhorst: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

10

Tribe Aleocharini Fleming, 1821

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.46 (a–h) Images of Aleochara (Coprochara) suffusa (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

125

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

126

Fig. 10.47 (a–h) Images of Aleochara (Coprochara) sulcicollis Mannerheim: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male

10

Tribe Aleocharini Fleming, 1821

sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c, d) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.48 (a–h) Images of Aleochara (Coprochara) verna Say: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

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male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 10.49 (a–j) Images of Aleochara (Emplenota) curtidens Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal

10

Tribe Aleocharini Fleming, 1821

view; (e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i, j) spermatheca. (c, d, i) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.50 (a–h) Images of Aleochara (Emplenota) litoralis (Mäklin): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

130

Fig. 10.51 (a–h) Images of Aleochara (Emplenota) pacifica (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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Tribe Aleocharini Fleming, 1821

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h), spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.52 (a–d) Images of Aleochara (Tinotus) acerba (Casey) [lectotype]: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in

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lateral view; (c) male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 10.53 (a–g) Images of Aleochara (Tinotus) morion Gravenhorst: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral

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Tribe Aleocharini Fleming, 1821

view; (c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.54 (a–e) Images of Aleochara (Xenochara) carmanah Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) female tergite VIII; (c)

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female sternite VIII; (d, e) spermatheca. (e) modified from Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 10.55 (a–i) Images of Aleochara (Xenochara) castaneipennis Mannerheim: (a) habitus in dorsal view; (b) magnified fragment of pronotum; (c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in

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Tribe Aleocharini Fleming, 1821

dorsal view; (e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i), spermatheca. (a–i) after Klimaszewski et al. (2017). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.56 (a–j) Images of Aleochara (Xenochara) quadrata Sharp: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male

135

sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i, j) spermatheca. (c, j) after Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

136

Fig. 10.57 (a–i) Images of Aleochara (Xenochara) fumata Gravenhorst: (a) habitus in dorsal view; (b) magnified fragment of pronotum; (c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in

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Tribe Aleocharini Fleming, 1821

dorsal view; (e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (a–i), after Klimaszewski et al. (2017). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

Fig. 10.58 (a–h) Images of Aleochara (Xenochara) lacertina Sharp: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

138

Fig. 10.59 (a–h) Images of Aleochara (Xenochara) lanuginosa Gravenhorst: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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Tribe Aleocharini Fleming, 1821

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) modified from Klimaszewski (1984). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Aleocharini Fleming, 1821

References Bernhauer M (1901) Neue exotische Arten der Gattung Aleochara Gravenhorst. Steittiner Entomologishe Zeitschrift 62(3–4):366–373 Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013). Checklist of beetles (Coleoptera) of Canada and Alaska, 2nd edn. Series Faunistica 109. Pensoft Publishers, Sofia-Moscow, 402 pp Casey TL (1884) Contributions to the descriptive and systematic Coleopterology of North America. Collins Printing House, Philadelphia. Part I, pp 1–60, pl. 1; Part II, pp 61–124 Casey TL (1893) Coleopterological notices. V. Ann N Y Acad Sci 7 (1892–1894):281–606 Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St Louis 16(6):125–434 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, Pennsylvania. 259 pp Cervenka VJ, Moon RD (1991) Arthropods associated with fresh cattle dung pats in Minnesota. J Kansas Entomol Soc 64(2):131–145 Downie NM, Arnett RH (1996) Introduction; suborders Archostemata, Adephaga, and Polyphaga, thru superfamily Cantharoidea. The beetles of northeastern North America, vol. I, The Sandhill Crane Press, Gainesville, FL, 880 pp Fauvel CAA (1889) Liste des coléoptères communes a l’Europe et a l’Amérique du Nord. D’après le catalogue de M.J. Hamilton. Avec rémarques et additions. Revue d’Ent 8(3):92–174 Fleming J (1821) Insecta [pp 41–56]. In: Supplement to the fourth, fifth and sixth editions of the Encyclopaedia Britannica, with preliminary dissertations on the history of sciences. vol. 5. A. Constable and Company, Edinburgh, 584 pp Fletcher J (1902) Report of the entomologist and botanist. Division of Entomology. Exp Farm Rep 16 (1901):197–259 Gibson A, Treherne RC (1916) The cabbage root maggot and its control in Canada with notes on the imported onion maggot and the seed corn maggot. Dominion of Canada Department of Agriculture Entomological Branch. Bulletin 12:1–58 Goeze JAE (1777) Entomologische Beyträge zu des Ritter Linné zwölften Ausgabe des Natursystems. Erster Theil Weidmanns, Erben & Reich, Leipzig. xvi + 736 pp Gouix N, Klimaszewski J (2007) Catalogue of aleocharine rove beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae). Pensoft Publishers, Sofia-Moscow. 165 pp Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in

139 genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353 (1):1–134 Gyllenhal L (1810) Insecta Svecica. Classis I: Coleoptera sive Eleuterata. Tomi I pars II. F.J. Leverentz, Scaris. xix + [1] + 660 pp Hatch MS (1957) The Beetles of the Pacific Northwest. Part II: Staphyliniformia. University of Washington Publications in Biology 16: IX + 384 pp Hemachandra KS, Holliday NJ, Klimaszewski J, Mason PG, Kuhlmann U (2005) Erroneous records of Aleochara bipustulata from North America: an assessment of the evidence. Can Entomol 137(2):182–187 Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Memoirs of the Entomological Society of Canada No. 129, [2] + 211 pp Klimaszewski J (1985) Nomenclatoral changes in the genus Aleochara Gravenhorst (Coleoptera: Staphylinidae: Aleocharinae). Coleopt Bull 39(4):376 Klimaszewski J, Cervenka VJ (1986) A revision of the genus Aleochara (Coleoptera: Staphylinidae) of America north of Mexico (Supplement 3). New distribution data. Entomol News 97(3):119–120 Klimaszewski J, Frank JH (1992) New distributional data for Nearctic Myllaena Erichson (Coleoptera: Staphylinidae). Supplement 2. Coleopt Bull 46 (4):397–402 Klimaszewski J, Génier F (1987) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Supplement 4. New distribution data and description of two new species. Coleopt Bull 41(3):241–248 Klimaszewski J, Jansen R (1993) Systematics, biology and distribution of Aleochara Gravenhorst from southern Africa. Part I: subgenus Xenochara Mulsant and Rey (Coleoptera: Staphylinidae). Ann Transvaal Mus 36 (7):53–107 Klimaszewski J, Peck SB (1986) A review of the cavernicolous Staphylinidae (Coleoptera) of eastern North America: Part I. Aleocharinae. Quaestiones Entomol 22(2):51–113 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc belge d’Entomol 40:3–126 Klimaszewski J, Pelletier G, Sweeney J (2002) Genus Tinotus (Coleoptera: Staphylinidae, Aleocharinae) from America north of Mexico: review of the types, distribution records, and key to species. Can Entomol 134(3):281–298 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48

140 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013) Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia-Moscow, 360 pp Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Langor DW, Bourdon C, Gilbert A, Labrecque M (2016a) Two new species and new provincial records of aleocharine rove beetles from Newfoundland and Labrador, Canada (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 593:49–89 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016b) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Brunke A (2017) A new criptic species of Aleochara associated with Marmota monax (L.) burrows in North America (Coleoptera: Staphylinidae, Aleocharinae). Insecta Mundi 0600: 1–11 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. 902 pp Majka CG, Klimaszewski J (2008) Introduced Staphylinidae (Coleoptera) in the Maritime Provinces of Canada. Can Entomol 140(1):48–72 Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39 Majka CG, Chandler DS, Donahue CP (2011) Checklist of the beetles of Maine, USA. Empty Mirrors Press, Halifax, Nova Scotia. 328 pp Mäklin FW (1853) New species and notes. In: Mannerheim CG: Dritter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soci Imp Natural Moscou 26(3):95–273 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp [separate, re-issued

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Tribe Aleocharini Fleming, 1821

in Mémoires Présentées à l’Académie Impériale des Sciences de St.-Pétersbourg 1:415–501, in 1831] Mannerheim CG (1843) Beitrag zur Kaefer-fauna der Aleutischen Inseln, der Insel Sitkha und Neu-Californiens. Bull Soci Imp des Natural Moscou 16(2):175–314 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). Div Agrc Sci Univ of Calif Special Publ 3015:1–514 Mulsant E, Rey C (1874) [Histoire naturelle des Coléoptères de France]. Tribu des brévipennes: Famille des aléochariens (suite): Sixième branche: Aléocharaires. Annales de la Société Linnéenne de Lyon (N.S.) 20(1873):285–447 Osswald J, Bachmann L, Gusarov VI (2013) Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae). Syst Entomol 38 (3):507–522 Say T (1833) Descriptions of new species of North American insects and observations on some already described. T. Say, New Harmony, 81 pp Schülke M, Smetana A (2015) Family Staphylinidae Latreille, 1802. In: Löbl I, Löbl D (eds) Catalogue of Palearctic Coleoptera, vol. 2, Revised and updated edition. Hydrophiloidea—Staphylinoidea. Brill, Leiden and Boston, pp 304–1134 Sharp D (1883) Fam. Staphylinidae [part]. In: Godman, FD, Salvin O (eds) Biologia Centrali-Americana. Insecta. Coleoptera. vol. 1, Part 2. Taylor & Francis, London, pp 145–312, pls. 5–7 Sikes D (2004) The Beetle Fauna of Rhode Island: an annotated checklist. Volume 3 of The Biota of Rhode Island. Rhode Island Natural History Survey, Kingston, RI, vi + 296 pp Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:171–248 Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016) Further contributions to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including descriptions of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J. (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216 Yamamoto S, Maruyama M (2016) Revision of the subgenus Thinotus Sharp, stat.n., of the parasitoid rovebeetle genus Aleochara Gravenhorst (Coleoptera, Staphylinidae, Aleocharinae) from Japan, Taiwan, and the Russian Far Est. ZooKeys 559: 81–106

Tribe Oxypodini C.G. Thomson, 1859

Tribe Oxypodini C.G. Thomson, 1859 (Figs. 11.60–11.104) (Seevers 1978; Ashe 2001; Klimaszewski et al. 2006, 2018; Osswald et al. 2013) Diagnosis. Body narrowly elongate or ovoid, body segments loosely articulated; head without distinct neck in most species; frontal suture of head absent or present (Oxypoda); mouthparts generalized (similarly developed across the tribe and without distinctive features for the tribe); pronotal hypomera not visible in lateral view in many (e.g., Devia, Oxypoda); mesocoxae narrowly separated in most, moderately so in few; process of mesoventrite slender in most; abdominal tergite IX narrowly subdivided at base; tarsal formula in most taxa 5-5-5, in some 4-5-5 or 4-44. Highly heterogenous assemblage, difficult to characterize uniformly. Key to Genera of Oxypodini 1. Head with distinct broad neck [sometimes not visble in dry specimens when head concealed by pronotum] (Figs. 11.60a, 11.63a, and 11.101a– 11.104a) .............................................................2 – Head without distinct neck (exception Ocalea but body differently shaped) ..............................4 2. Head in dorsal view with lateral margins very angular posteriorly, body subparallel, all pronotal pubescence directed straight posteriad (Figs.

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11.101a–11.104a); often found under bark of dead trees ..........................Phloeopora Erichson – Head in dorsal view with lateral margins rounded posteriorly, body and pronotal pubescence pattern not as above (Figs. 11.60a and 11.63a); found in different habitats....................3 3. Body slender, somewhat ant-shaped, with abdomen clavate, distinctly constricted at base and broadest at about middle, abdominal base significantly narrower than width of elytra (Fig. 11.60a); basal abdominal tergites deeply, transversely impressed at base, impressions coarsely punctate, interstices between punctures forming longitudinal ridges (Fig. 11.60a)............ ....................................... Blepharhymenus Solier – Body robust, not ant-shaped, with abdomen subparallel and tapering apically, not constricted at base and broadest at base, abdominal base slightly narrower than elytra (Fig. 11.63a); basal abdominal tergites deeply, transversely impressed at base, impressions without longitudinal ridges (Fig. 11.63a)...................Calodera Mannerheim 4. Pronotal hypomera not visible in lateral view ............................................................................5 – Pronotal hypomera visible in lateral view.....6 5. Body robust, narrowly oval with abdomen subparallel and not strongly tapering apically (Fig. 11.64a); pronotum subelliptical, without apparent hind angles (Fig. 11.64a); antennomere

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_11

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III distinctly longer than II; each mandible with a subapical tooth ..................... Devia Blackwelder – Body slender, subparallel or ovate with abdomen tapering apically (Figs. 11.69a–11.87a); pronotum often horseshoe-shaped with distinct posterior angles (Figs. 11.69a–11.87a); antennomere III as long as or shorter than II; mandibles without subapical tooth, right mandible may have basal median tooth......................... ........................................ Oxypoda Mannerheim 6. Mandibles extremely long and sickle-shaped, apices very slender, each crossing the other in resting position, each mandible bearing a long spine or tooth; infraorbital carina strong and complete; ligula more-or-less deeply bilobed; terminal palpomere needle-shaped; frontal suture of head absent; body as illustrated (Fig. 11.65a) .............. ................................................. Gnathusa Fenyes – Mandibles short to moderately long and shaped differently, not crossing each other in resting position, other characters different.........7 7. Body compact and robust (Aleochara like), subparallel, convex (Figs. 11.62a, 11.67a, and 11.68a); integument of forebody with meshed microsculpture....................................................8 – Body not as above; integument of forebody with or without meshed microsculpture........... 10 8. Pronotum distinctly transverse, about 1.5 times as wide as long (Fig. 11.62a); posterolateral margin of elytra with strong sinuate emargination (Fig. 11.62a); right mandible with strong median tooth, left mandible with obtuse subapical one; ligula rounded, not bifid; found in plant and animal refuse in stables, granaries, in bird and occasionally ant nests ....... Crataraea C.G. Thomson – Pronotum moderately transverse, about 1.3 times as wide as long (Figs. 11.67a and 11.68a); posterolateral margin of elytra with emargination scarcely noticeable (Figs. 11.67a and 11.68a); each mandible with median tooth; ligula bifid apically; found in riparian or forest habitats ......9 9. All pronotal setae distributed along midline of disc directed posteriad (Fig. 11.68a); frontoclypeal phragma present and visible externally

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Tribe Oxypodini C.G. Thomson, 1859

as frontal suture between antennal pits (seen better in diffused light); spermatheca S-shaped with spherical capsule bearing deep and broad invagination, stem sinuate and moderately broad (Fig. 11.68g); median lobe of aedeagus with tubus markedly produced ventrally in lateral view (Fig. 11.68b); Canadian species are known from wetland habitats..................Ocyusa Kraatz – Pronotal setae along midline of disc directed anteriad over about apical one-third of midline and posteriad elsewhere (Fig. 11.67a); frontal suture absent; spermatheca capsule spherical or tubular and narrowly extended forming L-shaped neck, connected to thin and elongate stem (Fig. 11.67g); median lobe of aedeagus with tubus almost straight in lateral view (except basally) and slightly produced ventrally (Fig. 11.67b); Canadian species are known from forest habitats ...........Mniusa Mulsant and Rey. 10. Antennomeres VII–X each up to three times as wide as long (Fig. 11.61a); body length 1.0–2.0 mm; elytra at suture about as long as pronotum along midline; apical lobe of parameres strongly sclerotized and appearing black or dark brown; spermatheca U-shaped or sinuate, appearing as a simple tube, capsule without apical invagination (Fig. 11.61g, h) ............................... .................................. Meotica Mulsant and Rey – Antennomeres VII–X each elongate or subquadrate to slightly transverse; body length 2.0–8.0 mm; elytra at suture slightly longer than pronotum along midline; apical lobe of parameres not strongly sclerotized; spermatheca shaped differently [Ocalea group of genera]....................11 11. Integument of forebody moderately glossy (Figs. 11.89a, 11.90a, 11.91a, 11.93a–11.98a, and 11.100a) and with strong isodiametric microsculpture consisting of convex sculpticells; punctation moderately dense and coarse and usually separated by a distance of two diameters of a puncture; body medium sized to larger (length 4.5–6.0mm) ......................................................12 – Integument of forebody strongly glossy (Figs. 11.88a, 11.92a, and 11.99a) and with weak, scarcely visible microsculpture consisting of usually flat sculpticells; punctation fine and

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Tribe Oxypodini C.G. Thomson, 1859

sparse and usually separated by a distance of two to three diameters of a puncture; body small to medium sized (length 3.0–4.4 mm) .................16 12. Basal tarsomere of metatarsus as long as the three following tarsomeres combined (Fig. 11.90a); antennomere IV elongate, about twice as long as wide (Fig. 11.90a); ligula entire and not divided apically; basal palpomere of labial palpus swollen and contrasting with the remaining palpomeres; mesoventrite broadly triangular with very short and sharply pointed mesoventral process, which is separated from the metaventral process by a space of about one-third of diameter of a mesocoxa; body large, length 5.5–6.0 mm ............................................... ....................................Megocalea Klimaszewski – Basal tarsomere of metatarsus approximately as long as the two following tarsomeres combined, or slightly longer but distinctly shorter than the three following tarsomeres combined; antennomere IV usually shorter than twice maximum width; ligula deeply bifid or slightly emarginated apically; basal palpomere of labial palpus only slightly broader than the remaining ones; mesoventrite usually narrowly triangular with narrowly elongate and apically sharply pointed mesoventral process, which is often separated from the metaventral process by a distance of less than one-third of diameter of mesocoxa; body medium sized to large, length 2.8–6.0 mm, average 4.5–5.0 mm ...................13 13. Elytra as broad as pronotum at base, at suture at most as long as pronotum, (Fig. 11.91a) ......... .....................................Metocalea Klimaszewski – Elytra broader that pronotum at base, at suture longer than pronotum (Figs. 11.89a, 11.93a– 11.98a, and 11.100a)........................................ 14 14. Antennomeres VII–X strongly transverse; body narrowly elongate, head and pronotum only slightly narrower than elytra (Fig. 11.100a) .............................................................................. .......................................... Parocalea Bernhauer – Antennomeres VII–X subquadrate to slightly transverse; body broader and less evenly subparallel (Figs. 11.89a and 11.93a–11.98a)............ 15

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15. Antennomeres VII–X subquadrate (Fig. 11.89a); mesoventrite partially carinate, carina about as long as half of mesoventral length; mesoventral process narrow and sharply pointed, wedging between mesocoxae and as long as three-fourths of mesocoxal length; median lobe of aedeagus with short projection in front of crista apicalis of bulbus (Fig. 11.89b); spermatheca with small capsule bearing deep invagination and slender S-shaped stem (Fig. 11.89h)............. ..................................... Betocalea Klimaszewski – Antennomeres VII–X slightly transverse (Figs. 11.93a–11.98a); mesoventrite fully, or partially carinate and reaching about one-third of mesoventrite length, mesoventral process long, narrow, and acutely pointed; other characters different or in different combination ........................ ...................................Neothetalia Klimaszewski 16. Pronotum as wide and as long as elytra along suture (Fig. 11.88a) ..... Alfocalea Klimaszewski – Pronotum narrower than elytra and shorter than elytra along suture (Figs. 11.92a and 11.99a) .............................................................17 17. Antennomeres VII–X elongate (Fig. 11.99a); pronotum distinctly narrower than elytra (Fig. 11.99a); median lobe of aedeagus and spermathecal as illustrated (Fig. 11.99b, c, h) ..... .................................................. Ocalea Erichson – Antennomeres VII–X transverse (Fig. 11.92a); mesoventral basal carina very short, about one-fifth of the mesoventrum; median lobe of aedeagus and spermathecal not as above (Fig. 11.92b, c, h)..............Neoisoglossa (Casey)

Genus Blepharhymenus Solier, 1849 (Fig. 11.60a–g) (Casey 1893; Klimaszewski and Peck 1986) Diagnosis. Body shape distinctive, head rounded posteriorly with distinct neck, head and pronotum subequal in width, pronotum about 3/5 as wide as elytra; abdomen swollen dorsoventrally and laterally in apical half, at fifth segment almost as wide as elytra; first three visible abdominal

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tergites with deep, coarsely punctate transverse impressions, intervals between punctures ridgelike, prominent median carina present; mesoventrite long with basal carina extending up to one-third of its length, process of mesoventrite V-shaped and pointed apically; tarsal formula 5-5-5. 27. Blepharhymenus illectus (Casey) (Fig. 11.60a–g) Echidnoglossa illecta Casey, 1911. Syntypes: USA: Oregon, Portland, Echinoglossa illecta Casey, Type USNM 39709; Casey bequest 1925 (USNM) 1 male, 1 female. Examined. Diagnosis. Body length 3.0–3.5 mm, colour dark brown with sutural part of elytra and appendages partially reddish, integument glossy, body shape distinctive (see generic description) (Fig. 11.60a). MALE. Tergite VIII with apical margin truncate, antecostal suture distant from basal margin (Fig. 11.60c); sternite VIII triangularly produced apically (Fig. 11.60d); tubus of median lobe of aedeagus in lateral view with ventral margin broadly arcuate, with apex broad, lancet-shaped (Fig. 11.60b); internal sac sclerites complex as illustrated (Fig. 11.60b). FEMALE. Tergite VIII with apical margin arcuate, antecostal suture distant from basal margin (Fig. 11.60e); sternite VIII triangularly produced apically (Fig. 11.60f); spermatheca with capsule small, spherical, stem thin, long, sinuate, and looped and twisted posteriad (Fig. 11.60g). Females of this species can be distinguished from those of externally similar B. brendeli Casey by the shape of spermatheca and triangularly produced apically sternite VIII, which is rounded and slightly emarginate medially in B. brendeli.

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Tribe Oxypodini C.G. Thomson, 1859

Columbia, Vancouver Island, Goldstream Park, 27.V.1968, B), A. Smetana (CNC) 4 males, 3 females, 2 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: associated with riparian habitats in forests. Collecting period: V. Collecting method: sifting leaf litter and moss and other forest litter. Subtribe Meoticina Seevers, 1978 (Seevers 1978; Ashe 2001) Diagnosis. Body slender, dorsoventrally compressed, sides subparallel in most species; head with neck about one-third as wide as head, without frontal suture; terminal antennomere with pair of coeloconic sensillae; mesocoxae narrowly separated; process of mesoventrite slender and acutely pointed, not extended beyond middle of coxae; process of metaventrite very short, usually almost absent; mesocoxal acetabula not or weakly margined in most species; pronotal hypomera fully visible in lateral view; first two or three visible abdominal tergites with transverse impressions; tarsal claws scythe-shaped (falcate) [except in Meotica]; tarsal formula 4-4-4 or 5-5-5.

Genus Meotica Mulsant and Rey, 1873 (Fig. 11.61a–h) (Klimaszewski et al. 2007, 2018)

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific provincial record]. USA: OR.

Diagnosis. Body slender, parallel-sided, length 1.0–2.0 mm, colour light reddish-brown; antennomeres VI–X distinctly transverse, each up to three times as wide as long; eyes small, about one-third length of genae in dorsal view; pronotal hypomera visible in lateral view; mesocoxal cavities not margined or weakly so; process of mesoventrite short and acute; process of metaventrite short and poorly delimited; paramere with apical lobe small and heavily sclerotized; spermatheca U-shaped or L-shaped, forming simple tube.

We record this genus and species from BC for the first time based on the following: British

28. Meotica pallens (Redtenbacher){ (Fig. 11.61a–h)

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Tribe Oxypodini C.G. Thomson, 1859

Homalota pallens Redtenbacher, 1849. Benick and Lohse 1974, Muona 1991, Gusarov 2002, Klimaszewski et al. 2007, 2018, Majka and Klimaszewski 2008a, Brunke et al. 2012, Webster et al. 2016. Diagnosis. Body narrow, subparallel, minute, length 1.3–1.6 mm, colour light rust-brown, integument moderately glossy, forebody with pubescence fine and sparse, head convex, eyes small; antennomeres VI–X distinctly transverse (Fig. 11.61a); pronotum subquadrate; elytra short and truncate posteriad; abdomen slightly broadening apicad. MALE. Tergite VIII broad, truncate apically, antecostal suture arcuate (Fig. 11.61c); sternite VIII transverse and truncate apically, antecostal suture almost straight (Fig. 11.61d); tubus of median lobe of aedeagus in lateral view with ventral margin slightly sinuate, apical part broadly triangular, internal sac structures of distinctive shape (Fig. 11.61b). FEMALE. Tergite VIII elongate with apical margin broadly truncate medially, rounded laterally, antecostal suture arcuate (Fig. 11.61e); sternite VIII elongate with apical margin narrowly truncate medially, broadly rounded laterally, antecostal suture almost straight (Fig. 11.61f); spermatheca U-shaped, capsule and stem without distinct separation (Fig. 11.61g,h). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: BC, NB, NS, ON, YT [NPR] USA: NJ, RI. We record this species from the Yukon Territory for the first time based on the following: Whitehorse, Long Lake, 2016-VI-29, grasses, 60.7395
N, 135.0398
W, 641 m, sweeping, coll. B. Godin and D. Horwood (BGC) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: old-growth northern hardwood forest, mixed forests (NB), and a Sphagnum bog (ON). Adults in muskrat nest, among sticks, debris and mud on beaver dam, in leaf litter on clay near small brook, one captured in Lindgren funnel trap in old-growth

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hardwood forest. Collecting period: V, VI. Collecting method: sweeping grasses near lake (new data); netted in a mixed forest opening during the evening, sifting from forest litter and among sticks, debris and clay on a beaver dam, and from litter on brook margin, Lindgren funnel trap. Comments. The Canadian specimens listed under this name are conspecific with the central European specimens sensu Benick and Lohse, 1974. For details, see Majka and Klimaszewski 2008b. Subtribe Microglottina Fenyes, 1918 (Osswald et al. 2013) Diagnosis. Body compact and robust; head and pronotum with moderately coarse, small mesh reticulation; punctation fine; right mandible with strong median tooth, left mandible with obtuse, apical tooth; ligula rounded apically; pronotum distinctly transverse, hypomera slightly visible laterally; process of mesoventrite very long and moderately wide; anterior and middle tibiae with stiff setae.

Genus Crataraea C.G. Thomson, 1858 (Figs. 11.62a–h) (Klimaszewski et al. 2018) Diagnosis. Body compact and robust (Aleochara-like in form); head and pronotum with fine meshed reticulation and fine punctation; right mandible with strong median tooth, left mandible with obtuse, apical tooth; ligula rounded apically; pronotum distinctly transverse, about 1.5 times as wide as long; posterolateral margin of elytra with strong sinuate emargination, hypomera slightly visible laterally; process of mesoventrite very long and moderately wide; anterior and middle tibiae with stiff setae; male sternite VII strongly produced medially; median lobe of aedeagus with tubus bearing basal tooth; spermatheca with club-shaped capsule, stem thin and sinuate.

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11

Comments. Adults are usually found in plant and animal refuse in stables, granaries, and occasionally in bird and ant nests. 29. Crataraea (Figs. 11.62a–h)

suturalis

(Mannerheim){

Bolitochara suturalis Mannerheim, 1830. Klimaszewski et al. 2007, 2010, 2011, 2013, 2018, Moore and Legner 1975, Seevers 1978, Downie and Arnett 1996, Ashe 2001, Webster et al. 2009. Diagnosis. Body robust and compact (Aleochara-like), length 2.3–3.0 mm; colour dark brown with antennae, sutural and posterior margin of elytra, tip of abdomen, and legs/tarsi reddish-brown; integument slightly glossy, forebody with fine punctation and short and dense pubescence (Fig. 11.62a); head distinctly narrower than pronotum; antennomeres V–X moderately transverse; pronotum as long as elytra at suture, and about as wide as elytra; abdomen subparallel, slightly narrower basally than elytra. MALE. Tergite VIII transverse, truncate apically (Fig. 11.62d); sternite VIII semicircularly produced apically at middle, angulate at sides (Fig. 11.62e); median lobe of aedeagus in lateral view with ventral margin of tubus arcuate, with large acute basal tooth (Fig. 11.62b), in dorsal view as illustrated (Fig. 11.62c). FEMALE. Tergite VIII broadly arcuate apically (Fig. 11.62f); sternite VIII unevenly rounded apically, antecostal suture sinuate obtusely angulate medially (Fig. 11.62g); spermatheca with club-shaped capsule and sinuate narrow stem (Fig. 11.62h). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: BC, LB, NB, NS, ON, QC, SK, YT. USA: CA, IA, IL, IN, MA, MO, PA, RI, SC, VA, VT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mainly synantropic habitats but also found in spruce and other forests. Adults in heaps of straw, haystacks, stables, barns, on floor in house,

Tribe Oxypodini C.G. Thomson, 1859

pigeon and horse dung, carrion (dead fish, seabirds, sea mammals, horses), rotting mushrooms, rotting tree trunks with ants (Formica sp., Lasius sp.), bird nests, and forest leaf litter. One from barred owl nest contents in tree hole. Collecting period: V–VIII. Collecting method: sifting litter, pitfall traps. Subtribe Oxypodina C.G. Thomson, 1859 (Ashe 2001) Diagnosis. Head with frontal suture present in most species; pronotal hypomera visible or not in lateral view; tarsal formula 5-5-5. This is a heterogeneous assemblage that cannot be satisfactorily characterized (Ashe 2001).

Genus Calodera Mannerheim, 1830 (Fig. 11.63a–h) (Ashe 2001; Assing 2008; Klimaszewski et al. 2018) Diagnosis. Body robust, narrowly elongate, colour dark brown to black; head with neck about two-thirds as wide as maximum width of head; antennae thick, antennomeres V–X about 1.5–2.0 times as wide as long; ligula short, bifid at apex; eyes small, about half as long as genae in dorsal view; pronotum subquadrate, hypomera fully visible in lateral view; middle coxae narrowly separated; process of mesoventrite long and spine-like; process of metaventrite short and rounded apically; abdomen with first four visible tergites bearing deep, coarsely punctured basal impressions; median lobe of aedeagus simple with ventral margin of tubus arcuate; spermatheca with capsule small, spherical with large apical invagination, stem thin, long, and looped. Comments. Adults of Calodera inhabit moist habitats such as swamps, moist leaf litter near water such as rivers, streams, lakes, and floodplain forests. 30. Calodera bennetti Klimaszewski, sp. n. (Fig. 11.63a–h)

11

Tribe Oxypodini C.G. Thomson, 1859

147

Holotype (male): Canada, British Columbia, Queen Charlotte Is., 5.3 km SW Tow Hill, 22. VIII.1983, treading floating Sphagnum and Carex patches, J.M. Campbell (CNC). Paratypes: British Columbia, Queen Charlotte Is., 5.3 km SW Tow Hill, 22.VIII.1983, treading floating Sphagnum and Carex patches, J.M. Campbell (CNC) 2 male, 3 female; Queen Charlotte Is., 8.8 km SW Tow Hill, 22.VIII.1983, treading Sphagnum and Carex at edge of marsh, J.M. Campbell (CNC) 3 sex undetermined; Queen Charlotte Is., 7.3 km NW Rennell, Sound Road, Ghost Main Rd., 883-23, 800 ft., 19.VII.1983, sifting moss, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., 10.5 km NW Rennell, Sound Road, Ghost Main Rd., 900 ft., 21.VIII.1983, 83-102, sifting cold moss, J.M. Campbell (CNC) 2 male, 1 female; Queen Charlotte Is., Louise Is., 1.5 km W Skedans, 6.VIII.1983, 83-60, ex moss along seepage, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., 14 km E Masset, 13.VII.1983, 83-5, treading Carex marsh and sifting Alnus litter, J.M. Campbell (CNC) 2 sex undetermined. USA, Alaska, 8 mi 8 mi NW Haines, 3.VII.1968, J.M. Campbell and A. Smetana (CNC) 1 male.

Tergite VIII broadly arcuate apically (Fig. 11.63d); sternite VIII with apical margin broadly rounded at apex (Fig. 11.63e); median lobe of aedeagus in lateral view with ventral margin of tubus arcuate, moderately bent ventrad at base, apical part staright, narrowly elongate and produced ventrad (Fig. 11.63b); median lobe in dorsal view as illustrated (Fig. 11.63c); internal sac structures as illustrated (Fig. 11.63b, c). FEMALE. Tergite VIII truncate apically (Fig. 11.63f); sternite VIII rounded apically and slightly pointed medially, antecostal suture slightly sinuate laterally (Fig. 11.63g); spermatheca with capsule spherical, stem sinuate with single broad loop posteriorly (Fig. 11.63h).

Etymology. This species is dedicated to Dr. Rob Bennett, entomologist of the Royal British Columbia Museum, Victoria, Canada, who greatly contributed to the knowledge of insects of this province.

Comments. This is the only species of Calodera reported from western North America.

Diagnosis. Body slender, narrowly elongate, subparallel (Fig. 11.63a), length 2.9–3.5 mm; colour dark brown to almost black with basal antennomeres and legs/tarsi reddish-brown; integument moderately glossy, with punctation moderately coarse and dense, pubescence short; head distinctly narrower than pronotum; antennomeres VII–X distinctly transverse, about 1.5 times as broad as long; pronotum slightly shorter than elytra at suture, and about 1/4 narrower than maximum width of elytra; abdomen subparallel and tapering apically, basally slightly narrower than elytra, four basal abdominal impressions with coarse punctation. MALE.

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: adults in floating Sphagnum and Carex patches, and in Carex marshes, Sphagnum and Carex at edge of marsh, moss along seepage. Collecting period: V–IX. Collecting method: sifting moss, treading Sphagnum and Carex patches.

Genus Devia Blackwelder, 1952 (Fig. 11.64a–h) (Gusarov 2003; Klimaszewski et al. 2018) Diagnosis. Body large and robust, narrowly oval, flattened; integument with pubescence short and moderately dense, appearing velvety; pronotum subelliptical apart from hind angles, shield-like with hypomera not visible laterally; antennomere III longer than II, IV elongate, V–X moderately to disctinctly transverse; terminal palpomere thin and minute; mandibles bifid apically, right mandible with a small median tooth; abdomen with three first abdominal tergites shallowly impressed; genitalia distinctive (Fig. 11.64b, c, h).

148

31. Devia prospera (Erichson) (Fig. 11.64a–h) Oxypoda prospera Erichson, 1839. Gusarov 2003, Gouix and Klimaszewski 2007, Klimaszewski et al. 2007, 2010, 2011, 2015, 2018, Webster et al. 2009, Majka and Klimaszewski 2010. Diagnosis. Body broadly oval, slightly flattened, length 2.8–3.3 mm, colour reddish-brown to brown, with appendages rusty-brown, head and posterior abdomen dark brown (Fig. 11.64a); pubescence short, evenly distributed; pronotum large, shield-shaped, slightly wider than maximum width of elytra, hypomera not visible in lateral view; antennae robust, antennomeres V– X elongate; abdomen with first three visible tergites deeply impressed; hind tarsus with basal article highly elongate. MALE. Tergite VIII arcuate apically (Fig. 11.64d); sternite VIII produced medially at apex (Fig. 11.64e); median lobe of aedeagus in dorsal view with moderately large bulbus, tubus elongate (Fig. 11.64c); in lateral view ventral margin of tubus almost straight, bent obliquely ventrad at base (Fig. 11.64d). FEMALE. Tergite VIII arcuate apically (Fig. 11.64f); sternite VIII broadly rounded apically (Fig. 11.64g); spermatheca with capsule small, spherical, stem highly sinuate, and thin (Fig. 11.64h). DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, LB, MB, NB, NT, ON, QC, SK, YT. USA: AK, CA, CO, MA, MI, MN, MT, NH, NM, OR, SD, UT, WA, WI, WY. Palearctic: Europe, Asia. New locality data: British Columbia, BCol., Hwy 16, rest stop at Cluculz Creek, 53
540 5200 N, 123
350 2200 W, 776 m, 31.VII.2012, in Salix and Alnus litter with moss, 10 m from lake, A. Davies (CNC) 3 males, 22 sex undetermined; Valemont, Yellowjacket Creek, 52.707
N, 110.049
W, 771 m, 3-8.VI.2014, pitfall, Kinbasket # 1061, R-mature, Charlene Wood (RBCM) 13 males, 10 females; Valemont, N Packside Creek,

11

Tribe Oxypodini C.G. Thomson, 1859

52.3778
N, 119.163
W, 759 m, pitfall, Kinbasket # 1031, C. Wood debris, Charlene Wood (RBCM) 2 males, 4 females. COLLECTION AND HABITAT DATA. Habitat (Klimaszewski et al. 2018). In NB, commonly found in moist leaf litter on the margins of vernal ponds in silver maple swamps and mixed forests. Elsewhere, in human settlements in stables, barns where it occurs in heaps of straw, haystacks. Also uncommon in a disturbed field with forbs and grasses as well as in rotting organic debris, mushrooms, willow humus and moss, and forest litter. In BC, specimens were found in Salix and Alnus litter with moss, 10 m from a lake. Collecting period: IV–VIII. Collecting method: sifting litter, pitfall traps.

Genus Gnathusa Fenyes, 1910 (Figs. 11.65a–g and 11.66a–g) (Klimaszewski et al. 2014, 2018) Diagnosis. Body compact, robust, subparallel, length 3.5–4.3 mm; colour dark brown to black, integument with distinct meshed microsculpture; head large with mandibles extremely long and sickle-shaped, apices very slender, crossing each other in resting position, each bearing long spine or tooth; infraorbital carina strong and complete; ligula moderately deeply bilobed; apical palpomere needle-shaped; frontal suture of head absent; pronotal hypomera visible in lateral view; anterior margin of mesoventrite with short V-shaped basal carina, remaining mesoventer not carinate; process of mesoventrite long, triangular basally, then narrowly produced and extending to about three-quarters length of mesocoxal cavities, process of mesoventrite triangular in shape and short, isthmus short; median lobe of aedeagus with large crista apicalis, internal sac structures variable in shape, paramere with apical lobe narrowly elongate, bearing four macrosetae; spermatheca pipe-shaped with capsule spherical or tubular, stem elongate, narrow.

11

Tribe Oxypodini C.G. Thomson, 1859

149

Key to Species of Gnathusa

33. Gnathusa tenuicornis Fenyes (Fig. 11.66a–g)

1. Anterior margin of labrum spinose ................. .......................................... Gnathusa eva Fenyes – Anterior margin of labrum with short fine setae but lacking coarse spines ..................................... ............................. Gnathusa tenuicornis Fenyes

Gnathusa tenuicornis Fenyes Klimaszewski and Winchester Klimaszewski et al. 2012, 2014, 2018.

32. Gnathusa eva Fenyes (Fig. 11.65a–g) Gnathusa eva Fenyes, 1909. Klimaszewski et al. 2012, Webster et al. 2016. Diagnosis. Body length 2.9–3.6 mm, sides subparallel; colour dark brown to almost black, with tarsi and antennae reddish-brown (Fig. 11.65a); head round, equal in length to pronotum; anterior margin of labrum spinose; antennomeres V–X slightly to strongly transverse; pronotum transverse, slightly narrower than maximum width of elytra, anterior angles fairly evident; abdomen subparallel, middle segments as wide as elytra, slightly narrowing basad and apicad. MALE. Tergite VIII distinctly, very truncate apically (Fig. 11.65c); sternite VIII with apical margin broadly triangularly produced, apex slightly rounded (Fig. 11.65d); median lobe of aedeagus in lateral view with ventral margin of tubus arcuately bent ventrad basally, apex narrow and rounded (Fig. 11.65b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 11.65e); sternite VIII parabolically rounded apically (Fig. 11.65f); spermatheca pipe-shaped, with capsule small, narrowly tubular, stem long, thin and slightly sinuate (Fig. 11.65g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, SK, YT. USA: CA. COLLECTION AND HABITAT DATA. Habitat: in YT specimens were captured in coniferous woodchip pile. Collecting period: V, IX. Collecting method: pitfall traps.

1921. 2002,

Diagnosis. Body length 2.5–3.7 mm, sides subparallel; colour light brown to almost black, with antennae, tarsi and often elytra and apical part of abdomen rust-brown (Fig. 11.66a); integument with microsculpture dense and surface highly glossy; head round to somewhat quadrate and almost as wide as pronotum, labrum with short fine setae but lacking coarse spines; pronotum transverse, subequal to slightly narrower than maximum width of elytra, slightly shorter than elytra at suture, angles somewhat evident; abdomen subparallel; antennomeres VI–X subquadrate, apical one short and broadly oval. MALE. Tergite VIII broadly arcuate apically (Fig. 11.66c); sternite VIII with apical margin obtusely produced, at apex broadly rounded (Fig. 11.66d); median lobe of aedeagus in lateral view with ventral margin of tubus bent obliquely ventrad at base, slightly sinuate, apex thin, pointed (Fig. 11.66b). FEMALE. Tergite VIII truncate apically (Fig. 11.66e); sternite VIII rounded apically (Fig. 11.66f); spermatheca pipeshaped, with capsule spherical, stem long, thin, highly sinuate, neck weakly sclerotized, neck to capsule angle variable (Fig. 11.66g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, YT. USA: AK, CA, OR. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in western Canada recorded from a Sitka spruce forest, Upper Cordilleran coniferous forests, including subxeric lodgepole pine forests, mesic white spruce and lodgepole pine stands, sprucedominated subhygric and hygric forests, and a subalpine meadow at 3000 feet in moss seepage on slope. At other sites in cold moss along a

150

stream, from gravel at edge of stream, and from litter in a mixed aspen and spruce forest. In NB, in moss and leaves under alders near a brook in an eastern white-cedar swamp, from under cobblestones and gravel in sand on a partially shaded cobblestone bar near outflow of brook into a river, and in moss along a shaded springfed brook in a boreal forest area. Collecting period: V–VIII. Collecting method: sifting forest litter and cold moss, pitfall traps, aspirating from gravel and sand along stream margins.

Genus Mniusa Mulsant and Rey, 1875 (Fig. 11.67a–g) (Klimaszewski et al. 2014, 2018) Diagnosis. Body moderately robust with sides subparallel or body narrowly oval, length 2.0–3.2 mm; colour dark brown to black, integument with distinct meshed microsculpture and moderately dense punctation and pubescence; head large with mandibles broad and long, not crossing each other in resting position left mandible with a small tooth, right one with a slightly larger tooth at base of arcuate cutting edge, apices distinctly narrowly elongate [more than in Ocyusa and less than in Gnathusa]; infraorbital carina strong and complete; ligula shallowly split apically; penultimate maxillary palpomere minute, apical one needle-shaped; labrum narrow and transverse, apical edge entire; frontal suture of head absent; pronotum moderately transverse, about 1.3 times as wide as long, disc with pubescence along midline directed anteriad or obliquely anteriad in about apical one-third, and posteriad or obliquely posteriad medio-basally; process of mesoventrite without longitudinal carina, triangular basally and then produced and extending from one-third to almost two-thirds length of mesocoxal cavities; process of metaventrite triangular and short; isthmus short to long; median lobe of aedeagus strongly produced ventrally; spermatheca with capsule spherical or tubular, narrowed posteriorly into

11

Tribe Oxypodini C.G. Thomson, 1859

L-shaped neck, invagination small, stem narrowly elongate. Species occur in forested habitats. 34. Mniusa yukonensis (Klimaszewski and Godin) (Fig. 11.67a–g) Ocyusa yukonensis Klimaszewski and Godin, in Klimaszewski et al. 2012. Klimaszewski et al. 2014, 2018. Diagnosis. Body narrowly elongate and broadest at elytra, length 2.8–3.0, mm,colour dark brown to almost black, elytra sometimes with reddish tinge, antennae and legs reddishbrown (Fig. 11.67a); forebody highly glossy, with dense microsculpture, dense punctation and pubescence; head round, narrower than pronotum; pronotum transverse, with sides distinctly arcuate, narrowed anteriad, slightly narrower than elytra; elytra at suture about as long as pronotum or slightly longer; abdomen broadly arcuate laterally, slightly narrower than elytra, first three visible tergites with basal impressions; antennomeres V–X slightly transverse. MALE. Tergite VIII truncate apically (Fig. 11.67c); sternite VIII with apical margin very broadly produced, very broadly rounded at apex (Fig. 11.67d); median lobe of aedeagus with ventral margin of tubus about straight and slightly bent ventrad near base, internal sac with complex structures (Fig. 11.67b). FEMALE. Tergite VIII truncate apically, lateral angles sharp (Fig. 11.67e); sternite VIII with apical margin very obtusely produced, broadly rounded at apex (Fig. 11.67f); spermatheca with capsule narrowly elongate, sac-shaped, neck L-shaped, stem narrow and long (Fig. 11.67g). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, NS, QC, YT. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: nothing is known about specific micro-habitat associations

11

Tribe Oxypodini C.G. Thomson, 1859

of this species. All known adults were captured in various kinds of traps in various forest types: rich Appalachian hardwood forest with some conifers, old-growth white spruce/balsam fir forest, balsam fir and sugar maple stands, mature white spruce with feather moss, and a red spruce forest. Collecting period: V–VII. Collecting method: pitfall traps, flight intercept traps, and Lindgren funnel traps.

Genus Ocyusa Kraatz, 1856 (Fig. 11.68a–g) (Lohse et al. 1990; Klimaszewski et al. 2014, 2018) Diagnosis. Body moderately robust, subparallel, narrowly oval, length 2.5–3.5 mm; colour dark brown to almost black, integument with distinct meshed microsculpture, punctation and pubescence moderate to dense; head large with mandibles broad and long, not crossing each other in resting position, left mandible with a small tooth, right one with a slightly larger tooth at base of arcuate cutting edge, apices moderately narrowly elongate (less than in Mniusa); infraorbital carina strong and complete; ligula shallowly split apically; penultimate maxillary palpomere minute, apical one needle-shaped; labrum narrow and transverse, apical edge entire; fronto-clypeal phragma present and visible externally as frontal suture between antennal pits; pronotum moderately transverse, about 1.3 times as wide as long, disc with pubescence directed posteriad or obliquely posteriad along midline, pronotal hypomera visible in lateral view; anterior margin of mesoventer without longitudinal carina; process of mesoventrite triangular basally, then produced and extending to about two-thirds length of mesocoxal cavities, process of metaventrite narrowly triangular in shape and short, isthmus short; median lobe of aedeagus strongly bent ventrad at base; spermatheca S-shaped with capsule spherical, neck short, stem elongate and sinuate. Canadian species are known from wetland habitats.

151

35. Ocyusa canadensis Lohse (Fig. 11.68a–g) Ocyusa canadensis Lohse, in Lohse et al. 1990. Brunke et al. 2012, Klimaszewski et al. 2014, 2015, 2018. Diagnosis. Body narrowly elongate, broadest at elytra, length 2.5–3.0 mm; colour dark brown to almost black, with tarsi, two basal antennal articles and tibiae rust-brown, rest of antennal segments brown (Fig. 11.68a); forebody highly glossy, with dense microsculpture, punctation and pubescence moderate; head round, narrower than pronotum; pronotum transverse, rectangular in shape with sides distinctly arcuate, narrower than elytra; elytra at suture about as long as pronotum; abdomen broadly arcuate laterally, slightly narrower than elytra, first three visible tergites with basal impressions; antennomeres V–X subquadrate to slightly elongate. MALE. Tergite VIII broadly arcuate apically (Fig. 11.68c); sternite VIII broadly parabolic apically (Fig. 11.68d); median lobe of aedeagus with ventral margin of tubus almost straight, strongly bent ventrad at base, internal sac with complex structures (Fig. 11.68b). FEMALE. Tergite VIII truncate apically (Fig. 11.68e); sternite VIII evenly rounded apically (Fig. 11.68f); spermatheca with capsule semi-spherical, apical invagination deep and evenly narrow, stem broadly S-shaped (Fig. 11.68g). DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific provincial record], NB, NF, ON, SK, YT. USA: AK. We record this genus and species from BC for the first time based on the following: British Columbia, 20 mi. E. Hope, Manning Pk., 21. VI.1968, Campbell and Smetana (CNC) 1 male; BColl, Hwy 16 btw, Milligan Rd. and Wilhite Rd., W of Rose Lake, 54
24.00N, 126
04.22W, 784 m, 31.VII.2012, sedges in slough, A. Davies (CNC) 1 female; BColl., Smithers, perimeter trail wetland, S of Hwy 16, 490 m, 31.VII.2012, in moss in swamp, A. Davies (CNC) 1 female.

152

COLLECTION AND HABITAT DATA. Habitat: lake margins in emergent Carex and grasses. Some also on moist soil/gravel among sedges (Klimaszewski et al. 2018). In BC, specimens were collected in sedges in slough and in moss in swamp. Collecting period: VI– VII. Collecting method: treading Carex and grasses into water and examining moist soil and hand collecting (aspirating specimens).

Genus Oxypoda Mannerheim, 1830 (Figs. 11.69–11.87) (Klimaszewski et al. 2006, 2018) Diagnosis. Body subparallel, narrowly oval or fusiform, usually with abdomen strongly tapering; integument with pubescence fine, short and dense, in some species appearing silky, with numerous macrosetae especially on abdomen; head with frontal suture, infraorbital carina present, neck absent; mouthparts generalized (similarly developed across the genera); mandibles simple except right mandible with median tooth; pronotum usually highly convex with posterior angles very narrowly rounded, hypomera not visible in lateral view; process of mesoventrite slender and acute; tarsal formula 5-5-5; parameres with elongate apical lobe bearing four macrosetae: short (reduced) one at apex, moderately long one at middle, and two long ones at base; tergite VIII and sternite VIII usually similar in both sexes but in some taxa male sternite shaped differently, antecostal suture different in male and/or female in various species; median lobe of aedeagus with tubus well developed, in dorsal view usually subparallel for most of its length and gradually or strongly narrowed at apex, bulbus moderately large and bearing frontal and ventral carinae, the latter not always easily visible due to weak sclerotization, internal sac with complex structures, length of flagellum variable, usually shorter than tubus but occasionally longer than median lobe; spermatheca with capsule usually spherical but in some taxa elongate, stem long, sinuate and in some taxa looped posteriorly.

11

Tribe Oxypodini C.G. Thomson, 1859

Comments. Some members of Oxypoda may appear superficially like some species of Myllaena Erichson. However, Myllaena differs in having extremely elongate glossae extending over the labrum, and a 4-4-5 tarsal formula. There are at least ten subgenera of Oxypoda recognized in the western Palearctic but they are not all well defined. Many Nearctic species are difficult to assign to those subgenera, for some Nearctic species new subgenera would have to be erected to uniformly apply this system in the Nearctic. For this reason we decided to use informal species groups where species are presumably closely related. The species groups and species within each group are listed alphabetically. Species of Oxypoda are usually captured in forest litter, and are common in moss, damp leaf mold, fungi, and may also be found on carrion and dung. Key to Species of Oxypoda (There is some overlap in states of some external characters, and the genitalia should be routinely checked for all members of this genus to confirm the identifications arrived at in the key. Antennomere IV is often less elongate in females than in males) 1. Antennomeres VII–XI extremely swollen (Fig. 11.82a).... Oxypoda stanleyi Klimaszewski and McLean – Antennomeres VII–XI not as above.............. 2 2. Body dull in lustre, pubescence of forebody short and dense (Fig. 11.80a); pronotum large, distinctly broader than elytra (Fig. 11.80a); median lobe of aedeagus and spermatheca as illustrated (Fig. 11.80b, c, h, i) ............................ ....................................Oxypoda orbicollis Casey – Body more or less glossy, pubescence moderately long and dense in most; other characters not as above..............................................................3 3. Eyes small, almost three times shorter than postocular area of head (Fig. 11.69a); median lobe of aedeagus and spermatheca as illustrated

11

Tribe Oxypodini C.G. Thomson, 1859

153

(Fig. 11.69b, c, h)................................................. ....................Oxypoda canadensis Klimaszewski – Eyes moderate in size to large, two times or less as long as postocular area of head; median lobe of aedeagus and spermatheca different ......4

spermatheca as illustrated (Figs. 11.70b, c, h and 11.74b, c, h) .....................................................10 – Pronotum dark brown to almost black, median lobe of aedeagus and spermatheca not as above ..........................................................................11

4. Antennomere IV slightly to strongly elongate (Figs. 11.70a, 11.72a, 11.73a, 11.74a, 11.75a, 11.83a, and 11.86a)............................................5 – Antennomere IV subquadrate to transverse, (Figs. 11.71a, 11.76a, 11.77a, 11.78a, 11.79a, 11.81a, 11.84a, and 11.87a)............................. 12

10. Pronotum slightly longer than elytra at suture (Fig. 11.70a), median lobe of aedeagus and spermatheca as illustrated (Fig. 11.70b, c, h) ............. ................................ Oxypoda convergens Casey – Pronotum as long as elytra at suture (Fig. 11.74a); median lobe of aedeagus and spermatheca not as above (Fig. 11.74b, c, h) ............. ................Oxypoda longicarinata Klimaszewski

5. Antennomeres VII–IX subquadrate to slightly transverse (Figs. 11.72a and 11.86a) .................6 – Antennomeres VII–IX moderately to distinctly transverse (Figs. 11.70a, 11.73a, 11.74a, 11.75a, 11.83a, and 11.85) .............................................7 6. Elytra red brown in central/sutural area of disc, dark brown laterally and in scutellar area, integument densely punctate, strongly glossy, pubescence short and inconspicuous (Fig. 11.86a)....... .................................... Oxypoda impressa Casey – Elytra uniformly dark brown, integument moderately densely punctate, slightly glossy, pubescence long and distinct (Fig. 11.72a).......... ........................... Oxypoda opaca (Gravenhorst)

11. Pronotum slightly transverse, width to length ratio 1.5, distinctly narrower than elytra at base (Fig. 11.73a), median lobe of aedeagus and spermatheca as illustrated (Fig. 11.73b, c, h) ............. .......................... Oxypoda gatosensis Bernhauer – Pronotum moderately transverse, width to length ratio 1.4 (Fig. 11.75a); median lobe of aedeagus and spermatheca not as above (Fig. 11.75b, c, h)............ Oxypoda irrasa Casey

7. Body with head and abdomen dark brown and pronotum and elytra uniformly pale reddish brown (Fig. 11.85); genitalia missing in only known holotype specimen.................................... .....................................Oxypoda glenorae Casey – Body colour not as above..............................8

12. Pronotum densely and coarsely punctate, some punctures along middle line of disc confluent (Fig. 11.84a); median lobe of aedeagus and spermatheca as illustrated (Fig. 11.84b, c, h) ...... ....................Oxypoda vancouveri Klimaszewski – Pronotal punctures sparse or dense but always fine, punctures along midline not confluent (Figs. 11.71a, 11.76a, 11.77a, 11.78a, 11.79a, 11.81a, and 11.87a); median lobe of aedeagus and spermatheca not as above.......................... 13

8. Body with lateral and posterior margin of pronotum and parts of elytra yellow (Fig. 11.83a), median lobe of aedeagus and spermatheca as illustrated (Fig. 11.83b, c, h) ............. ..........................................Oxypoda sylvia Casey – Body approximately uniformly dark brown, median lobe of aedeagus and spermatheca not as above ..................................................................9

13. Body broad, oval, flattened, more or less uniformly brown (Fig. 11.71a); median lobe of aedeagus and spermatheca as illustrated (Fig. 11.71b, c, h)................................................. ..................................Oxypoda manitobae Casey – Body moderately broad to narrow, subparallel or ovoid, moderately convex; median lobe of aedeagus and spermatheca not as above .......... 14

9. Pronotum light brown, brown with lighter margins or reddish brown (Figs. 11.70a and 11.74a), median lobe of aedeagus and

14. Antennomeres VIII–X strongly transverse, up to 2 ½ as wide as long (Figs. 11.76a, 11.77a, and 11.79a).......................................................15

154

11

Tribe Oxypodini C.G. Thomson, 1859

– Antennomeres VIII–X moderately transverse, less than two times as wide as long, (Figs. 11.78a, 11.81a, and 11.87a).......................................... 17

spermatheca not described ................................... ..................................... Oxypoda regressa Casey

15. Pubescence very short, dense and with silky appearance, punctation fine, length 2.4–3.0 mm (Fig. 11.77a)................ Oxypoda lacustris Casey – Pubescence moderately long, moderately dense and not appearing silky, punctation moderate to coarse, length 1.7–2.6 mm (Figs. 11.76a and 11.79a).......................................................16

Convergens Species et al. 2006, 2018)

16. Head as broad as 2/3 of the maximum width pronotum (Fig. 11.79a), body moderately glossy, punctation of forebody moderately dense (Fig. 11.79a), male median lobe of aedeagus as illustrated (Fig. 11.79b, c), spermatheca with short median portion of stem and broad posterior loop (Fig. 11.79h, i)Oxypoda frigida Bernhauer – Head nearly as broad as 3/4 of the maximum width pronotum (Fig. 11.76a), body strongly glossy, punctation of forebody dense (Fig. 11.76a), male undescribed, spermatheca with long median portion of stem and narrow posterior loop (Fig. 11.76b) ..... Oxypoda famula Casey 17. Head narrow, about half as broad as the maximum width of pronotum (Fig. 11.78a), body length 2.0–2.4 mm, pronotum strongly converging apically (Fig. 11.78a), median lobe of aedeagus and spermatheca as illustrated (Fig. 11.78b, c, h)........ Oxypoda nimbata Casey – Head broad, at least as broad as 2/3 of the maximum width of pronotum (Figs. 11.81a and 11.87a), remaining characters different............ 18 18. Body length 2.5–2.7 mm, pronotum approximately oval in outline with sides slightly converging apically (Fig. 11.81a), median lobe of aedeagus and spermatheca as illustrated (Fig. 11.81b, c, h)................................................. ............................Oxypoda smithi Klimaszewski – Body length 1.7 mm, pronotum of different shape with sides strongly converging apically (Fig. 11.87a), median lobe of aedeagus and

Group (Klimaszewski

Diagnosis. Body narrowly oval or subparallel, length 3.0–4.5 mm; antennomere IV elongate in both sexes; pronotum distinctly transverse; elytra at suture as long as pronotum or only slightly shorter; median lobe of aedeagus with tubus broadly subparallel in dorsal view, in lateral view bulbus moderately large, ventral margin of tubus arcuate or slightly sinuate with apex narrow and rounded, flagellum shorter than tubus, internal sac with complex structures, base of flagellum heart- or V-shaped in dorsal view, apical lobe of paramere moderately elongate; shape of spermatheca variable, with capsule roughly spherical or slightly elongate, with broad apical invagination, stem moderately long to long, sinuate or coiled posteriorly. 36. Oxypoda (Fig. 11.69a–h)

canadensis

Klimaszewski

Oxypoda canadensis Klimaszewski, in Klimaszewski et al. (2006). Klimaszewski et al. (2011, 2012, 2018). Diagnosis. Body length 2.8–3.5 mm, subparallel, broadest at middle of elytra, colour uniformly dark brown with reddish tinge, or with legs, one to three basal antennomeres, and tip of abdomen slightly lighter, forebody with pubescence and punctation dense, integument moderately glossy with isodiametric microsculpture, glossy (Fig. 11.69a); head broadest at basal third, slightly more than half as wide as pronotum; eyes about one fourth as long as genae in dorsal view; antennae moderately long and slender, antennomeres I–IV elongate, V–X quadrate to slightly transverse; pronotum moderately convex, moderately transverse and about one fourth broader than long, broadest at middle, pubescence

11

Tribe Oxypodini C.G. Thomson, 1859

on disc directed anteriad apically along midline and obliquely posteriad from midline elsewhere; elytra slightly broader than pronotum, slightly shorter at suture than pronotum, pubescence on disc directed straight posteriad; abdomen gradually slightly tapering toward apex. MALE. Tergite VIII elongate, arcuate apically, antecostal suture slightly arcuate (Fig. 11.69d); sternite VIII elongate, with apical margin triangularly produced, apex narrowly rounded, antecostal suture almost straight (Fig. 11.69e); median lobe of aedeagus with tubus subparallel for most of its length in dorsal view (Fig. 11.69c), in lateral view ventral margin of tubus sinuate, at apex narrowly rounded and not bent (Fig. 11.69b), internal sac with complex structures (Fig. 11.69b, c). FEMALE. Tergite VIII elongate, arcuate apically, antecostal suture broadly arcuate (Fig. 11.69f); sternite VIII elongate, almost evenly rounded apically, antecostal suture sinuate, obtusely angular medially (Fig. 11.69g); spermatheca with capsule moderately large, spherical, with moderate, rounded apical invagination, stem long, with several tightly clustered posterior coils (Fig. 11.69h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], LB, MB, NF, NT, ON, QC, YT. USA: AK, NH. We record this species from BC for the first time based on the following: British Columbia, 55 km W. of Watson Lk., 29.VI.2015, 59.6843129.1627, 693 m, firekill, mushroom, sifting, coll. B. Godin and D. Horwood (LFC), 1 female. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mixed wood, deciduous birch-dominated or aspendominated forests, deciduous stands with developing white spruce understories, fir forests; also riparian habitats. Adults in forest litter, rotting fungi, mosses including Sphagnum sp., rotten logs, in lining of deserted beaver lodge, and in nest of Larus marinus L. (black-backed gull). Collecting period: V–IX, XII (most abundant in VI). Collecting method: pitfall traps, sifting

155

forest litter and processing through Berlese funnels, Luminoc® pitfall-light traps. 37. Oxypoda convergens Casey (Fig. 11.70a–h) Oxypoda convergens Casey, 1893. Klimaszewski et al. 2005, 2006, 2011, 2018. Diagnosis. Body length 3.0–3.4 mm, fusiform, broadest at middle of elytra, colour dark brown with legs, two to three basal antennomeres, and tip of abdomen rust-brown, in some specimens elytra and margins of pronotum also rust-brown (Fig. 11.70a); forebody with pubescence moderately long and dense and punctation dense; integument moderately glossy with faint isodiametric microsculpture; head broadest at level of posterior margins of eyes, three fifths as broad as pronotum; eyes about as long as genae in dorsal view; antennae moderately long, moderately stout, antennomere IV elongate, V–X slightly transverse, more so toward apex of antenna; pronotum moderately convex, moderately transverse, about one third broader than long, broadest in basal third, pubescence on disc directed anteriad apically and obliquely outward from midline elsewhere; elytra about as broad as pronotum and at suture slightly shorter than pronotum, pubescence on disc directed straight posteriad; abdomen tapering toward apex (Fig. 11.70a). MALE. Tergite VIII elongate, with apical margin truncate medially, broadly rounded laterally, antecostal suture almost straight except laterally (Fig. 11.70d); sternite VIII elongate with apical margin triangularly produced, apex rounded, antecostal suture sinuate (Fig. 11.70e); median lobe of aedeagus in dorsal view with tubus swollen at middle, sides broadly arcuate (Fig. 11.70c), in lateral view ventral margin of tubus almost straight, bent obliquely ventrad at base, narrowly rounded apically (Fig. 11.70b), internal sac with complex structures as illustrated (Fig. 11.70b, c). FEMALE. Tergite VIII elongate, with apical margin truncate medially, broadly rounded laterally, antecostal suture slightly sinuate (Fig. 11.70f); sternite VIII similar

156

to tergite VIII but antecostal suture distinctly sinuate (Fig. 11.70g); spermatheca with capsule large, roughly heart-shaped, apical invagination very deep, stem short, S-shaped (Fig. 11.70h). Oxypoda convergens differs from the remaining Nearctic Oxypoda species in having a large, usually dark brown body (length 3.0–3.4 mm), antennomeres VI–X slightly transverse, and the shape of the median lobe of the aedeagus (Fig. 11.70b, c) and the spermatheca (Fig. 11.70h) are distinctive. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], LB, NB, NF, NS, ON, QC. USA: IA, MO, NY. We record this species from BC for the first time based on the following: British Columbia, 160 km W. of Watson Lk., 27.VI.2015, 59.9056131.4924, 872 m, firekill, decidous forest, sifting litter, coll. B. Godin and D. Horwood (LFC), 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: various forest types including white spruce, mature black spruce, red spruce, mature 50-year-old balsam fir, oak, maple, yellow birch/balsam fir white pine forests, red oak forests, mixed forests, and old eastern white cedar forest; often very common in decaying gilled mushrooms in the above forest habitats, especially those dominated hardwoods. Also in riparian habitats, and a meadow with Salix shrubs. Several collected from decaying corncobs and cornhusks. Collecting period: V– XI (most common in August and September). Collecting method: unbaited and carrion-baited pitfall traps, processing forest litter through Berlese funnels, Luminoc pitfall-light traps, Lindgren traps, intercept traps, and treading leaf litter at the edge of a pond, sifting decaying mushrooms. 38. Oxypoda manitobae Casey (Fig. 11.71a–h)

11

Tribe Oxypodini C.G. Thomson, 1859

Oxypoda manitobae Casey, 1911. Lohse et al. 1990 [as O. lata]. Diagnosis. Body length 3.0–3.8 mm, broadly fusiform, broadest at middle of elytra, uniformly dark brown or with legs, two to three basal antennomeres, posterior parts of tergites and tip of abdomen slightly lighter, in some specimens elytra and edges of pronotum also rust-brown, pubescence and punctation of forebody dense, integument slightly glossy, with isodiametric microsculpture (Fig. 11.71a); head broadest at posterior level of eyes, about two thirds as broad as pronotum; eyes shorter than postocular region; antennae moderately long and moderately stout, antennomere IV subquadrate, V quadrate to slightly transverse, VI–X moderately transverse and more so toward apex of antenna (Fig. 11.71a); pronotum moderately convex, strongly transverse and about one third broader than long, broadest in basal part, pubescence fine, directed anteriad apically along midline and elsewhere obliquely posteriad from midline of disc, punctation fine; elytra, slightly broader than pronotum, about as long as pronotum at suture, pubescence directed straight posteriad; abdomen slightly tapering toward apex; basal tarsomere of hind tarsus as long as two following ones combined. MALE. Tergite VIII truncate apically and shallowly emarginated medially, antecostal suture approximately straight medially (Fig. 11.71d); sternite VIII with apex triangularly, sharply produced, antecostal suture strongly sinuate (Fig. 11.71e); median lobe of aedeagus with tubus subparallel in dorsal view (Fig. 11.71c), broadly arcuate ventrally in lateral view (Fig. 11.71b); internal sac with complex structures (Fig. 11.71b, c). FEMALE. Tergite VIII truncate apically, antecostal suture approximately straight medially (Fig. 11.71f); sternite VIII rounded apically, antecostal suture sinuate (Fig. 11.71g); spermatheca with moderately large spherical capsule bearing deep, broad invagination, stem sinuate and hooked posteriorly (Fig. 11.71h).

11

Tribe Oxypodini C.G. Thomson, 1859

DISTRIBUTION. Origin: Nearctic. CANADA: BC, MB, SK. USA: CO. It is probably broadly distributed in northern Canada and along the Rocky Mountain range. COLLECTION AND HABITAT DATA. Habitat: arctic habitats or in the Rocky Mts. Collecting period: VII, VIII. Collecting method: not recorded. 39. Oxypoda 11.72a–h)

opaca

(Gravenhorst){ (Fig.

Aleochara opaca Gravenhorst 1802. Horion 1967, Hoebeke 1990, Klimaszewski et al. 2006, 2007, 2011, 2018, McLean et al. 2009, Webster et al. 2009. Diagnosis. Body length 3.7–4.5 mm, broad, broadest at middle of elytra, colour dark brown with antennomeres I–II, posterior or middle of elytra, posterior margins of abdominal tergites and legs lighter brown (Fig. 11.72a); forebody with pubescence and punctation dense; integument moderately glossy, with isodiametric microsculpture; head broadest at basal third, about half as wide as pronotum; eyes longer than postocular region, moderately large; antennae slim, antennomeres I–IV elongate, V– X subquadrate; pronotum slightly flattened medially, distinctly transverse and about one third broader than long, broadest in basal third, pubescence on disc directed anteriad apically along midline and obliquely posteriad from midline elsewhere; elytra slightly broader and at suture about as long as pronotum, pubescence on disc directed straight posteriad; abdomen gradually narrowing toward apex, basal tarsomere of hind tarsus as long as two following ones combined. MALE. Tergite VIII elongate, apical margin broadly rounded on each side of shallow medial emargination, antecostal suture almost straight (Fig. 11.72d); sternite VIII elongate with apical margin triangularly produced, antecostal suture slightly arcuate or slightly sinuate (Fig. 11.72e); median lobe of aedeagus with tubus broad and triangular apically in dorsal view (Fig. 11.72c), in

157

lateral view bent ventral margin arcuately ventrad medially, slightly elongate apically (Fig. 11.72b); internal sac bearing complex structures (Fig. 11.72b, c). FEMALE. Tergite VIII elongate, apical margin rounded on each side of shallow median emargination, antecostal suture almost straight (Fig. 11.72f); sternite VIII elongate with apical margin broadly arcuate medially, broadly rounded laterally, antecostal suture straight or sinuate, closer to basal margin medially (Fig. 11.72g); spermatheca with capsule spherical, apical invagination large, stem S-shaped (Fig. 11.72h). DISTRIBUTION. Origin: Palearctic, and adventive in North America. CANADA: BC, LB, NB, NF, NS, ON, QC. USA: NC, NY, PA, SC, TN, VT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: various forest types including white spruce, mature black spruce, red spruce, mature 50-year-old balsam fir, oak, maple, yellow birch/balsam fir white pine forests, red oak forests, mixed forests, and old eastern white cedar forest; often very common in decaying gilled mushrooms in the above forest habitats, especially those dominated by hardwoods. Also in wetland habitats, and a meadow with Salix shrubs. Several collected from decaying corncobs and cornhusks. Collecting period: V–XI (most common in August and September). Collecting method: sifting decaying mushrooms and litter, treading leaf litter at edge of a pond, processing forest litter through Berlese funnels, unbaited and carrionbaited pitfall traps, Luminoc pitfall-light traps, Lindgren traps, intercept traps. Gatosensis Species Group (Klimaszewski et al. 2006) Diagnosis. The gatosensis species group may be characterized by the following combination of characters: body medium-sized, length 2.8–3.8 mm, broadly oval; antennomere IV strongly elongate, V–X quadrate to slightly

158

11

Tribe Oxypodini C.G. Thomson, 1859

transverse; pronotum slightly transverse and about one fourth narrower than elytra; elytra large and at suture as long as or longer than pronotum; abdomen broadly curved laterally in dorsal view; median lobe of aedeagus consisting of narrowly subparallel tubus with swelled sides at the middle in dorsal view, ventral margin of tubus broadly arcuate to nearly straight; flagellum shorter than tubus; internal sac with complex structures; bulbus broadly oval, moderate in size, with more or less protruding crista apicalis; apical lobe of paramere strongly elongate; spermatheca with capsule spherical and bearing large invagination, stem S- or U-shaped and twisted posteriorly.

(Fig. 11.73c), and broadly arched with apex sharply pointed in lateral view (Fig. 11.73b). FEMALE. Tergite VIII slightly produced apically, antecostal suture approximately straight (Fig. 11.73f); sternite VIII truncate apically, antecostal suture slightly sinuate (Fig. 11.73g); spermatheca approximately S-shaped with moderately large spherical capsule, deep, broad invagination and twisted posterior portion of stem (Fig. 11.73h).

40. Oxypoda 11.73a–h)

Bernhauer (Fig.

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA.

Oxypoda gatosensis Bernhauer 1905, Moore and Legner 1975.

New locality data: British Columbia, BCol., W of Terrace, Shames Mtn., road to ski area, 54
280 1400 N, 128
560 0500 W, 320 m, 29.VII.2012, in moss and alder litter at cold stream, A. Davies (CNC) 1 male.

gatosensis

Diagnosis. Body length 2.8–3.8 mm, mediumsized, subparallel, approximately uniformly brown or rust-brown, with antennae, legs, most of elytra and posterior margins of tergites light brown (Fig. 11.73a); pubescence of forebody dense; integument moderately glossy, with isodiametric microsculpture; head broadest behind eyes, about two thirds as broad as pronotum; eyes slightly shorter than the postocular region; antennomere IV elongate, V–X quadrate to slightly transverse; pronotum strongly convex, slightly transverse, broadest below middle, about one-fifth broader than long, pubescence directed anteriad apically along midline and obliquely posteriad from midline of disc elsewhere; elytra distinctly broader than pronotum, at suture longer than pronotum, pubescence directed straight posteriad; abdomen broadly arcuate laterally. MALE. Tergite VIII approximately as long as broad, rounded apically, antecostal suture approximately straight (Fig. 11.73d); sternite VIIII with slightly pointed apex, antecostal suture weakly sinuate and slightly pointed medially (Fig. 11.73e); median lobe of aedeagus swollen medially in dorsal view

Oxypoda gatosensis is externally similar to O. longicarinata from which it may be separated by the shape and the proportion of pronotum, and the shape of genitalia.

COLLECTION AND HABITAT DATA. Habitat: in moss and alder litter at cold stram. Collecting period: VII. Collecting method: sifting litter and moss. 41. Oxypoda longicarinata Klimaszewski (Fig. 11.74a–h) Oxypoda longicarinata Klimaszewski, in Klimaszewski et al. 2006. Diagnosis. Body 2.83.0 mm, broadly oval, approximately uniformly brown or rust-brown, antennae and in some specimens apical portion of abdomen light brown (Fig. 11.74a); pubescence of forebody dense; integument moderately glossy, with isodiametric microsculpture; head broadest behind eyes, about two thirds as broad as pronotum; eyes about as long as the postocular region; antennomere IV elongate, V–X quadrate to moderately transverse; pronotum strongly convex, distinctly transverse, broadest below the

11

Tribe Oxypodini C.G. Thomson, 1859

middle, about one third broader than long, pubescence directed anteriad apically along midline and obliquely posteriad from midline of disc elsewhere; elytra distinctly broader and at suture about as long pronotum, pubescence directed straight posteriad; abdomen broadly arcuate laterally in dorsal view; basal tarsomere of hind tarsus longer than two following ones combined. MALE. Tergite VIII slightly produced apically, antecostal suture straight (Fig. 11.74d); sternite VIII with rounded apex, antecostal suture strongly sinuate and pointed medially (Fig. 11.74e); tubus of the median lobe of the aedeagus slightly swollen medially in dorsal view (Fig. 11.74c), and broadly arched with apex narrowly rounded in lateral view (Fig. 11.74b); crista apicalis of bulbus strongly protruding (Fig. 11.74b, c). FEMALE. Tergite VIII rounded apically, antecostal suture approximately straight (Fig. 11.74f); sternite VIII truncate apically, antecostal suture sinuate (Fig. 11.74g); spermatheca approximately U-shaped with moderately large spherical capsule, deep, broad invagination and twisted posterior portion of stem (Fig. 11.74h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: coastal Sitka spruce forest and mixed wood forest. Collecting period: V–VII. Collecting method: intercept traps. Irrasa Species Group (Klimaszewski et al. 2006) Diagnosis. Body narrowly oval, length 3.2–3.5 mm; antennomere IV distinctly elongate, VII–X distinctly transverse; pronotum strongly transverse; elytra at suture slightly longer than pronotum; bulbus streamlined with tubus in dorsal view; in lateral view tubus bent ventrad in about half length, apical part narrow and rounded; spermathecal capsule spherical with small apical invagination, stem narrow, long and sinuate.

159

42. Oxypoda irrasa Mäklin (Fig. 11.75a–h) Oxypoda irrasa Mäklin 1853. Moore and Legner 1975, Lohse and Smetana 1985, Klimaszewski et al. 2006, Webster et al. 2016. Diagnosis. Body length 3.2–3.5 mm, narrowly oval, broadest at basal third of elytra, abdomen subparallel (Fig. 11.75a); uniformly black or dark brown, with paler reddish brown basal two antennal segments, legs, elytra (dark brown reddish posteriorly) and apical part of abdomen; pubescence long and moderately dense, and punctation of forebody moderately dense; integument glossy, with isodiametric microsculpture; head broadest at basal third, distinctly broader than half of pronotal width; eyes moderately large, about as long as postocular area; antennae slender, antennomeres I–III strongly elongate, IV slightly elongate, V–VI quadrate or slightly transverse, and VII–X moderately transverse; pronotum moderately convex, strongly transverse and about one third broader than long, broadest in basal third, pubescence directed anteriad apically along midline and obliquely posteriad from midline of disc elsewhere; elytra slightly broader than pronotum and at suture slightly longer than pronotum, pubescence directed approximately straight posteriad; abdomen subparallel and slightly tapering apically. MALE. Tergite VIII transverse and arcuate apically, antecostal suture approximately straight (Fig. 11.75d); sternite VIII slightly transverse and produced apically, antecostal suture broadly arcuate (Fig. 11.75e); median lobe of aedeagus approximately narrowly oval, consisting of anteriorly converging tubus, which is streamlined with bulbus in dorsal view (Fig. 11.75c); ventral margin of tubus angularly connected with bulbus, slightly sinuate and with apex narrow and produced ventrad in lateral view (Fig. 11.75b); flagellum shorter than tubus; internal sac with two elongate subapical structures (Fig. 11.75b, c); bulbus with large crista apicalis (Fig. 11.75b). FEMALE. Tergite VIII similar to that of male (Fig. 11.75f); sternite VIII similar to

160

that of male but apical margin truncate (Fig. 11.75g); spermatheca with capsule spherical and bearing small invagination, stem thin, long and sinuate (Fig. 11.75h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], SK, YT. USA: AK, OR. We record this species from BC for the first time based on the following: British Columbia, Atlin, Pine Creek, 21.VI.2015, 59.7116133.5787, 939 m, Alder soil sifting, coll. B. Godin and D. Horwood (LFC), 1 female; Boya Lake, 28. VI.2015, 59.3639-129.1035, 698 m, mushroom sifting, coll. B. Godin and D. Horwood (LFC), 1 male; Valemont, Yellowjacket ck., 3-8. VI.2014, pitfall, Kinbasket # 1054, 52.703119.049, 771 m, R-Mature, coll. Charlene Wood (LFC), 1 female; Valemont, Peatland, 23-28. VI.2014, pitfall, Kinbasket # 1106, 52.763119.163, 763 m, T. Wood removal, coll. Charlene Wood (LFC), 1 female; Valemont, N Packsaddle ck., pitfall, Kinbasket # 1024, 52.3778-119.163, 759 m, R-Mature forest, coll. Charlene Wood (BGC), 1 male, 1 female; same dat except: Kinbasket # 1028, 52.78-119.164, 767 m (BGC), 1 female. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2006). Habitat: in AB, adults occurred in all forest cover types and all retention treatments, but were most abundant in unharvested stands. In BC, one specimen was captured from mushroom, one was found in alder soil, and a few from mature and cut forests. Collecting period: V–VIII, most abundant in VIIII. Collecting method: pitfall traps, sifting forest litter and processing it through Berlese funnels, sifting mushrooms. Lacustris Species Group (Klimaszewski et al. 2006, 2018) Diagnosis. Body narrowly oval, length 1.6–2.5 mm; antennomere IV subquadrate to

11

Tribe Oxypodini C.G. Thomson, 1859

moderately elongate; pronotum transverse; elytra at suture as long as pronotum or distinctly shorter; median lobe of aedeagus with tubus narrowly subparallel in dorsal view, in lateral view ventral margin of tubus arcuate at base and nearly straight anteriorly, apex bluntly truncate (except for O. nimbata), bulbus moderately large, flagellum shorter than tubus, internal sac with semicircular apical structure consisting of several interconnected elements in lateral view and falciform or evenly narrow subapical sclerite, apical lobe of paramere slender and extremely elongate; spermatheca capsule roughly spherical, with apical invagination moderate, constricted at apex, stem moderately long, sinuate and spirally looped posteriorly. 43. Oxypoda famula Casey (Fig. 11.76a–e) Oxypoda famula Casey, 1911. Moore and Legner, 1975, Klimaszewski et al. 2006. Diagnosis (based on female lectotype). Body length 1.7 mm; compact, robust, ovoid in shape, broadest at middle of elytra (Fig. 11.76a); colour dark brown with darker head, antennae, tip of abdomen, legs reddish brown; forebody with pubescence and punctation dense; integument glossy, with isodiametric microsculpture; head nearly as broad as 3/4 of the maximum width pronotum; eyes moderately large; antennae moderately slender, antennomere IV slightly transverse, V–X strongly transverse, incrassate apically; pronotum convex, flattened mediobasally, moderately transverse and about one fourth broader than long, pubescence on disc directed anteriad along midline apically and obliquely posteriad from midline elsewhere; elytra slightly broader than pronotum, at suture slightly longer than pronotum, pubescence on disc directed straight posteriad; abdomen subparallel, tapering apically from about a third posteriorly. MALE. Undescribed. FEMALE. Tergite VIII broadly arcuate apically (Fig. 11.76c); sternite VIII broadly arcuate apically, antecostal suture arcuate (Fig. 11.76d); spermatheca with

11

Tribe Oxypodini C.G. Thomson, 1859

capsule small, spherical, apical invagination moderate, stem long, sinuate and looped posteriorly (Fig. 11.76b); pygidium as illustrated (Fig. 11.76e). DISTRIBUTION. Origin: Nearctic. CANADA: BC [QCI, Massett]. USA: AK. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. 44. Oxypoda lacustris Casey (Fig. 11.77a–i) Oxypoda lacustris Casey 1906. Klimaszewski et al. 2006, 2011, 2015, 2018, Webster et al. 2009. Diagnosis. Body length 2.4–3.0 mm, very flexible, broad, subparallel, broadest at middle of elytra (Fig. 11.77a); colour light to dark brown, with antennae, tip of abdomen, legs and rarely elytra lighter, in some specimens outline of body with lighter reddish tinge; forebody with pubescence short and punctation dense; integument moderately glossy, with isodiametric microsculpture; head broadest at basal third, slightly more than half as wide as pronotum; eyes moderately large as long as postocular area; antennae moderately slender, antennomere IV subquadrate or slightly elongate, V–X strongly transverse; pronotum slightly flattened medially, distinctly transverse and about one third broader than long, broadest at about middle, pubescence on disc directed anteriad along midline apically and obliquely posteriad from midline elsewhere; elytra slightly broader than pronotum, at suture at least as long as pronotum, pubescence on disc directed straight posteriad; abdomen gradually tapering apically. MALE. Tergite VIII elongate, broadly rounded apically, antecostal suture virtually straight (Fig. 11.77d); sternite VIII elongate, parabolically rounded apically, antecostal suture sinuate (Fig. 11.77e); median lobe of aedeagus with tubus in dorsal view gradually narrowing from bulbus, subparallel toward apex (Fig. 11.77c), in lateral view ventral margin

161

curved obliquely ventrad basally, straight to bluntly truncate apex (Fig. 11.77b); internal sac with falciform subapical sclerite angular ventrally at base, with elongate posterior projection (Fig. 11.77b, c); flagellum moderately long (Fig. 11.77b); apical structure semicircular, consisting of several articulately connected elements (Fig. 11.77b). FEMALE. Tergite VIII similar to that of male but truncate apically (Fig. 11.77f); sternite VIII similar to that of male but antecostal suture almost straight (Fig. 11.77g); spermatheca with capsule spherical, apical invagination moderate, constricted at apex, stem long, sinuate and spirally looped posteriorly (Fig. 11.77h, i). Oxypoda lacustris differs from O. hiemalis and O. pseudolacustris in having the elytra at least as long as the pronotum (shorter than the pronotum in the two latter species). The spermatheca of O. lacustris is similar to that of O. pseudolacustris. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NT, ON, QC, SK, YT. USA: AK, WA. New locality data: British Columbia, Osoyoos, Hayness E.R., 4-IV-1.V.1995, AN, BCxhl, recovery after fire, pitfall trapN0. A1, G.G.E. Scudder (UBC) 1 male; Galiano Is., Bodega ridge, 16.VII-25.VIII.1990, pitfall P5, G.G.E. Scudder (UBC) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in predominantly wet habitats; in moist leaves on vernal pond margins in mixed forests and seasonally flooded silver maple forests and swamps, in leaf and grass litter at base of red maple in a Carex marsh, in moist leaf and grass litter in Carex and cattail marshes, and in drift material on a lake margin. Also in deciduous forests (oak, birch, aspen), fir and conifer dominated forests, coastal dunes and coastal barrens. Specimens were also found in gopher burrows and muskrat nests. One

162

11

Tribe Oxypodini C.G. Thomson, 1859

individual captured in Lindgren funnel trap in an old silver maple swamp. Collecting period: III– X. Collecting method: sifting forest litter, pitfall traps, Lindgren funnel traps, and sifting debris from burrows and nests of rodents.

invagination moderate, constricted at apex, stem long, broadly looped posteriorly (Fig. 11.78h).

45. Oxypoda nimbata Casey (Fig. 11.78a–h)

New locality data: British Columbia, Oliver, IR1, Water tower, 119.31
N, 49.10
W, 1.VI-7. VII.1994, Purshia association, AN. BGxl, pitfall trap, U 2-3, G.G.E. Scudder (UBC) 1 male; Vasseux Creek, Kennedy bench, 119.30
N, 49.16
W, 9.VI-5.VII.1995, Purshia association, AN, BGxh, pitfall trap, Z 1-3 and 1-4, G.G.E. Scudder (UBC) 2 females; Guliano Is., Bodera ridge, 16.VII-25.VIII.1990, pitfall P 5, G.G.E. Scudder (UBC) 1 female; Lillooet, 50.7286
N, 121.6184
W, 835 m, 23.VI.2013, moss, Berlese, Darren and Claudia Copley (RBCM) 1 male; Victoria, Fort Rod Hill, 48.4348
N, 123.4521
W, 10 m, 23.V.2015, Gary Oak, soil sifting, B. Godin and D. Horwood (RBCM) 1 male.

Oxypoda nimbata Casey, 1911. Bernhauer and Scheerpeltz, 1926, Moore and Legner, 1975, Klimaszewski and Winchester, 2002. Diagnosis. Body length 2.7–3.0 mm, slender, narrowly fusiform, broadest at middle of elytra with abdomen slightly tapering apically (Fig. 11.78a); colour reddish brown, head dark brown and remainder of body light brown to reddish light brown; forebody with pubescence and punctation dense; integument moderately glossy, with isodiametric microsculpture; head broadest at basal third, half as broad as pronotum; eyes small, about half as long as genae in dorsal view; antennomere IV subquadrate to slightly elongate, V subquadrate, VI–X distinctly so, to about two times as wide as long; pronotum moderately convex, slightly transverse, about one fifth broader than long, broadest in basal third, pubescence on disc directed anteriad along midline apically, and obliquely posteriad from midline elsewhere; elytra as broad as pronotum, pubescence on disc directed straight posteriad; abdomen slightly tapering posteriad. MALE. Tergite VIII elongate, broadly rounded apically, antecostal suture almost straight (Fig. 11.78d); sternite VIII elongate, broadly parabolically rounded apically, antecostal suture sinuate (Fig. 11.78e); median lobe of aedeagus with tubus narrowly subparallel in dorsal view (Fig. 11.78c), in lateral view ventral margin curved ventrad basally, straight to near apex, apex bluntly truncate (Fig. 11.78b); internal sac with apical structure semicircular (Fig. 11.78b, c); flagellum highly sinuate. FEMALE. Tergite and sternite VIII similar to those of male (Fig. 11.78f, g); spermatheca with capsule spherical, apical

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA.

COLLECTION AND HABITAT DATA. Habitat: transitional zone of Sitka spruce forest on Vancouver Island, with Purshia association in Okanagen Valley, and in soil of Garry Oak parkland. Collecting period: VI, VII. Collecting method: pitfall traps. Orbicollis Species Group (Klimaszewski et al. 2006, 2018) Diagnosis. Median lobe of aedeagus in dorsal view with tubus narrow, subparallel, strongly narrowing in apical part, apex rounded, bulbus small, oval, in lateral view tubus with ventral margin broadly arcuate, narrowly elongate and rounded apically, bulbus with crista apicalis enlarged, flagellum about as long as tubus, internal sac with complex structures, apical lobe of paramere moderately elongate; spermatheca with capsule spherical, with medium to large invagination, stem sinuate with large posterior loop.

11

Tribe Oxypodini C.G. Thomson, 1859

163

46. Oxypoda frigida Bernhauer (Fig. 11.79a–i) Oxypoda frigida Bernhauer, Klimaszewski et al. 2006, 2011, 2018.

1907.

Diagnosis. Body compact, subparallel, length 2.0–2.6 mm, colour brown or rust-brown with head dark brown and elytra and abdomen darker than pronotum and appendages (Fig. 11.79a); forebody with pubescence dense and moderately long; integument moderately glossy with fine isodiametric microsculpture; head broadest behind eyes, less than two thirds as wide as pronotum; eyes small, shorter than postocular region; antennomere IV slightly transverse, V–X strongly transverse, at least twice as wide as long; pronotum highly convex, slightly transverse, broadest in basal third, about one fourth broader than long, pubescence on disc directed anteriad along midline apically, and obliquely posteriad from midline elsewhere; elytra about as broad as pronotum, at suture distinctly shorter than pronotum, pubescence on disc directed straight posteriad; abdomen subparallel, slightly tapering apically. MALE. Tergite VIII slightly produced apically, antecostal suture sinuate (Fig. 11.79d); sternite VIII with apical margin obtusely angulate, apex broadly rounded, antecostal suture sinuate (Fig. 11.79e); median lobe of aedeagus in dorsal view subparallel, tubus tapering apically (Fig. 11.79c), in lateral view tubus bent obliquely ventrad at base, almost straight to narrowly rounded apex, bulbus with crista apicalis very large (Fig. 11.79b); internal sac structures complex, flagellum shorter than tubus (Fig. 11.79b, c). FEMALE. Tergite VIII rounded apically, antecostal suture arcuate (Fig. 11.79f); sternite VIII broadly rounded apically, antecostal suture virtually straight (Fig. 11.79g); spermatheca with capsule roughly spherical, with invagination moderately large, stem with broad, slightly spiral posterior loop (Fig. 11.79h, i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, NT, ON, QC, YT. USA: AK, NH.

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in hardwood-dominated stands with white spruce understories, balsam fir, old fir, black spruce, white spruce, maple, Salix and Alnus forests, alpine forests, and among shrubs in coastal limestone barrens. Adults in moss and litter near a pond, in leaf litter, in Sphagnum, and in mushrooms. Collecting period: V–X. Collecting method: Luminoc pitfall-light traps, sifting forest litter and processing it through Berlese funnels, and Lindgren funnel traps. 47. Oxypoda orbicollis Casey (Fig. 11.80a–i) Oxypoda orbicollis Casey, 1911. Moore and Legner 1975, Klimaszewski et al. 2006, 2011, 2015, 2018. Diagnosis. Body medium-sized, length 3.0–3.8 mm, narrowly spindle-shaped, uniformly brown or rust-brown (Fig. 11.80a); pubescence of forebody dense and very short; integument dull in luster with small isodiametric microsculpture; head broadest behind eyes, about two thirds as broad as pronotum; eyes small, shorter than the postocular region; antennomere IV elongate, V– X quadrate to slightly transverse; pronotum strongly convex, slightly transverse, broadest at the middle, about one fifth broader than long, pubescence directed anteriad apically along midline and obliquely posteriad from midline of disc elsewhere; elytra slightly broader and at suture shorter than pronotum, pubescence directed straight posteriad; abdomen tapering apically. MALE. Tergite VIII elongate, and rounded apically, antecostal suture arcuate (Fig. 11.80d); sternite VIII elongate and rounded apically, antecostal suture slightly sinuate (Fig. 11.80e); median lobe of aedeagus subparallel with strongly converging apical part in dorsal view (Fig. 11.80c), and broadly arched tubus with apex narrowly rounded in lateral view (Fig. 11.80b); flagellum with heart-shaped structure at base (Fig. 11.80c). FEMALE. Tergite VIII elongate, and rounded apically, antecostal suture

164

11

arcuate (Fig. 11.80f); sternite VIII elongate and rounded apically, antecostal suture slightly sinuate and pointed medially (Fig. 11.80g); spermatheca with approximately spherical capsule bearing deep and wide invagination, stem broadly looped (Fig. 11.80h, i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], QC, YT. USA: WI. We record this species from BC for the first time based on the following: British Columbia, Valemont, Peatland, 23-28.VI.2014, pitfall, Kinbasket # 1086, 52.762-119.164, 762 m, R-Mature, coll. Charlene Wood (LFC), 1 male. COLLECTION AND HABITAT DATA. Habitat: in AB, adults were collected only in deciduous-dominated stands. Individuals were found both in clearcuts and uncut stands. Collecting period: VII, VIII. Collecting method: pitfall traps, Luminoc pitfall-light traps, Lindgren traps, sifting forest litter and Sphagnum and processing it through Berlese funnels. Smithi Species Group (Klimaszewski et al. 2006) Diagnosis. The smithi species group may be distinguished by the following combination of characters: body moderately large, length 2.5–2.7 mm, subparallel; integument moderately glossy; antennomere IV subquadrate, and V–X moderately to strongly transverse; pronotum smaller and narrower than elytra, approximately round in shape; elytra at suture at most as long as pronotum; the median lobe of the aedeagus distinctly shaped; and by the S-shape of spermatheca lacking capsular invagination. 48. Oxypoda 11.81a–h)

smithi

Klimaszewski (Fig.

Oxypoda smithi Klimaszewski, Klimaszewski et al. (2006).

in

Tribe Oxypodini C.G. Thomson, 1859

Diagnosis. Body length 2.5–2.7 mm, subparallel, uniformly brown or rust-brown (Fig. 11.81a); pubescence of forebody moderately dense; integument moderately glossy with small isodiametric microsculpture; head broadest behind eyes, about two thirds of maximum width of pronotum; eyes large, about as long as postocular region; antennomere IV subquadrate, V–X moderately to strongly transverse, about two times as wide as long; pronotum approximately circular, strongly convex, slightly transverse, broadest in basal third, about one sixth broader than long, pubescence directed anteriad apically along midline and obliquely posteriad from midline of disc elsewhere; elytra much shorter than pronotum and at suture at most as long as pronotum, pubescence directed straight posteriad; abdomen subparallel and slightly tapering apically. MALE. Tergite VIII broadly arcuate apically, antecostal suture approximately straight (Fig. 11.81d). Sternite VIII broadly rounded apically, antecostal suture arcuate (Fig. 11.81e); median lobe of aedeagus gradually converging apically in dorsal view (Fig. 11.81c), tubus broadly arched with apex pointed ventrad in lateral view (Fig. 11.81b); flagellum approximately as long as tubus (Fig. 11.81b), bulbus broader than base of tubus in dorsal view (Fig. 11.81c) with small crista apicalis in lateral view (Fig. 11.81b). FEMALE. Tergite VIII slightly transverse and truncate apically, antecostal suture arcuate (Fig. 11.81f); sternite VIII truncate apically, antecostal suture sinuate and slightly pointed medially (Fig. 11.81g); spermatheca with spherical capsule lacking invagination, stem S-shaped (Fig. 11.81h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: VI, VII. Collecting method: not recorded. Stanleyi Species Group (McLean et al. 2009)

11

Tribe Oxypodini C.G. Thomson, 1859

165

Diagnosis. Oxypoda stanleyi species group is distinct from other Nearctic species of Oxypoda by the short macrosetae on the apical tergites and sternites VIII, and by its swollen antennae. 49. Oxypoda stanleyi Klimaszewski McLean (Fig. 11.82a–h)

and

Oxypoda stanleyi Klimaszewski and McLean, in McLean et al. 2009. Diagnosis. Body small, reddish-brown with dark brown head and at least posterior half of abdomen; length 1.6–1.9 mm, approximately narrowly oval, slightly broadest at middle of elytra, abdomen subparallel basally and gradually narrowed posteriad (Fig. 11.82a); forebody with scarcely visible microsculpture, sculpticells of flattened hexagonal shape; integument moderately glossy; pubescence sparse on head and abdomen and denser elsewhere; head broadest at posterior eye level, narrower than pronotum, pubescence directed anteriad centrally and slightly obliquely laterad laterally, frontal suture and infraorbital carina strong; eyes moderately sized, approximately as long as postocular area; antennae strongly swollen, antennomeres IV–X strongly transverse and incrassate; maxillary palpus with four palpomeres, last one needle-shaped; labial palpus with three palpomeres; glossae separated, V-shaped; pronotum moderately convex, strongly transverse ratio of maximum pronotal width/ length 1.3, pubescence directed straight posteriad along midline of disc and obliquely posteriorly elsewhere; elytra slightly broader at middle than pronotum and at suture slightly shorter than pronotum, pubescence directed approximately straight to slightly obliquely posteriad; abdomen subparallel basally and slightly tapering apically; three basal tergites with strong basal impressions; metatarsus with basal article elongate and at least as long as two following articles combined. MALE. Tergite VIII truncate apically, antecostal suture slightly sinuate medially (Fig. 11.82d); sternite VIII rounded apically, antecostal suture slightly sinuate medially (Fig. 11.82e); median lobe of aedeagus with moderately sized bulbus

and subparallel tubus abruptly tapering apically in dorsal view (Fig. 11.82c); venter of tubus arcuate basally and straight subapically ending sharply in lateral view (Fig. 11.82b); crista apicalis large (Fig. 11.82b); median lobe in dorsal view as illustrated (Fig. 11.82c); internal sac with complex structures (Fig. 11.82b, c). FEMALE. Tergite VIII similar to that of male (Fig. 11.82f); sternite VIII approximately truncate apically, antecostal suture straight medially (Fig. 11.82g); spermatheca with small spherical capsule with invagination facing up and long and narrow sinuate stem (Fig. 11.82h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: all specimens were captured in a closed canopy second-growth forest, mainly Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), with a bare forest floor with old needles and some ivy around traps (site 1), and in a closed canopy second-growth stand of mainly western hemlock (Tsuga heterophylla (Raf.) Sarg.), and a few Douglas-fir and western cedar (Thuja plicata Donn ex D. Don) with a bare forest floor with old needles. The thick antennae indicate that this species may be affiliated with ants, but this needs to be confirmed by field observations. Collecting period: IV–VII. Collecting method: pitfall traps, funnel trap. Sylvia Species Group (Klimaszewski et al. 2006, as grandipennis group; Klimaszewski et al. 2018, as sylvia group) Diagnosis. Body length 3.5–4.6 mm, narrowly oval, pronotum small, elytra large and long; median lobe of aedeagus in dorsal view with tubus broad, gradually narrowing apically, base of bulbus bearing elongate projection, which is split along median line and forms two subparallel arms in ventral view, bulbus oval, evenly coalescent with tubus dorsally, with crista apicalis narrowly elongate, flagellum about as long as tubus, internal sac with complex structures; apical lobe

166

of paramere moderately elongate; spermatheca with capsule spherical, with large invagination, stem with large posterior loop, twisted posteriorly. 50. Oxypoda sylvia Casey (Fig. 11.83a–h) Oxypoda sylvia Casey 1906. Klimaszewski et al. 2018. Moluciba grandipennis Casey, 1911; Seevers 1978. As Oxypoda: Gusarov 2003, Klimaszewski et al. 2005, 2006, 2011, 2015. Synonymized by Klimaszewski et al. (2018). Diagnosis. Body large, length 3.5–4.6 mm, narrowly oval, broadest at apex of elytra, colour brown yellow, head and posterior abdomen usually dark brown, with apical antennomeres, marginal parts of pronotum, basal and posterior portions of elytra and base of abdomen yellowbrown (Fig. 11.83a); forebody with pubescence moderately dense and long, punctation fine, moderately dense; integument moderately glossy, with isodiametric microsculpture; head broadest at level of eyes, two thirds as broad as pronotum; eyes about as long as genae in dorsal view; antennae long and slender, antennomere IV moderately elongate, V–X moderately transverse; pronotum flattened, transverse, broadest in basal third, about one fifth broader than long, pubescence on disc directed obliquely apicad along apical part of median line and posterolaterad from midline elsewhere; elytra distinctly elongate, longer than pronotum at suture, pubescence on disc directed slightly obliquely outward from suture; abdomen gradually narrowed to apex. MALE. Tergite VIII slightly elongate and rounded apically, antecostal suture almost straight (Fig. 11.83d); sternite VIII elongate, with apex produced, antecostal suture sinuate (Fig. 11.83e); median lobe of aedeagus broad dorsally (Fig. 11.83c), with long narrow projection diverging from base of bulbus ventrally below median lobe in lateral view (Fig. 11.83b). FEMALE. Tergite VIII slightly elongate, rounded apically,

11

Tribe Oxypodini C.G. Thomson, 1859

antecostal suture arcuate (Fig. 11.83f); sternite VIII rounded apically, antecostal suture distinctly sinuate (Fig. 11.83g); last two abdominal segments with internal sclerotized structure; spermatheca with capsule moderately large, spherical, apical invagination broad, stem broadly looped posteriorly (Fig. 11.83h). Oxypoda sylvia is probably closely related to the Palearctic Oxypoda spectabilis Märkel. Both have a similar subdivided process attached to the base of the aedeagus (in ventral view). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK, YT. USA: AK, ME, NH. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: many forest types, including hardwood, red oak, mixed, mature red spruce, red spruce and birch, old red pine, spruce and balsam fir forests, and old-growth eastern white cedar forests. Adults common in decaying and rotten mushrooms in many of the above forests types. Some in group of Pholiota mushrooms of trunk of dead Populus; another under coyote dung on forest road. Collecting period: V–X (most common in September). Collecting method: sifting mushrooms, forest litter, dung, and moss; some of above material also processed through Berlese funnels, Luminoc® pitfall-light traps, unbaited pitfall traps. Vancouveri et al. 2006)

Species

Group (Klimaszewski

Diagnosis. Body length 2.7–3.0 mm, narrowly elongate, pronotum small, elytra distinctly longer and wider than pronotum; antennomere IV subquadrate, V–X moderately transverse; median lobe of aedeagus in dorsal view with tubus moderately broad, gradually narrowing apically, in lateral view tubus stronngly arcuate with narrow and rounded apex, flagellum slightly shorter than tubus, internal sac with apical structures;

11

Tribe Oxypodini C.G. Thomson, 1859

167

spermatheca with capsule tubular and L-shaped, with small invagination, stem narrow and long and strongly sinuate. 51. Oxypoda vancouveri Klimaszewski (Fig. 11.84a–h) Oxypoda vancouveri Klimaszewski, Klimaszewski and Winchester (2002)

in

COLLECTION AND HABITAT DATA. Habitat: interior and transition zone of Sitka spruce forest, and red cedar-dominated forest. Collecting period: VII–IX. Collecting method: pitfall and pan traps. Species Incertae Sedis [known only from the type specimens with missing genitalia] 52. Oxypoda glenorae Casey (Fig. 11.85)

Diagnosis. Body length 2.7–3.0 mm; narrowly ovoid, broadest at elytra (Fig. 11.84a); dark brown with paler apex of abdomen, parts of elytra and legs; punctation and pubescence of forebody dense; integument moderately glossy and with small isodiametric microsculpture; head broadest behind eyes, about two-thirds width of pronotum; eyes slightly longer than postocular area; antennomere IV subquadrate, V–X slightly to moderately transverse; pronotum strongly converging apically, broadest in basal third, about one fifth broader than long (Pronotal Index ¼ 1.2), pubescence on disc directed obliquely apicad along apical part of median line and posterolaterad from midline elsewhere; elytra subparallel, slightly broader than maximum width of pronotum, at suture slightly longer than pronotum; abdomen tapering posteriad. MALE. Tergite VIII slightly elongate and rounded apically, slightly pointed medially, antecostal suture arcuate (Fig. 11.84d); sternite VIII with apex produced and rounded, antecostal suture slightly sinuate (Fig. 11.84e); median lobe of aedeagus moderately broad and gradually narrowed apically in dorsal view (Fig. 11.84c), with small projection near base of bulbus ventrally in lateral view (Fig. 11.84b). FEMALE. Tergite VIII slightly elongate, rounded apically, antecostal suture arcuate (Fig. 11.84f); sternite VIII rounded apically, antecostal suture slightly sinuate (Fig. 11.84g); spermatheca with capsule tubular and L-shaped, apical invagination small, stem narrow and strongly sinuate (Fig. 11.84h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

Oxypoda glenorae Klimaszewski et al. 2006.

Casey,

1893.

Diagnosis. (based on lectotype with missing apical part of abdomen). Body length 2.9 mm; narrowly elongate (Fig. 11.85); reddish-yellow with dark brown head, and darker basal tergites mottled with reddish-brown patches; punctation and pubescence of forebody fine and very dense; integument strongly glossy and with small isodiametric microsculpture; head broadest behind eyes, about two-thirds width of maximum width of pronotum; eyes about as long as postocular area; antennomere IV elongate, V–X slightly to moderately transverse; pronotum strongly converging apically, broadest in basal third, about one fifth broader than long (Pronotal Index ¼ 1.2), pubescence on disc directed obliquely apicad along apical part of median line and posterolaterad from midline elsewhere; elytra at suture shorter than pronotum along midline; abdomen subparallel; basal tarsomere of hind tarsus extremely elongate, about as long as three following tarsomeres combined. Genitalia and terminalia of male and female unknown. DISTRIBUTION. Origin: Nearctic. CANADA: BC [Glenora]. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. 53. Oxypoda impressa Casey (Fig. 11.86a–d)

168

Oxypoda impressa Klimaszewski et al. 2006.

11

Casey,

1893.

Diagnosis (based on female type). Body length 2.7 mm; ovoid in shape (Fig. 11.86a); dark brown almost black, with reddish-brown central part of elytra, two basal antennomeres, and legs; punctation and pubescence of forebody fine and very dense; integument moderately glossy and with small isodiametric microsculpture; head broadest just behind eyes, about two-thirds width of pronotum; antennomere IV elongate, V–X subquadrate or moderately transverse; pronotum about as broad as elytra, strongly converging apically, broadest in basal third, about one fifth broader than long (Pronotal Index ¼ 1.2), pubescence on disc directed obliquely apicad along apical part of median line and posterolaterad from midline elsewhere, punctation dense and coarse; elytra at suture longer than pronotum along midline; abdomen subparallel; basal tarsomere of hind tarsus about as long as two following tarsomeres combined. MALE. Undescribed. FEMALE. Spermatheca unknown; tergite VIII truncate apicomedially and sinuate laterally (Fig. 11.86b); sternite VIII rounded apically and slightly produced medially (Fig. 11.86c); pygidium as illustrated (Fig. 11.86d). DISTRIBUTION. Origin: Nearctic. CANADA: BC [Glenora]. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. 54. Oxypoda regressa Casey (Fig. 11.87a–g) Oxypoda regressa Casey, 1911. Klimaszewski et al. 2006. Diagnosis. Body length 1.7 mm; narrowly elongate (Fig. 11.87a); reddish brown, with dark brown head and abdomen except for its apex; punctation and pubescence of forebody fine and

Tribe Oxypodini C.G. Thomson, 1859

moderately dense; integument moderately glossy and with small isodiametric microsculpture; head broadest behind eyes, about two-thirds width of pronotum; eyes moderately sized; antennomere IV subquadrate to slightly transverse, V–X moderately transverse; pronotum strongly converging apically, broadest in basal third, about one fourth broader than long (Pronotal Index ¼ 1.3), pubescence on disc directed obliquely apicad along apical part of median line and posterolaterad from midline elsewhere; elytra at suture distinctly longer than pronotum along midline; abdomen subparallel; basal tarsomere of hind tarsus about as long as two following tarsomeres combined. MALE. Aedeagus not described (missing from male syntype); tergite VIII arcuate apically (Fig. 11.87b); sternite VIII sinuate laterally and strongly produced medially (Fig. 11.87c); pygidium (Fig. 11.87d). FEMALE. Spermatheca not described (missing from female syntype); tergite VIII arcuate apically (Fig. 11.87e); sternite VIII arcuate apically (Fig. 11.87f); pygidium as illustrated (Fig. 11.87g). DISTRIBUTION. Origin: Nearctic. CANADA: BC [Victoria]. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded.

Ocalea Group of Genera (Figs. 11.88–11.100) (Klimaszewski and Pelletier 2004) Includes medium- to large-sized oxypodine genera, presumably closely related, and with majority of species found in moss at high altitudes. This group, except for Neothetalia canadiana Klimaszewski and Parocalea pseudobaicalica Lohse, is represented only in western North America. Pronotal index used in descriptions relates to ratio of pronotal maximum width to length, and elytral index relates to ratio of elytral maximum width to length.

11

Tribe Oxypodini C.G. Thomson, 1859

Genus Alfocalea Klimaszewski, 2004 (Fig. 11.88a–h) (Klimaszewski and Pelletier 2004) Diagnosis. Adults of the genus Alfocalea may be distinguished from those of other aleocharine genera by the following combination of characters: body narrowly elongate, subparallel, dark brown with brown to rust brown pronotum, elytra, and appendages; size medium, length 4.3–4.7 mm, integument more or less densely punctate and pubescent, forebody with distinct isodiametric microsculpture; antennomere IV quadrate in female to slightly elongate in male, VII–X transverse; maxillary palpus with four palpomeres, apical palpomere narrowly elongate and one-third length of penultimate one; galea expanding slightly beyond apex of lacinia, its apical lobe bearing dense short pubescence; apex of lacinia with row of closely spaced spines; labial palpus with three palpomeres; ligula slender and shallowly emarginated apically; mentum trapezoidal, apical margin slightly concave, with three long apical setae on each side; labrum transverse and usually slightly projecting medially; right mandible with distinct medial tooth; head quadrate to slightly elongate and almost as wide and as large as pronotum; pronotum transverse, broadest in about middle, and with arcuate edges evenly converging apically and basally, pubescence directed anteriad medially except for very basal part, laterad on each side of disc and often forming broadly arcuate lines; mesoventrite about as long as maximum diameter of mesocoxa, with partial median carina, carina flattened and broad, about one-third of mesoventrite length, mesoventral process long, narrow, and acutely pointed, extending about two-thirds length of mesocoxal cavities, isthmus short and no more than one-fourth of maximum diameter of mesocoxa; elytra about as broad as pronotum, moderately transverse, at suture at most as long as pronotum; abdomen with three basal tergites shallowly impressed basally; basal tarsomere of mesotarsus slightly longer than following one, and basal tarsomere of metatarsus as long as two following segments combined. MALE. Median lobe of

169

aedeagus with small and narrowly elongate crista apicalis at base of bulbus, tubus long, approximately arcuate medially, sinuate anteriorly, and strongly produced ventrally in lateral view, apex narrowly rounded. FEMALE. Spermatheca with small, spherical capsule and with short, narrow, apical invagination facing obliquely downwards, stem sinuate and slightly hooked posteriorly. Adults were collected at higher altitudes ranging from 975–2133 m. The species apparently prefer cool, high mountain habitats (Klimaszewski and Pelletier 2004). 55. Alfocalea montana Klimaszewski (Fig. 11.88a–h) Alfocalea montana Klimaszewski, Klimaszewski and Pelletier (2004).

in

Diagnosis. Body length 4.3–4.7 mm, dark brown with elytra, basal antennal segment, tarsi, and pronotum rust brown, forebody moderately glossy and abdomen strongly so (Fig. 11.88a); antennae stout, antennomere IV quadrate in female to slightly elongate in male, V–X slightly to strongly transverse, and terminal palpomere about as long as two preceding ones combined; pronotum strongly transverse (Pronotal Index ¼ 1.3), lateral edges broadly arcuate, punctures sparse, distance between punctures about 2–3 times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad, usually forming arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.6), about as long as pronotum; abdomen arcuate laterally and broadest in apical third, three basal sternite with deep impressions. MALE. Tergite VIII transverse, truncate apically, and without median emargination (Fig. 11.88d); sternite VIII slightly transverse, apical part broadly triangularly produced (Fig. 11.88e); median lobe of aedeagus with small and narrowly elongate crista apicalis at base of bulbus, tubus long, arcuate medially, sinuate anteriorly and strongly produced ventrally in lateral view

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(Fig. 11.88b), apex narrowly rounded (Fig. 11.88b); median lobe in dorsal view as illustrated (Fig. 11.88c). FEMALE. Tergite VIII similar to that of male (Fig. 11.88f); sternite VIII transverse and broadly rounded apically (Fig. 11.88g); spermatheca (Fig. 11.88h) with small, approximately spherical capsule and with short, narrow apical invagination facing obliquely downwards, stem sinuate and slightly hooked posteriorly. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: high mountain habitats at altitudes 975–2133 m. Collecting period: VI, VII, VIII. Collecting method: pitfall traps.

Genus Betocalea Klimaszewski, 2004 (Fig. 11.89a–h) (Klimaszewski and Pelletier 2004) Diagnosis. Body slightly flattened, narrowly elongate, subparallel, dark brown with rust brown pronotum, elytra, and appendages; medium sized, length 4.0–5.0 mm, integument moderately densely punctate and pubescent, forebody with distinct isodiametric microsculpture; antennomere IV nearly twice as long as wide, antennomeres V–VII elongate, and VIII– X subquadrate; maxillary palpus with four palpomeres, apical one narrowly elongate and one-third to nearly one-half length of penultimate palpomere; galea expanding slightly beyond apex of lacinia, its apical lobe bearing dense short pubescence; apex of lacinia with row of closely spaced spines; labial palpus with three palpomeres; ligula slender, deeply bifid; mentum trapezoidal, apical margin slightly concave, with three long apical setae on each side; labrum transverse and usually slightly projecting medially; right mandible with distinct medial tooth; head quadrate or slightly elongate and distinctly narrower than pronotum; pronotum transverse,

11

Tribe Oxypodini C.G. Thomson, 1859

broadest medially, lateral edges broadly arcuate, gradually converging basally and apically from middle of lateral edge, length ratio of base to apex 1.3 times, pubescence directed anteriad medially except for very basal part, and laterad on each side of disc; mesoventrite approximately as long as maximum diameter of mesocoxa, with median carina about as long as half of mesoventrite, mesoventral process long, narrow, and acutely pointed, extending to about two-thirds length of mesocoxal cavities, isthmus short and no more than one-fourth of maximum diameter of mesocoxa; elytra broader than pronotum, slightly transverse, at suture longer than pronotum; abdomen subparallel, basal three abdominal tergites deeply impressed basally. MALE. Median lobe of aedeagus, tubus narrowly triangular at apex in lateral view and bearing small projection in front of crista apicalis of bulbus, bulbus approximately as long as tubus in lateral view, crista apicalis oval and large; internal sac with two inconspicuous subapical structures; paramerite with welldeveloped and narrowly elongate apical lobe bearing two long subapical setae and two additional short setae, one on apex and one at base of apical lobe; median part of paramerite moderately elongate. FEMALE. Spermatheca with small, approximately spherical capsule bearing deep apical invagination facing downwards, attachment of spermathecal gland located at capsule base near neck of spermatheca; basal, medial, and posterior portions of spermatheca consist of narrow, tube-like, S-shaped stem. Adults occur in mountainous areas and in Douglas-fir dominated coastal forest in British Columbia (Klimaszewski and Pelletier 2004). 56. Betocalea pacifica Klimaszewski (Fig. 11.89a–h) Betocalea pacifica Klimaszewski, Klimaszewski and Pelletier (2004).

in

Diagnosis. Body length 4.0–5.0 mm, dark brown with elytra, basal antennomere, tarsi, and pronotum light brown with rust tinge, forebody

11

Tribe Oxypodini C.G. Thomson, 1859

moderately glossy and abdomen strongly so (Fig. 11.89a); antenna moderately stout, antennomeres IV–VII strongly elongate, and VIII–X slightly elongate in male and quadrate in female, terminal antennomere about as long as two preceding ones combined; pronotum strongly transverse (Pronotal Index ¼ 1.3), lateral edges slightly converging basally in basal half and moderately in apical half, length ratio of base to apex 1.2 times, pronotal punctures sparse, distance between punctures about two times diameter of a puncture, space between punctures with strong isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad on both sides; elytra transverse (Elytral Index ¼ 1.5). MALE. Tergite VIII transverse, truncate apically, and without median emargination (Fig. 11.89d); sternite VIII slightly transverse, apical part broadly triangularly produced (Fig. 11.89e); median lobe of aedeagus with large, oval crista apicalis at base of bulbus and with small triangular ventral projection anterior to crista apicalis, tubus long, approximately arcuate medially, moderately produced ventrally at apex, apex narrowly triangular and slightly pointed in lateral view (Fig. 11.89b); median lobe in dorsal view as illustrated (Fig. 11.89c). FEMALE. Tergite VIII similar to that of male (Fig. 11.89f); sternite VIII transverse and broadly rounded apically (Fig. 11.89g); spermatheca with small, approximately spherical capsule and with short, narrow apical invagination, stem slender and S-shaped (Fig. 11.89h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: transition zone of the Douglas-fir dominated coastal forest in west Vancouver, and the remaining two specimens were found in the Mt. Garibaldi area. Collecting period: V, IX, X. Collecting method: Malaise trap.

171

Genus Megocalea Klimaszewski, 2004 (Fig. 11.90a–i) (Klimaszewski and Pelletier 2004) Diagnosis. Body broad, moderately elongate, subparallel, large, length 5.5–6.0 mm, dark brown with paler and slightly rust brown pronotum, elytra, apical part of abdomen, and appendages; integument moderately densely punctate and pubescent, distance between punctures 2–3 times diameter of a puncture, forebody with distinct isodiametric microsculpture, strongly glossy; head elongate, narrowly produced in front of eyes and gradually narrowed posteriad; all antennomeres elongate; maxillary palpus strongly elongate, with four palpomeres, apical palpomere narrow and one-half length of penultimate one; galea expanding slightly beyond apex of lacinia, its apical lobe bearing dense short pubescence; apex of lacinia with row of closely spaced spines; labial palpus with three strongly elongate palpomeres, basal one swollen; ligula entire bearing two apical sensillae; mentum trapezoidal, apical margin slightly concave, with three longer apical setae on each side; labrum broad and transverse; right mandible with strong medial tooth; head elongate and distinctly narrower than pronotum; pronotum strongly transverse, broadest in the middle, and strongly narrowed anteriad and posteriad, pubescence directed anteriad medially and laterad on each side of disc, forming more or less arcuate lines; mesoventrite short, slightly less than maximum diameter of mesocoxa, with partial median carina, slightly longer than one-half length of mesoventrite, mesoventral process short, narrow, and acutely pointed, extending less than one-half length of mesocoxal cavities, isthmus long; elytra transverse, broader than pronotum, at suture slightly longer than pronotum; abdomen tapering apically, with three basal tergites deeply impressed basally. MALE. Median lobe of aedeagus with tubus triangularly shaped and bulbus strongly expanded in dorsal aspect, bulbus

172

11

shorter than tubus in lateral view, crista apicalis of bulbus small and attached apically to two long and arcuate processes; internal sac with flagellum shorter than tubus and with two pairs of subapical strong structures; paramerite strongly elongate medially, with well-developed, moderately long, and broad apical lobe bearing four moderately long setae. FEMALE. Spermatheca moderatelysized spherical capsule bearing moderately deep apical invagination facing downwards; stem tubelike, strongly sinuate, and S-shaped. Species of this genus are associated with moss and litter on the forest floor (Klimaszewski and Pelletier 2004). 57. Megocalea lemieuxi Klimaszewski (Fig. 11.90a–i) Megocalea lemieuxi Klimaszewski, Klimaszewski and Pelletier (2004)

Tribe Oxypodini C.G. Thomson, 1859

flagellum shorter than tubus and with two pairs of subapical structures. FEMALE (new description). Tergite VIII with apical margin arcuate (Fig. 11.90g); sternite VIII slightly elongate and with triangularly produced apical part (Fig. 11.90h); spermatheca moderately-sized spherical capsule bearing moderately deep apical invagination facing downwards; stem tube-like, strongly sinuate, and S-shaped (Fig. 11.90i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: hybrid spruce and subalpine fir in Smithers area, the prevailing climate was cool and wet. Collecting period: VIII, IX. Collecting method: unbaited pitfall traps.

in

Genus Metocalea Klimaszewski, 2004 Diagnosis. Body length 5.5–6.0 mm, dark brown with paler pronotum, elytra, base of antennae, and tarsi; entire body strongly glossy (Fig. 11.90a); antenna moderately stout, all antennomeres strongly elongate, and terminal one about as long as two preceding antennomeres combined; pronotum strongly transverse and distinctly narrower than elytra (Pronotal Index ¼ 1.3), broadest at middle and strongly converging apically and basally, punctures sparse, distance between punctures 2–3 times diameter of a puncture, space between punctures with distinct isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad, forming arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.6), about 1.4 times longer than pronotum. MALE. Tergite VIII with apical margin truncate (Fig. 11.90e); sternite VIII slightly elongate and with triangularly produced apical part (Fig. 11.90f); median lobe of aedeagus with dorsally triangular tubus and expanded bulbus (Fig. 11.90d), bulbus shorter than tubus in lateral view, crista apicalis of bulbus small and attached apically to two long and arcuate processes (Fig. 11.90b,c); internal sac with

(Fig. 11.91a–i) (Klimaszewski and Pelletier 2004) Diagnosis. Head narrower than pronotum, moderately sized and distinctly smaller than pronotum; antennomeres IV–VII slightly elongate, VIII–X quadrate, and terminal antennomeres about as long as two preceding ones combined; ligula with small apical emargination, about one-third to one-fourth of its length; terminal palpomere of labial and maxillary palps without pseudosegments; terminal palpomere of maxillary palpus less than one-half length of penultimate one; labrum with apical margin slightly produced medially; pronotum broadest in middle, lateral sides broadly arcuate, with distinct, isodiametric microsculpture and moderately dense punctation, distance between punctures about two times diameter of a puncture, pubescence directed anteriad in upper median line, posteriad basally and lateroposteriad, forming arcuate lines, on sides; elytra at most as long as pronotum; mesoventrite with vestigial carina, about one-fourth length of mesoventrite; abdomen subparallel, first three visible tergites with deep

11

Tribe Oxypodini C.G. Thomson, 1859

173

impression and fourth with shallow impression, basal line of impressions more or less sinuate. MALE. Median lobe of aedeagus with bulbus bearing oval crista apicalis and additional pseudocrista in proximity, tubus broadly arcuate laterally and with rounded narrow apex; internal sac membranous, subapical structures distinct and strongly sclerotized. FEMALE. Spermatheca with large, approximately spherical capsule and with moderately sized, narrow invagination, stem looped posteriorly; a unique U-shaped, strongly sclerotized structure present inside pygidium of female. Adults of this genus were found in predominantly hybrid spruce, Picea glauca  engelmannii (Parry) Engelmann, and subalpine fir, Abies lasiocarpa (Hook.) Nutt., in cool and wet environment (Klimaszewski and Pelletier 2004). 58. Metocalea lindgreni Klimaszewski (Fig. 11.91a–i) Metocalea lindgreni Klimaszewski, Klimaszewski and Pelletier (2004).

in

Diagnosis. Body length 4.0–4.5 mm, dark rust brown with paler pronotum, elytra, base of antennae, and tarsi, forebody moderately glossy and abdomen strongly so (Fig. 11.91a); antenna moderately stout in both sexes, antennomeres IV– VII slightly elongate, VIII–X quadrate, and terminal antennomere about as long as two preceding ones combined; pronotum strongly transverse (Pronotal Index ¼ 1.2–1.3), lateral edges broadly arcuate, punctures sparse, distance between punctures about two times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad, usually forming arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.7), about as long as pronotum. MALE. Tergite VIII truncate apically, and with shallow median emargination (Fig. 11.91d); sternite VIII with apical part broadly and triangularly produced (Fig. 11.91e);

median lobe of aedeagus with small, oval crista apicalis at base of bulbus and with additional small crista in front, tubus long, broadly arcuate, and slightly produced ventrally in lateral view, apex rounded (Fig. 11.91b); median lobe in dorsal view as illustrated (Fig. 11.91b); internal sac structures elongate and well sclerotized (Fig. 11.91b, c). FEMALE. Tergite VIII truncate apically, slightly emarginated medially (Fig. 11.91f); sternite VIII broadly rounded apically (Fig. 11.91g); spermatheca with large, approximately spherical capsule and with short, narrow apical invagination, stem looped posteriorly (Fig. 11.91h); U-shaped and strongly sclerotized structure present at ventral part of pygidium (Fig. 11.91i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. New locality data: British Columbia, Vancouver Is., Mt. Cain, 50.13
N, 126.21
W, 10.VIII-23.VIII.1996, 16.1 PAN 2, N. Winchester (CNC) 3 females. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: forest types were predominantly hybrid spruce, Picea glauca engelmannii (Parry) Engelmann, and subalpine fir, Abies lasiocarpa (Hook.) Nutt., and the prevailing climate was cool and wet. Collecting period: V–VIII. Collecting method: unbaited pitfall traps, pantraps.

Genus Neoisoglossa Casey, 1893 (Fig. 11.92a–h) (Casey 1893; Klimaszewski and Pelletier 2004) Diagnosis. Body subparallel, length 3.0–4.0 mm, strongly glossy, microsculpture of forebody weak and scarcely visible, punctation sparse and fine, distance between punctures 2–3 times diameter of a puncture, pubescence short; colour rust brown or dark brown with paler and slightly rust brown pronotum, elytra, apical part

174

of abdomen, and appendages; head narrowly produced in front of eyes; antennomere IV slightly to strongly elongate, V–VII quadrate or elongate, VIII–X quadrate to slightly transverse, and terminal antennomere slightly less than two preceding ones combined; ligula with two apical sensillae and appearing slightly emarginated apically (not examined in all species); terminal palpomere of labial and maxillary palps without pseudosegment; terminal palpomere of maxillary palpus less than one-half length of penultimate one; labrum with apical margin slightly produced medially; pronotum rectangular in shape, slightly narrower than elytra, emarginated laterally, convex, lateral edges arcuate, disc with fine and sparse punctation, distance between punctures 2–3 times diameter of a puncture; mesoventral basal carina very short, about one-fifth of the mesoventrum (not examined in all species). Tergite and sternite VIII of both sexes with numerous very long and protruding apically distributed macrosetae. Includes one, semi-desert habitat species, found in association with ArtemisiaPurshia plants. 59. Neoisoglossa agnita (Casey) (Fig. 11.92a–h) Ocalea agnita Casey, 1911. Moore and Legner 1975, Klimaszewski and Pelletier 2004. Diagnosis. Body length 3.3–3.4 mm, dark brown with paler pronotum, elytra, base of antennae, and tarsi, entire body strongly glossy (Fig. 11.92a); antenna moderately stout, antennomere IV slightly elongate, V–VII slightly and VIII–X strongly transverse, and terminal antennomere less than two preceding segments combined; pronotum strongly transverse (Pronotal Index ¼ 1.3), lateral edges broadly arcuate, punctures sparse, distance between punctures 2–3 times diameter of a puncture, pubescence directed anteriad along median line of disc and laterad on both sides; elytra transverse (Elytral Index ¼ 1.6). MALE. Tergite VIII rounded apically with long macrosetae

11

Tribe Oxypodini C.G. Thomson, 1859

(Fig. 11.92d); sternite VIII broadly triangularly produced apically, apex rounded with long macrosetae (Fig. 11.92e); median lobe of aedeagus with large and oval crista apicalis of bulbus, tubus long, broadly arcuate, and moderately produced ventrally, apex narrowly elongate and pointed in lateral view (Fig. 11.92b), tubus broad and subparallel in dorsal view (Fig. 11.92c); internal sac structures small and weakly sclerotized (Fig. 11.92b, c). FEMALE. Tergite VIII arcuate apically with long macrosetae (Fig. 11.92f); sternite VIII broadly rounded apically with long macrosetae (Fig. 11.92g); spermatheca with moderately large, approximately spherical capsule and with deep, narrow apical invagination, stem narrow, sinuate medially, and irregularly looped posteriorly (Fig. 11.92h). DISTRIBUTION. Origin: Nearctic. CANADA: AB [Waterton Pk.], BC [new provincial record]. USA: AZ. We report this species from BC for the first time here based on the following: Canada, BC, Osoyoos, Haynes Ecol. Res., pitfall traps, Artemisia-Purshia assoc., 4.IV.1995/1.V.1995, 19. VIII.1999/16.IX.1999, 6.X.1994/4-IV.1995, 11. X.1998/11.IV.1999, 14.X.2001/27.IV.2002, coll. G.G.E. Scudder (UBC, LFC), 19 males, 29 females; Osoyoos, Haynes E.R., pitfall traps, recovery after fire, 6.X.1994/4.IV.1995, 1. V.1995/26.V.1995, 10.X.1997/12.IV.1998, 24. V.1999/30.VI.1999, 30.VI.1999/22.VII.1999, 11.X.1998/11.IV.1999, 9.X.1999/9.IV.2000, 11. V.2001/15.V.2001, 15.V.2001/17.VI.2001, 11. IV.2001/15.V.2001, 9.V.2001/15.V.2001, 14. X.2001/27.IV.2002, coll. G.G.E. Scudder (UBC, LFC), 50 males, 73 females; Osoyoos, 49.03
N, 119.31
W, pitfall traps, 11.IV.1999/27.V.1999, 10.X.1999/8.IV.2000, 10.IV.2001/14.V.2001, coll. G.G.E. Scudder (UBC, LFC), 16 males, 14 females; Osoyoos Desert Centre, pitfalls, 10. X.1999/8.IV.2000, coll. G.G.E. Scudder (UBC, LFC), 5 males, 8 females; BC, Vaseux Cr.,

11

Tribe Oxypodini C.G. Thomson, 1859

Kennedy flats, 49.15
N, 119.31
W, 2.V.1995/8. VI.1995, Purshia association, pitfall trap, coll. G.G.E. Scudder (LFC), 1 female. COLLECTION AND HABITAT DATA. Habitat: dry habitat, Artemisia-Purshia association, and habitat recovery after fire both in Okanagan Valley, BC. Collecting period: IV– X. Collecting method: pitfall traps.

Genus Neothetalia Klimaszewski, 2004 (Figs. 11.93–11.98) (Klimaszewski and Pelletier 2004; Klimaszewski et al. 2018) Diagnosis. Adults of the genus Neothetalia may be distinguished from those of other aleocharine genera by the following combination of characters: body narrowly elongate, subparallel, dark brown to nearly black with brown to rustbrown elytra; size medium to large, length 4.0–6.0 mm, integument moderately glossy and moderately densely punctate and pubescent, forebody with distinct isodiametric microsculpture; antennomere IV quadrate to elongate, VII–X slightly transverse; maxillary palpus with four palpomeres, apical one narrowly elongate and one-third to nearly one-half length of penultimate palpomere; galea expanding slightly beyond apex of lacinia, its apical lobe bearing dense short pubescence; apex of lacinia with row of closely spaced spines; labial palpus with three palpomeres; ligula slender, deeply bifid (N. nimia, N. pallitarsis) or slightly emarginated apically (N. smetanai, N. canadiana, and N. columbiana); mentum trapezoidal, apical margin slightly concave, with three long apical setae on each side; labrum transverse and usually slightly projecting medially; right mandible with distinct medial tooth; head quadrate or slightly elongate and distinctly narrower than pronotum; pronotum broadest medially and rather abruptly narrowing anteriorly but slightly so posteriorly, moderately to strongly transverse; pubescence along median line of disc directed anteriad except for very basal part, laterad on each side of disc and often forming broadly arcuate lines;

175

mesoventrite long, longer than maximum diameter of mesocoxa, with complete (N. pallitarsis) or partial median carina (remaining species), mesoventral process long, narrow, and acutely pointed, extending about two-thirds length of mesocoxal cavities, isthmus no more than one-fourth of maximum diameter of mesocoxa; elytra broader than pronotum, slightly transverse; abdomen with three basal tergites deeply impressed basally. MALE. Median lobe of aedeagus with tubus more or less narrowly elongate at apex in lateral view, bulbus approximately as long as tubus in lateral view, crista apicalis of bulbus broadly or narrowly oval, medium sized; internal sac with usually two pairs of inconspicuous subapical structures. FEMALE. Spermatheca with moderately sized, approximately spherical capsule bearing shallow or deep apical invagination, attachment of spermathecal gland located at capsule base near neck of spermatheca; basal, medial, and posterior portions of spermatheca consist of narrow tube-like stem, which is more or less broadly S-shaped or looped. Adults of this genus are associated with cold wet moss found alongside streams, on vertical slopes, on logs, and on and under rocks. They are cold-loving creatures and have been captured at elevations of up to 2133 m (Klimaszewski and Pelletier 2004). Key to Species of the Genus Neothetalia 1. Antennomere IV subquadrate (Figs. 11.94a and 11.98a); pronotum with edges strongly converging apically (Figs. 11.94a and 11.98a); median lobe of aedeagus and spermatheca as illustrated (Figs. 11.94b, c, h and 11.98b, c, h) ............................................................................2 – Antennomere IV slightly elongate; pronotum with edges moderately converging apically; median lobe of aedeagus and spermatheca differently shaped .......................................................3 2. Apical antennomere strongly elongate, at least as long as the two preceding antennomeres combined (Fig. 11.94a); pronotal edges slightly converging basally from basal third (Fig. 11.94a);

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median lobe of aedeagus and spermatheca as illustrated (Fig. 11.94b, c, h)................................ ........... Neothetalia columbiana (Klimaszewski) – Apical antennomere moderately elongate, shorter than the two preceding antennomeres combined (Fig. 11.98a); pronotal edges strongly converging basally from basal third (Fig. 11.98a); median lobe of aedeagus and spermatheca as illustrated (Fig. 11.98b, c).................................... ...................Neothetalia smetanai Klimaszewski 3. Pronotum dark brown to virtually black (Fig. 11.96a); elytra 1.2 times as long as pronotum (Fig. 11.96a); tubus of median lobe of aedeagus sinuate in lateral view (Fig. 11.96b) and sharply pointed in dorsal view (Fig. 11.96c); spermatheca strongly sinuate, stem broadly looped (Fig. 11.96h)....... Neothetalia pallitarsis (Kirby) – Pronotum brown with rust tinge (Figs. 11.93a, 11.97a, and 11.95a); elytra 1.2–1.4 times as long as pronotum (Figs. 11.93a, 11.97a, and 11.95a); tubus of median lobe of aedeagus and spermatheca not as above ..............................................4 4. Pronotum subquadrate (Fig. 11.97a); elytra at suture about as long a pronotum (Fig. 11.97a); median lobe of aedeagus with evenly arcuate tubus and moderately broad, triangular apex in lateral view (Fig. 11.97b)..................................... .........Neothetalia robergei Klimaszewski. sp. n. – Pronotum transverse (Figs. 11.93a and 11.95a); elytra at suture longer than pronotum (Figs. 11.93a and 11.95a); median lobe of aedeagus differently shaped ...............................5 5. Elytra 1.4 times as long as pronotum (Fig. 11.95a); body length 4.0–6.0 mm; pronotal sides moderately converging apically (Fig. 11.95a); ligula deeply emarginated; median lobe of aedeagus with narrowly elongate crista apicalis of bulbus in lateral view (Fig. 11.95b), and tubus without apical lateral projections in dorsal view (Fig. 11.95c); spermatheca with stem narrowly looped (Fig. 11.95h)..................... ...................................Neothetalia nimia (Casey) – Elytra 1.2 times as long as pronotum (Fig. 11.93a); body length 4.8–5.5 mm; pronotal sides strongly converging apically (Fig. 11.93a);

11

Tribe Oxypodini C.G. Thomson, 1859

ligula shallowly emarginated; median lobe of aedeagus with short and oval crista apicalis of bulbus in lateral view (Fig. 11.93b), and tubus with two apical lateral projections in dorsal view (Fig. 11.93c); spermatheca with stem broadly looped (Fig. 11.93h)............................................. ................ Neothetalia canadiana Klimaszewski

60. Neothetalia canadiana Klimaszewski (Fig. 11.93a–h) Neothetalia canadiana Klimaszewski, in Klimaszewski and Pelletier (2004), Klimaszewski et al. (2018). Diagnosis. Body length 4.8–5.5 mm, dark brown with elytra, basal antennomere, tarsi, and sometimes pronotum light brown with rust tinge, forebody moderately glossy and abdomen strongly so (Fig. 11.93a); antenna moderately stout, antennomere IV strongly elongate in both sexes, antennomeres V–X strongly elongate in male and slightly so in female, terminal antennomere about as long as two preceding ones combined; pronotum slightly transverse (Pronotal Index ¼ 1.2), lateral edges slightly converging basally in basal half and moderately strong in apical half, length ratio of base to apex 1.5 times, pronotal punctures sparse, distance between punctures about three times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad forming arcuate lines on both sides of disc; elytra transverse (Elytral Index ¼ 1.5), 1.2 times as long as pronotum. MALE. Tergite VIII transverse, broadly arcuate apically (Fig. 11.93d); sternite VIII slightly transverse, apical part broadly triangularly produced (Fig. 11.93e); median lobe of aedeagus with small, oval crista apicalis at base of bulbus, tubus long, approximately straight medially, moderately produced ventrally at apex, apex narrowly rounded in lateral view (Fig. 11.93b), a subapical hemispherical carina occurs on each side of tubus and is visible in dorsal view (Fig. 11.93c), internal sac with two strong subapical structures (Fig. 11.93b, c). FEMALE. Tergite

11

Tribe Oxypodini C.G. Thomson, 1859

VIII similar to that of male (Fig. 11.93f); sternite VIII transverse and broadly rounded apically (Fig. 11.93g); spermatheca (Fig. 11.93h) with capsule approximately spherical and slightly compressed, bearing short apical invagination, stem forming characteristically broadly looped tube. DISTRIBUTION. Origin: Nearctic. CANADA: BC, QC, YT. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: adults have been captured from gopher burrows, on carrion, and by sifting forest debris. The Yukon specimen was captured in a white spruce-feathermoss forest. Collecting period: V–VII. Collecting method: unbaited pitfall traps. 61. Neothetalia columbiana (Klimaszewski) (Figs. 11.94a–h) Ocalea columbiana Klimaszewski, in Klimaszewski and Winchester 2002, Klimaszewski and Pelletier 2004. Diagnosis. Body length 4.3–5.5 mm, dark brown with elytra, basal antennomere, tarsi, and sometimes pronotum light brown with rust tinge, forebody moderately glossy and abdomen strongly so (Fig. 11.94a); antenna stout, antennomere IV moderately elongate, IV slightly elongate, antennomeres VI–X quadrate to slightly transverse in male and distinctly transverse in female, and terminal antennomere about as long as two and a half preceding ones combined; pronotum strongly transverse (Pronotal Index ¼ 1.3), lateral edges slightly converging basally in basal half and strongly so in apical half, length ratio of base to apex about 1.6 times, pronotal punctures sparse, distance between punctures about 2–3 times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad on both sides; elytra transverse (Elytral Index ¼ 1.5). MALE. Tergite VIII truncate apically and without

177

median emargination (Fig. 11.94d); sternite VIII slightly transverse, apical part broadly triangularly produced, apex pointed (Fig. 11.94e); median lobe of aedeagus with small, oval crista apicalis at base of bulbus, tubus long, approximately straight medially, moderately produced ventrally at apex, apex narrowly rounded in lateral view (Fig. 11.94b); median lobe in dorsal view as illustrated (Fig. 11.94c). FEMALE. Tergite VIII truncate apically (Fig. 11.94f); sternite VIII transverse and broadly rounded apically (Fig. 11.94g); spermatheca similar to that of P. canadiana, capsule approximately spherical and slightly compressed, with short apical invagination, spermathecal stem forming narrow, characteristically broadly looped tube (Fig. 11.94h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002; Klimaszewski and Pelletier 2004). Habitat: The Carmanah (Vancouver Island) specimens were captured in the forest interior and the transition zone of the Sitka spruce dominated forest. Some specimens were found on rotting piles of mushrooms and under deciduous shrubs around bases of cedar. One female was taken from a dung and carrion trap. One male was captured in the transition zone of Douglas-fir (Pseudotsuga menziesii (Mirb.)) dominated forest in west Vancouver, and the remaining two males were found in the Kenai Mountains of Alaska. Collecting period: V–VIII. Collecting method: pitfall traps, dung trap, Malaise and pan traps. 62. Neothetalia nimia (Casey) (Fig. 11.95a–h) Atheta (Athetalia) nimia Casey 1910. As Atheta (Hypatheta): Bernhauer and Scheerpeltz 1926. As Ischnopoda (Stethusa) metlakatlana: Hatch 1957. As Atheta (Stethusa): Moore and Legner 1975. As Athetalia: Campbell and Davies 1991. As Ocalea: Klimaszewski and Winchester 2002. As Neothetalia: Klimaszewski and Pelletier 2004.

178

Diagnosis. Body length 4.0–6.0 mm, dark brown to nearly black with elytra, basal antennal segment, tarsi, and often pronotum light brown with rust tinge, forebody moderately glossy and abdomen strongly so (Fig. 11.95a); antennae slender, antennomere IV strongly elongate, V–VII slightly elongate, VIII–X quadrate in male and slightly transverse in female, terminal antennomere about as long as two preceding ones combined; pronotum slightly transverse (Pronotal Index ¼ 1.2), lateral edges slightly converging basally and strongly so in apical half, length ratio of base to apex 1.4 times, pronotal punctures sparse, distance between punctures about 2–3 times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad forming approximately arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.5), about 1.4 times as long as pronotum. MALE. Tergite VIII transverse, truncate apically and bearing shallow median emargination (Fig. 11.95d); sternite VIII with apical part triangularly produced, apex pointed (Fig. 11.95e); median lobe of aedeagus with long and narrowly crista apicalis at the base of bulbus, tubus approximately straight medially, strongly produced ventrally at apex, apex narrow and sharply pointed in lateral view (Fig. 11.95b); tubus in dorsal view as illustrated (Fig. 11.95c). FEMALE. Tergite VIII similar to that of male (Fig. 11.95f); sternite VIII broadly triangular posteriorly, apex rounded (Fig. 11.95g); spermatheca with capsule small, spherical, bearing deep and narrow invagination facing downwards, stem S-shaped with broad posterior loop (Fig. 11.95h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. New locality data: British Columbia, W of Terrace, Shames Mtn., road to skee area, 52
28.140 N, 128
56.050 W, 320 m, 29.VII.2012, in moss and alder litter, A. Davies (CNC) 3 males, 22 sex undetermined; Kitsumkalum Lake at Hwy. 13, S of Rosswood, 54
47.340 N, 128
45.970 W, 146 m, 30.VII.2012, in moss and leaf litter at

11

Tribe Oxypodini C.G. Thomson, 1859

creek, A. Davies (CNC) 2 females, 2 sex undetermined. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002; Klimaszewski and Pelletier 2004). Habitat: Specimens have been captured in sifted forest litter; in cold wet moss on rocks, on slopes, along streams, and in the splash zone; on dung, carrion, and rotting mushrooms. This species occurred mainly in samples with cold wet moss in the interior of Sitka spruce (Picea sitchensis (Bong.) Carrière) dominated ancient forest on Vancouver Island. Collecting period: VI–IX. Collecting method: sifting forest litter, pitfall traps, pan traps, and Malaise traps. 63. Neothetalia 11.96a–h)

pallitarsis

(Kirby) (Fig.

Aleochara pallitarsis Kirby 1837. As Atheta: Bernhauer and Scheerpeltz 1926; Moore and Legner 1975. As undescribed genus near Ocalea: Klimaszewski 1984. As Neothetalia: Klimaszewski and Pelletier 2004. Diagnosis. Body length 4.7–6.0 mm, dark brown to nearly black with elytra, basal antennal segment, and tarsi light brown with rust tinge, forebody moderately glossy and abdomen strongly so (Fig. 11.96a); antennae moderately stout, antennomere IV slightly elongate, V–VII quadrate in male and transverse in female, VIII–X quadrate to slightly elongate in male and transverse in female, terminal antennomere about as long as two preceding ones combined; pronotum slightly transverse (Pronotal Index ¼ 1.2), lateral edges slightly converging basally in basal half and strongly so in apical half, length ratio of base to apex 1.6 times, pronotal punctures sparse, distance between punctures about three times diameter of a puncture, space between punctures with strong isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad forming slightly arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.6). MALE. Tergite VIII truncate apically and bearing

11

Tribe Oxypodini C.G. Thomson, 1859

shallow median emargination (Fig. 11.96d); sternite VIII transverse, apical part broadly triangularly produced posteriorly, apex slightly pointed (Fig. 11.96e); median lobe of aedeagus with short crista apicalis at base of bulbus, tubus slightly sinuate and strongly produced ventrally in lateral view (Fig. 11.96b), median lobe with internal sac bearing strongly sclerotized, reversed narrowly V-shaped structure in dorsal view (Fig. 11.96c). FEMALE. Tergite VIII similar to that of male (Fig. 11.96f); sternite VIII broadly rounded posteriorly (Fig. 11.96g); spermathecal capsule subspherical and with short apical invagination, stem forming narrow tube of a more or less semicircular shape, median portion of stem variably curved (Fig. 11.96h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: in forest litter at altitudes 122–2133 m. Collecting period: V–IX. Collecting method: sifting forest litter, pitfall traps. 64. Neothetalia robergei Klimaszewski, sp.n. (Fig. 11.97a–h) Holotype (male): Canada, British Columbia, Vancouver Island, 4 km SW Bamberton, Spectacle Lake, 48
340 4100 N, 123
340 1900 W, 7.X.2011, moss on Douglas fir logs in second growth forest, A. Davies (CNC). Paratypes: label data same as the holotype (CNC) 6 females. Etymology. This species is dedicated to Sylvain Roberge, an extraordinary artist, who provided many images for species included in this and the previous book on aleocharine beetles in Canada. Diagnosis. Body length 5.7–6.0 mm, head, posterior part of abdomen and antennomeres III–XI dark brown, pronotum reddish brown, and remainder of the body yellow brown; forebody

179

strongly glossy and abdomen more so (Fig. 11.97a); antennae slender, antennomere IV distinctly elongate, V–VII slightly transverse, VIII–X moderately transverse; pronotum subquadrate (Pronotal Index ¼ 1.06), lateral edges slightly converging basally in basal half and more so in apical half, length ratio of base to apex 1.2 times, pronotal punctures sparse, distance between punctures about three times the diameter of a puncture, space between punctures with strong isodiametric microsculpture, pubescence directed anteriad along median line of disc, except at base of disc, and laterad on both sides; elytra slightly transverse (Elytral Index ¼ 1.2). MALE. Tergite VIII truncate apically and bearing shallow median emargination (Fig. 11.97d); sternite VIII with apical part triangularly produced posteriorly with pointed apex (Fig. 11.97e); median lobe of aedeagus with moderately sized crista apicalis at base of bulbus, tubus evenly arcuate and moderately produced ventrally, apex narrowly triangular in lateral view (Fig. 11.97b), in dorsal view bulbus oval and tubus subparallel basally and triangular apically, sides slightly sinuate (Fig. 11.97c); internal sac with two narrow, basally converging structures in bulbus, and several prominent elongate apical structures in tubus (Fig. 11.97b, c). FEMALE. Tergite VIII truncate apically and slightly emarginated medially (Fig. 11.97f); sternite VIII rounded triangularly produced apically (Fig. 11.97g); spermathecal capsule small, spherical and with short apical invagination, stem forming narrow sinuate tube with broad posterior loop (Fig. 11.97h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: moss on Douglas fir logs in second growth forest. Collecting period: X. Collecting method: sifting moss. 65. Neothetalia (Fig. 11.98a–h)

smetanai

Klimaszewski

180

Neothetalia smetanai Klimaszewski, Klimaszewski and Pelletier (2004).

11

in

Diagnosis. Body length 5.2–5.6 mm, dark brown to nearly black with elytra and sometimes pronotum paler, basal antennal segment and tarsi brown with rust tinge, forebody moderately glossy and abdomen strongly so (Fig. 11.98a); antennae moderately stout (Fig. 11.71a), antennomere IV subquadrate to slightly elongate, V–VII quadrate to slightly transverse, VIII–X slightly transverse in male to more so in female, terminal antennomere about as long as two preceding ones combined; pronotum slightly transverse (Pronotal Index ¼ 1.2), often bearing slight elongate impression along median line of disc, lateral edges strongly converging basally and apically from about middle of disc, length ratio of base to apex 1.2 times, punctures sparse, distance between punctures from two to three times diameter of a puncture, space between punctures with strong, isodiametric microsculpture, pubescence directed anteriad along median line of disc and laterad forming approximately arcuate lines on both sides; elytra transverse (Elytral Index ¼ 1.4). MALE. Tergite VIII arcuate apically (Fig. 11.98d); sternite VIII with apical part narrowly triangularly produced (Fig. 11.98e); median lobe of aedeagus with short and narrowly elongate crista apicalis at base of bulbus, tubus approximately straight medially, strongly produced ventrally at apex, apex short, triangular, moderately broad and pointed in lateral view (Fig. 11.98b); median lobe in dorsal view as illustrated (Fig. 11.98c). FEMALE. Tergite VIII arcuate apically (Fig. 11.98f); sternite VIII rounded posteriorly (Fig. 11.98g); spermathecal capsule small, spherical, bearing deep and narrow invagination facing downwards, stem S-shaped with bends slightly angular (Fig. 11.98h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

Tribe Oxypodini C.G. Thomson, 1859

COLLECTION AND HABITAT DATA (Klimaszewski and Pelletier 2004). Habitat: in forest litter at altitudes 1036–1371 m. Collecting period: V, VI. Collecting method: not recorded.

Genus Ocalea Erichson, 1837 (Fig. 11.99a–h) (Klimaszewski and Pelletier 2004) Diagnosis. Body narrowly elongate, elytra and abdomen subparallel, medium sized, length 4.0–4.4 mm, dark brown with paler and slightly rust brown pronotum, elytra, apical part of abdomen, and appendages; integument strongly glossy, moderately densely punctate and pubescent, distance between punctures about two times diameter of a puncture, forebody with weak, scarcely visible microsculpture; head narrowly produced in front of eyes and rounded behind eyes; antenna: antennomeres IV–VII strongly elongate, VIII–X slightly elongate, and terminal antennomere about as long as two preceding ones combined; ligula slender, shallowly split apically; terminal palpomere of labial and maxillary palps without pseudosegments; terminal palpomere of maxillary palpus as long as about one-half length of penultimate one; labrum with apical margin slightly produced medially; pronotum gradually broadening to about one-third to one-half of its length and then strongly narrowed apically, broadly club-shaped, distinctly narrower than elytra, and slightly transverse, broadest in apical third, strongly converging apically and slightly so basally, slightly impressed basally, punctation moderately large and moderately dense, distance between punctures 2–3 times diameter of a puncture, pubescence directed anteriorly along arcuate lines from midline of disc; mesoventrite with partial carina approximately as long as one-fourth length of mesoventrite, mesoventral process narrowly triangular, sharp apically, and wedging to about one-half length of mesocoxae, distance between apex of mesoventrite and metaventrite long and about one-third of mesocoxal length; mesocoxae contiguous

11

Tribe Oxypodini C.G. Thomson, 1859

181

medially. MALE. Median lobe of aedeagus with short, narrowly oval crista apicalis of bulbus, tubus sinuate toward apex in lateral view, apical part broad, elongate and rounded, tubus moderately broad and subparallel in dorsal view, slightly narrower than bulbus; internal sac with two small subapical structures. FEMALE. Spermatheca with capsule approximately spherical, slightly compressed and with shallow and narrow apical invagination, stem narrowly elongate, sinuate (posterior part damaged in only known female syntype). Bionomics not available for Nearctic species. Palearctic species are affiliated with wet moss near water and particularly in the splash zone near waterfalls (Lohse 1974 and label data for O. picata). 66. Ocalea vancouveri Casey (Fig. 11.99a–h) Ocalea vancouveri Casey, Klimaszewski and Pelletier, 2004.

1893.

Diagnosis. Body length 4.0–4.4 mm, dark brown with paler pronotum, elytra, base of antennae, and tarsi; entire body strongly glossy (Fig. 11.99a); antennae slender in both sexes, antennomeres IV–X strongly elongate and terminal antennomere slightly shorter than two preceding ones combined; pronotum slightly transverse and distinctly narrower than elytra (Pronotal Index ¼ 1.1), broadest in about apical third, strongly converging apically and slightly so basally, impressed basally, punctures sparse, distance between punctures 2–3 times diameter of a puncture, space between punctures with weak microsculpture, pubescence directed anteriad along median line of disc and laterad, usually forming arcuate lines on both sides of disc; elytra slightly transverse (Elytral Index ¼ 1.4) and 1.4 times as long as pronotum. MALE. Tergite VIII broadly arcuate apically and sinuate laterally (Fig. 11.99d); sternite VIII with apical part rounded and slightly produced (Fig. 11.99e); median lobe of aedeagus with short, oval crista apicalis at base of bulbus, tubus long, broad, and

subparallel in dorsal view (Fig. 11.99c), sinuate and strongly produced ventrally in lateral view, apex broad and rounded, internal sac structures small and weakly sclerotized (Fig. 11.99b). FEMALE. Tergite and sternite VIII broadly rounded apically (Fig. 11.99f, g); spermatheca with capsule moderately large, approximately spherical, with short, narrow apical invagination, stem broken posteriorly in the only known female syntype (Fig. 11.99h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. Ocalea vancouveri is known only from the type locality in Victoria, Vancouver Island. It is the only species of the genus Ocalea in North America, and all remaining previous Ocalea records from this region should be considered erroneous, with species belonging to different genera (see Klimaszewski and Pelletier 2004). We have compared O. vancouveri with the type specimens of Ocalea picata (Stephens) (originally described as Aleochara), the type species for the genus Ocalea, and have found general agreement in their characteristics. Externally, O. picata has stronger microsculpture on the forebody; slightly longer pronotum, mesosternal carina, and mesosternal process; and strong macrosetae on sides of pronotum, elytra, and abdomen. These macrosetae were not observed in the two specimens of O. vancouveri, in which they may have been broken.

Genus Parocalea Bernhauer, 1902 (Fig. 11.100a–g) (Lohse Klimaszewski et al. 2018)

et

al.

1990;

Diagnosis. Body narrowly elongate, subparallel, head rounded, slightly narrower than pronotum; integument of forebody moderately glossy and with strong isodiametric microsculpture

182

11

Tribe Oxypodini C.G. Thomson, 1859

consisting of convex sculpticells, punctation fine; head and pronotum only slightly narrower than elytra; eyes small, infraorbital carina complete; antennae slightly incrassate, antennomere IV quadrate or slightly transverse, V–X distinctly transverse; pubescence on midline of disc of pronotum directed anteriad along anterior half to two-thirds, laterad on each side, and posteriad in basal one-third to half; mesocoxae contiguous or narrowly separated; elytra at suture slightly longer than pronotum along midline; abdomen subparallel, bases of first four visible tergites with deep transverse impressions; legs moderately long, basal tarsomere at least as long as two following ones combined, tarsal formula 5-5-5.

VIII arcuate apically (Fig. 11.100e); sternite VIII slightly produced at apex (Fig. 11.100f); spermatheca with capsule subspherical, stem irregularly coiled and twisted posteriorly (Fig. 11.100g). Parocalea pseudobaicalica differs from P. nearctica Lohse in having the pronotum subquadrate, the punctation in the impressions of the basal tergites denser, the tubus of the median lobe of the aedeagus evenly arcuate (tubus feebly sinuate in P. nearctica), and different structures in the internal sac.

67. Parocalea pseudobaicalica Lohse (Fig. 11.100a–g)

We record this species from BC for the first time based on the following: British Columbia, Atlin, Pine Creek, 21.VI.2015, 59.7116133.5787, 939 m, Alder soil sifting, coll. B. Godin and D. Horwood (BGC, LFC) 4 males, 2 females.

Parocalea pseudobaicalica Lohse, in Lohse et al. (1990). Diagnosis. Body medium-sized, narrowly linear, length 3.5–4.2 mm, colour brown or brownishblack, apex of abdomen paler, appendages light to dark brown (Fig. 11.100a); head one tenth narrower than length of pronotum, broadest in apical half; eyes small, each about half length of genae in dorsal view; antennae about as long as head and pronotum combined, antennomeres I–III elongate, IV quadrate, V–X transverse and incrassate; pronotum subquadrate, broadest in apical half, hind angles obtuse; elytra at suture as long as pronotum along midline and 1 1/3 as broad as pronotum; abdomen subparallel; antennae with basal antennomere as long as antennomeres II–IV combined. MALE. Tergite VIII truncate apically (Fig. 11.100c); sternite VIII triangularly produced apically (Fig. 11.100d); median lobe of aedeagus in dorsal view broad, narrowly oval, with two pairs of elongate sclerites; in lateral view bulbus moderately large, with large and oval crista apicalis, tubus with ventral margin strongly arched ventrad in lateral view (Fig. 11.100b). FEMALE. Tergite

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific provincial record], NT, QC, YT. USA: AK, NH.

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: near Salix, Larix, and Picea trees, edges of standing or running water, under stones on silt and gravel, among grasses, moss, and Ranunculus. The BC specimens were found in alder soil. Collecting period: VI–VIII. Collecting method: sifting forest litter and soil. Subtribe Phloeoporina C.G. Thomson, 1859 (Ashe 2001; Osswald et al. 2013) Diagnosis. Body slender, subparallel, convex, colour brown to black, forebody with pubescence short and dense; head nearly as broad as pronotum or broader, strongly constricted basally, with distinct neck, eyes as long as or slightly shorter than genae in dorsal view, postocular carina absent; antennomeres V–X moderately to distinctly transverse; pronotum angular, subquadrate to slightly transverse, pubescence on disc directed obliquely inward basally, straight posteriad elsewhere; elytra broader than

11

Tribe Oxypodini C.G. Thomson, 1859

pronotum, posterior margin distinctly sinuate; first three to four visible abdominal tergites with basal impressions deep, transverse, with very coarse punctation. Comments. Osswald et al. (2013), based on molecular studies, considered Phloeopora to be a separate lineage with unresolved relationships to the other clades of Oxypodini, and based on inferred phylogeny they treated Phloeopora as the only member of the subtribe Phloeoporina.

Genus Phloeopora Erichson, 1837 (Figs. 11.101–11.104) (Lohse et al. 1990; Ashe 2001; Klimaszewski et al. 2018) Diagnosis. Body slender, subparallel, convex, length 2.2–3.5 mm; colour brown to black, forebody with pubescence short and dense; head nearly as broad as pronotum or broader, strongly constricted basally, with distinct neck, eyes as long as or slightly shorter than genae in dorsal view, postocular carina absent; antennomeres V– X moderately to distinctly transverse; pronotum angular, subquadrate to slightly transverse, pubescence on disc directed obliquely inward basally, straight posteriad elsewhere; elytra broader than pronotum, posterior margin distinctly sinuate; first three to four visible abdominal tergites with basal impressions deep, transverse, with very coarse punctation; male sternite VIII triangularly produced apically; median lobe of aedeagus narrow, tubus subparallel, bulbus oval and small; spermatheca with capsule spherical, with invagination deep, moderately broad, stem sinuate with posterior coils and loops. Comments. All species of this genus are externally similar to each other and differ in small details such as body proportions (e.g., width of pronotum, length of elytra), density of punctation and pubescence on forebody, body colour, shape of the apical part of the median lobe of the aedeagus in lateral view, and shape of the spermatheca.

183

Key to Species of Phloeopora 1. Body broad, head and elytra distinctly broader than pronotum (Fig. 11.101a); forebody with pubescence long and sparse; integument glossy; median lobe of aedeagus with apex of tubus thin and moderately strong produced ventrally in lateral view (Fig. 11.101b); spermathecal stem with several broad coils and loops (Fig. 11.101h)....... ................................... Phloeopora arctica Lohse – Body narrowly subparallel, head at most as broad as pronotum and elytra slightly broader than pronotum (Figs. 11.102a, 11.103a, and 11.104a); forebody with pubescence short and dense; integument less glossy; median lobe of aedeagus with apex of tubus moderately thin apically and strongly produced ventrally in lateral view (Figs. 11.102b, 11.103b, and 11.104b); spermathecal stem with a few tighter coils and loops (Figs. 11.102h and 11.103g) ....................2 2. Forebody with pubescence moderately dense; integument highly glossy (Fig. 11.102a); male sternite VIII strongly triangularly produced with apex pointed (Fig. 11.102e); median lobe of aedeagus arcuate with apex of tubus strongly produced ventrally in lateral view (Fig. 11.100b); body length usually less than 3.0 mm.................. Phloeopora canadensis Klimaszewski and Langor – Forebody with pubescence dense; integument slightly glossy (Figs. 11.103a and 11.104a); male sternite VIII moderately triangularly produced with apex rounded (Figs. 11.103d and 11.104d); median lobe of aedeagus with apex of tubus straight and strongly produced ventrally in lateral view (Figs. 11.103b and 11.104b); body length usually greater than 3.0 mm................................. ................................. Phloeopora oregona Casey

68. Phloeopora arctica Lohse (Fig. 11.101a–h) Phloeopora arctica Lohse, in Lohse et al. (1990). Majka and Klimaszewski 2008a, Klimaszewski et al. 2018.

184

Diagnosis. Body moderately broad, elongate, length 3.0–3.5 mm, colour dark brown with tarsi, maxillary palpi, labrum and antennae reddish-brown (Fig. 11.101a); integument moderately glossy, moderately densely punctate and pubescent, pubescence long, adhering to body; head slightly wider and longer than pronotum, pubescence on disc directed inward from sides to midline; eyes large, slightly shorter than genae in dorsal view; antennae slightly incrassate, antennomeres V–X distinctly transverse; pronotum trapezoidal in shape, broadest subapically, ¼ narrower than width of elytra, pubescence on disc directed almost straight posteriad; elytra elongate, sides arcuate, with shoulders distinct, narrowly rounded, pubescence on disc directed obliquely and straight posteriad; abdomen subparallel with first three visible tergites deeply impressed basally MALE. Tergite VIII with apical margin very broadly arcuate medially, sinuate laterally (Fig. 11.101d); sternite VIII with apical margin strongly, narrowly triangularly produced, rounded at apex (Fig. 11.101e); median lobe of aedeagus in lateral view with ventral margin of tubus curved obliquely ventrad near base, thin and slightly curved ventrad apically (Fig. 11.101b), in dorsal view bulbus round and tubus narrowly subparallel (Fig. 11.101c). FEMALE. Tergite VIII broadly arcuate apically (Fig. 11.101f); sternite VIII with apical margin arcuate (Fig. 11.101g); spermatheca with capsule spherical, acutely bent laterad at base, stem with several broad, irregular loops (Fig. 11.101h). This species differs from its Nearctic congeners in having the body broad, the colour uniformly dark brown except for the appendages, and the shapes of the median lobe of the aedeagus in lateral view and the spermatheca are distinctive.

11

Tribe Oxypodini C.G. Thomson, 1859

DISTRIBUTION. Origin: Nearctic. CANADA: AB and BC [new generic and specific records], NT, ON, YT. USA: not recorded. We record this species from BC for the first time based on the following: British Columbia, BColl, Hwy 16 W of Fishpan Lake, N of Barrett, 54
300 6600 N, 126
450 3700 W, 834 m, 26.VII.2012, log pile, in Pinus contorta bark, A. Davies (CNC) 1 male, 2 females. Alberta: Whitney Cr., 10. VII.1985, ex Dendroctonus ponderosae gallery in Lodgepole pine, D. Langor (CNC) 1 male. COLLECTION AND HABITAT DATA. Habitat: one specimen sifted from litter, otherwise nothing is known about the habitat association. In AB, one male was found in mountain pine beetle (Dendroctonus ponderosae (Hopkins)) gallery in Lodgepole pine. In BC, specimens were collected from log pile, in Pinus contorta Douglas bark. Collecting period: VII, VIII. Collecting method: sifting litter, hand picking in bark. 69. Phloeopora canadensis and Langor (Fig. 11.102a–h)

Klimaszewski

Phloeopora canadensis Klimaszewski and Langor, in Klimaszewski et al. 2011. Bousquet et al. 2013, Webster et al. 2016, Klimaszewski et al. 2018. Diagnosis. Body narrowly elongate, length 2.4–3.0 mm, subparallel, colour brown with head dark brown and antennae, legs, palps and apical part of abdomen light brown with rustybrown tinge (Fig. 11.102a); integument moderately glossy with meshed microsculpture; head as wide and as long as pronotum; pronotum 1/5 narrower than elytra; antennomeres V–X moderately transverse; elytra narrow, arcuate laterally, slightly longer than pronotum; abdomen subparallel, slightly narrower than elytra. MALE. Tergite VIII arcuate apically (Fig. 11.102d); sternite

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Tribe Oxypodini C.G. Thomson, 1859

VIII with apical margin strongly triangularly produced, slightly emarginate on each side of narrowly rounded apex (Fig. 11.102e); tubus of median lobe of aedeagus in lateral view broadly bent obliquely ventrad near base, slightly elongate and slightly curved ventrad subapically, apex narrowly rounded (Fig. 11.102b), in dorsal view bulbus oval and tubus subparallel (Fig. 11.102c). FEMALE. Tergite VIII broadly arcuate apically (Fig. 11.102f); sternite VIII with apical margin very obtusely produced, apex broadly rounded (Fig. 11.102g); spermatheca with capsule spherical, stem irregularly coiled posteriorly (Fig. 11.102h). Phloeopora canadensis is externally very similar to P. oregona Casey but has a narrower and smaller body. Phloeopora oregona has a more sinuate pronotal base, slightly longer elytra in comparison with the pronotum, and in lateral view the median lobe of the aedeagus has the apical part more elongate and the anterior part of the bulbus more angulate. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific record], NF, NB. USA: not recorded. We record this species from BC for the first time based on the following: British Columbia, BColl, Hwy 16 W of Fishpan Lake, N of Barrett, 54
30.66’N, 126
45.37’W, 834 m, 26.VII.2012, log pile, in Pinus contorta bark, A. Davies (CNC) 3 males, 3 females, 9 sex undetermined; BC, Liard Riv., Hot Springs, 2.VI.1981, Picea sp., D.E. Bright (CNC) 1 male. COLLECTION AND HABITAT DATA. Habitat: most specimens from NB were captured in Lindgren funnel traps in conifer and mixed forests. These were an old black spruce forest, an old-growth white spruce and balsam fir forest, a mature red and white spruce forest, an old white pine stand, an old mixed forest, and a red oak forest adjacent to a black spruce stand. The only

185

specimens with microhabitat data were collected from under bark of red spruce infested with Dendroctonus in a red spruce stand. In NF, this species was found under the bark of tamarack, Larix laricina (Du Roi) K. Koch, recently killed by the eastern larch beetle, Dendroctonus simplex LeConte. In BC, specimens were collected from log pile, in Pinus contorta Douglas bark. Collecting period: V–VII, X. Collecting method: Lindgren funnel traps, aspirating from under bark. 70. Phloeopora oregona Casey (Figs. 11.103a– g and 11.104a–d) Phloeopora oregona Casey, 1906. Webster et al. 2012, 2016, Klimaszewski et al. 2018. Diagnosis. Body subparallel, narrowly elongate, length 2.9–3.3 mm, colour dark brown except tarsi and basal antennomeres reddish-brown; integument slightly glossy, densely and finely punctate and pubescent, pubescence short, dense, adhering to body (Figs. 11.103a (NB) and 11.104a (BC)); head as large as or slightly smaller than pronotum, pubescence on disc directed inward from sides to midline; eyes large, slightly shorter than genae in dorsal view; antennomeres V–X moderately transverse; pronotum trapezoidal in shape, broadest subapically, 1/5 narrower than elytra, pubescence on disc directed almost straight posteriad; elytra elongate, sides arcuate, with shoulders narrowly rounded, pubescence on disc directed obliquely and straight posteriad; abdomen subparallel with first three visible tergites deeply impressed basally. MALE. Tergite VIII slightly sinuate apically (Figs. 11.103c (NB) and 11.104c (BC)); sternite VIII with apical margin obtusely triangularly produced, slightly emarginate on either side of rounded apex (Figs. 11.103d (NB) and 11.104d (BC)); tubus of median lobe of aedeagus in lateral view bent obliquely ventrad at base, arcuately curved ventrad in apical half, slightly

186

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Tribe Oxypodini C.G. Thomson, 1859

elongate apically, apex narrowly rounded (Figs. 11.103b (NB) and 11.104b (BC)). FEMALE (NB and BC no differences). Tergite VIII sinuate apically and slightly produced medially (Fig. 11.103e); sternite VIII broadly rounded apically (Fig. 11.103f); spermathecal capsule spherical and with shallow apical invagination, stem long, strongly sinuate and looped posteriorly (Fig. 11.103g).

COLLECTION AND HABITAT DATA. Habitat: in NB, most specimens were collected from Lindgren funnel traps in old red pine forest, old jack pine forest, old white pine forest and old red spruce forests. Some were collected from under bark of red spruce and under bark of spruce in a wood pile in a mixed forest. Collecting period: IV–VII Collecting method: Lindgren funnel traps, aspirating from under bark.

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific record], NB. USA: OR.

Comments. The BC specimens are similar externally to those from NB (Fig. 11.103a–g) but have slightly broader pronotum, more lightly coloured posterior parts of elytra and slightly thinner apical part of tubus of median lobe of aedeagus. We attribute these small differences to geographic variability.

We record this species from BC for the first time based on the following: British Columbia, Summerland, 10.X.1932, A.N. Gartrell (CNC) 2 males, 2 females.

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.60 (a–g) Images of Blepharhymenus illectus (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

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sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

188

Fig. 11.61 (a–h) Images of Meotica pallens (Redtenbacher): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

11

Tribe Oxypodini C.G. Thomson, 1859

female tergite VIII; (f) female sternite VIII; (g, h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.62 (a–h) Images of Crataraea suturalis (Mannerheim): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

189

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

190

Fig. 11.63 (a–h) Images of Calodera bennetti Klimaszewski, sp. n.: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

11

Tribe Oxypodini C.G. Thomson, 1859

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–e), holotype; (f–h), paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.64 (a–h) Images of Devia prospera (Erichson): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

192

Fig. 11.65 (a–g) Images of Gnathusa eva Fenyes: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

11

Tribe Oxypodini C.G. Thomson, 1859

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.66 (a–g) Images of Gnathusa tenuicornis Fenyes: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

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male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

194

Fig. 11.67 (a–g) Images of Mniusa yukonensis (Klimaszewski and Godin): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.68 (a–g) Images of Ocyusa canadensis Lohse: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

195

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

196

Fig. 11.69 (a–h) Images of Oxypoda canadensis Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.70 (a–h) Images of Oxypoda convergens Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

197

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

198

Fig. 11.71 (a–h) Images of Oxypoda manitobae Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.72 (a–h) Images of Oxypoda opaca (Gravenhorst): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

200

Fig. 11.73 (a–h) Images of Oxypoda gatosensis Bernhauer: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

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Tribe Oxypodini C.G. Thomson, 1859

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.74 (a–h) Images of Oxypoda longicarinata Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

201

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

202

Fig. 11.75 (a–h) Images of Oxypoda irrasa Mäklin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

11

Tribe Oxypodini C.G. Thomson, 1859

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.76 (a–e) Images of Oxypoda famula Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) spermatheca; (c) female tergite VIII; (d) female sternite

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VIII; (e) pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 11.77 (a–i) Images of Oxypoda lacustris Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

11

Tribe Oxypodini C.G. Thomson, 1859

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.78 (a–h) Images of Oxypoda nimbata Casey: (a) habitus in dorsal view (apical part of abdomen and legs removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g)

205

female sternite VIII (partially damaged); (h) spermatheca (partially damaged). (b, c, h) modified from Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 11.79 (a–i) Images of Oxypoda frigida Bernhauer: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.80 (a–i). Images of Oxypoda orbicollis Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

207

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

208

Fig. 11.81 (a–h). Images of Oxypoda smithi Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.82 (a–h). Images of Oxypoda stanleyi Klimaszewski and McLean: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

209

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

210

Fig. 11.83 (a–h). Images of Oxypoda sylvia Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.84 (a–h). Images of Oxypoda vancouveri Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

211

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

212 Fig. 11.85 Images of Oxypoda glenorae Casey: habitus in dorsal view (apical part of abdomen removed). Scale bar for habitus ¼ 1 mm

11

Tribe Oxypodini C.G. Thomson, 1859

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.86 (a–d). Images of Oxypoda impressa Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) female tergite VIII; (c) female sternite

213

VIII; (d) pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

214

Fig. 11.87 (a–g). Images of Oxypoda regressa Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) male tergite VIII; (c) male sternite VIII; (d) male pygidium; (e) female tergite VIII; (f) female

11

Tribe Oxypodini C.G. Thomson, 1859

sternite VIII; (g) female pygidium. Median lobe of aedeagus and spermathecal missing. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.88 (a–h). Images of Alfocalea montana Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

215

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

216

Fig. 11.89 (a–h). Images of Betocalea pacifica Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.90 (a–i). Images of Megocalea lemieuxi Klimaszewski: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in ventral view; (e) male tergite VIII; (f) male

217

sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

218

Fig. 11.91 (a–i). Images of Metocalea lindgreni Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca; (i) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.92 (a–h). Images of Neoisoglossa agnita (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

219

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

220

Fig. 11.93 (a–h). Images of Neothetalia canadiana Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.94 (a–h). Images of Neothetalia columbiana (Klimaszewski and Winchester): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

221

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

222

Fig. 11.95 (a–h). Images of Neothetalia nimia (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.96 (a–h). Images of Neothetalia pallitarsis (Kirby): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

223

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

224

Fig. 11.97 (a–h). Images of Neothetalia robergei Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

11

Tribe Oxypodini C.G. Thomson, 1859

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b–e) holotype, (a, f–h) paratypes. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.98 (a–h). Images of Neothetalia smetanai Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

225

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) after Klimaszewski and Pelletier (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

226

Fig. 11.99 (a–h). Images of Ocalea vancouveri Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–e) lectotype, (f–h) paralectotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.100 (a–g). Images of Parocalea pseudobaicalica Lohse: (a) habitus in dorsal view; (b) male tergite VIII; (c) male sternite VIII; (d) male pygidium; (e) female tergite VIII; (f) female sternite VIII; (g)

227

female pygidium. (a–g) type specimens. Median lobe of aedeagus and spermathecal missing. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

228

Fig. 11.101 (a–h). Images of Phloeopora arctica Lohse: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

11

Tribe Oxypodini C.G. Thomson, 1859

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.102 (a–h). Images of Phloeopora canadensis Klimaszewski and Langor: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

229

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

230

Fig. 11.103 (a–g). Images of Phloeopora oregona Casey (NB specimens): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite

11

Tribe Oxypodini C.G. Thomson, 1859

VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

11

Tribe Oxypodini C.G. Thomson, 1859

Fig. 11.104 (a–d). Images of Phloeopora oregona Casey (BC specimen): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in

231

lateral view; (c) male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

232

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press, Boca Raton, London, New York, Washington, DC, 443 pp [publ. 28 Dec 2000 acccording to CRC Press] Assing V (2008) The genus Calodera Mannerheim in Canada (Insecta, Coleoptera, Staphylinidae, Aleocharinae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian coleoptera. ZooKeys [Special Issue] 2:203–208 Benick G, Lohse GA (1974) Tribus 14 (Callicerini) [pp 72–221]. In: Freude H, Harde KW, Lohse A (eds) Die Käfer Mitteleuropas. Band 5: Staphylinidae II (Hypocyphtinae und Aleocharinae), Pselaphidae. Goecke & Evers, Krefeld: 381 pp Bernhauer M (1902) Die Staphyliniden der paläarktischen Fauna. 1. Tribus: Aleocharini (part II). Verh kaiserkönigl zool-bot Ges Wien 52(Beiheft):87–284 Bernhauer M (1905) Neue Aleocharinen aus Nordamerika. Deutsch Entomol Z 1905(2):249–256 Bernhauer M (1907) Neue Aleocharini aus Nordamerika. (Col.) (3. Stück.). Deutsche Entomol Z 1907 (4):381–405 Bernhauer M, Scheerpeltz O (1926) Staphylinidae 6. In: Junk W, Schenkling S (eds) Coleopterorum Catalogus 82. W Junk, Berlin Blackwelder RE (1952) The generic names of the beetle family Staphylinidae, with an essay on genotypy. US Nat Museum Bull 200:I–IV, 1–483 Bousquet Y, Bouchard P, Davies AE, Sikes D (2013) Checklist of beetles (Coleoptera) of Canada and Alaska. Pensoft Publishers, Sofia-Moscow, 402 pp Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and Ecology of canadian Staphylinidae (Coleoptera) II. Zookeys [Special Issue] 186:119–206 Campbell JM, Davies A (1991) Family Staphylinidae rove beetles. pp 86–124. In: Bousquet Y (ed) Checklist of beetles of Canada and Alaska. Research Branch Agriculture Canada Publication 1861/E, vi + 430 pp Casey TL (1893) Coleopterological notices. V. Ann N Y Acad Sci 7[1892–1894]:281–606 [see reference in Chapter 7 for explanation of date] Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St. Louis 16(6):125–434 Casey TL (1910) New species of the staphylinid tribe Myrmedoniini [pp 1–183]. In: Memoirs on the Coleoptera. I. New Era Printing Co., Lancaster, PA, 205 pp

11

Tribe Oxypodini C.G. Thomson, 1859

Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, PA, 259 pp Downie NM, Arnett RH (1996) Introduction; Suborders Archostemata, Adephaga, and Polyphaga, thru Superfamily Cantharoidea. The beetles of northeastern North America. vol. I. The Sandhill Crane Press, FL, 880 pp Erichson WF (1837) Die Käfer der Mark Brandenburg. In: Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Erichson WF (1839) Genera et species Staphylinorum insectorum coleopterorum familiae. Erster Band. F. H. Morin, Berlin, viii + 400 pp Fenyes A (1910) Two new species of Aleocharinae from California. Proc Entomol Soc Wash 11[1909] (4):197–199 Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera Insectorum vol. XXVII, Fascicule 173A. M. Nijhoff, Den Haag, 453 pp + 7 pls Fenyes A (1921) New genera and species of Aleocharinae with a polytomic synopsis of the tribes. Bull Mus Comp Zool 65(2):17–36 Gouix N, Klimaszewski J (2007) Catalogue of aleocharine rove beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae). Pensoft, Sofia. 165 pp Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Gusarov VI (2002) A revision of Nearctic species of the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 81:1–88 Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353 (1):1–134 Hatch MS (1957) The Beetles of the Pacific Northwest. Part II: Staphyliniformia. Univer Wash Publ Biol 16: IX + 384 pp Hoebeke ER (1990) First record of the Palearctic species Oxypoda opaca (Gravenhorst) from North America (Coleoptera: Staphylinidae: Aleocharinae). J N Y Entomol Soc 97 [1989] (4):448–454 Horion A (1967) Faunistik der Mitteleuropäischen Käfer. Bd. XI: Staphylinidae. 3. Teil: Habrocerinae bis Aleocharinae (ohne Subtribus Athetae). P.C.V. Schmidt, Űberlingen – Bodensee, xxiv + 419 pp Kirby W (1837) The Insects. Coleoptera. In: Richardson J (ed), Fauna Boreali-Americana; or the zoology of the northern parts of British America. . . Part the fourth and last, J. Fletcher, Norwich, xxxix þ325 pp Klimaszewski J, Peck SB (1986) A review of the cavernicolous Staphylinidae (Coleoptera) of eastern North America: Part I. Aleocharinae. Quaest Entomol 22 (2):51–113

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Tribe Oxypodini C.G. Thomson, 1859

Klimaszewski J, Pelletier G (2004) Review of the Ocalea group of genera (Coleoptera, Staphylinidae, Aleocharinae) in Canada and Alaska: new taxa, bionomics, and distribution. Can Entomol 136 (4):443–500 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc belge d’Entomol 40:3–126 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48 Klimaszewski J, Pelletier G, Germain C, Work T, Hébert C (2006) Review of Oxypoda species in Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics, and distribution. Can Entomol 138(6):737–852 Klimaszewski J, Assing V, Majka CG, Pelletier G, Webster RP, Langor D (2007) Records of adventive aleocharine beetles (Coleoptera: Staphylinidae: Aleocharinae) found in Canada. Can Entomol 139 (1):54–79 Klimaszewski J, Langor D, Majka CG, Bouchard P, Bousquet Y, LeSage L, Smetana A, Sylvestre P, Pelletier G, Davies A, DesRochers P, Goulet H, Webster R, Sweeney J (2010) Review of adventive species of Coleoptera (Insecta) recorded from eastern Canada. Pensoft Series Faunistica No. 94. Pensoft, Sofia-Moscow, 272 pp Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Bourdon C (2012) Further contributions to the aleocharine fauna of the Yukon Territory, Canada (Coleoptera, Staphylinidae). In: Klimaszewski J, Anderson R (eds): Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. Zookeys [Special Issue] 186:207–237 Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013) Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia, 360 pp Klimaszewski J, Webster RP, Langor DW, Bourdon C, Hammond HEJ, Pohl GR, Godin B (2014) Review of Canadian species of the genera Gnathusa Fenyes, Mniusa Mulsant & Rey and Ocyusa Kraatz (Coleoptera, Staphylinidae, Aleaocharinae). ZooKeys 412:9–40 Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91

233 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. xvi þ 902 pp Kraatz G (1856) Staphylinii. Naturgeschichte der Insecten Deutschlands. Erste Abtheilung Coleoptera. Zweiter Band. Erste, zweite Lieferung. Bogen 1–24. Nicolai, Berlin, viii + 376 pp Lohse GA (1974) Hypocyphtinae, Aleocharinae [in part, pp 7–72, 221–292]. In: Freude H, Harde KW, Lohse A (eds) Die Käfer Mitteleuropas. Band 5: Staphylinidae II (Hypocyphtinae und Aleocharinae), Pselaphidae. Goecke & Evers, Krefeld, 381 pp Lohse GA, Smetana A (1985) Revision of the types of species of Oxypodini and Athetini (sensu Seevers) described by Mannerheim and Mäklin from North America (Coleoptera: Staphylinidae). Coleopt Bull 39 (3):281–300 Lohse GA, Klimaszewski J, Smetana A (1990) Revision of arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44(2):121–202 Majka C, Klimaszewski J (2008a) Adventive Staphylinidae (Coleoptera) of the Maritime Provinces of Canada: further contributions. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:151–174 Majka CG, and Klimaszewski J (2008b) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In: Majka CG, Klimaszewski J. (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys 2:85–114 Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39 Mäklin FW (1853) [New species and notes]. In: Mannerheim CG: Dritter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Imp Natural Moscou 26(3):95–273 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp. [separate, re-issued in Mémoires Présentées à l’Académie Impériale des Sciences de St.-Pétersbourg 1:415–501, in 1831] McLean JA, Klimaszewski J, Li A, Savard K (2009) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species. Part 1. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian coleoptera II. ZooKeys [Special Issue] 22:5–17 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514

234 Mulsant E, Rey C (1873) Description de divers coléoptères brévipennes nouveaux ou peu connus. Opuscules Entomol 15:147–189 Mulsant E, Rey C (1875) Histoire naturelle des Coléoptères de France: Brévipennes Aléochariens (suite). Myrmédoniaires (2eme partie). Deyrolle, Paris, [2] + 470 pp. + 4 pls Muona J (1991) The north European and British species of the genus Meotica Mulsant & Rey (Coleoptera, Staphylinidae). Deutsche Entomol Z 38(1–3):225–246 Osswald J, Bachmann L, Gusarov VI (2013) Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae). Syst Entomol 38 (3):507–522 Redtenbacher L (1849) Fauna Austriaca. Die Käfer. Nach der analytischen Methode bearbeitet. C. Gerold, Wien, pp i–xxvii + 641–883 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana Zool 71: vi + 289 pp Solier AJJ (1849) Zoología. Tomo cuarto. Insectos. Orden III. Coleópteros. [pp. 105–380, 414–511, pls. 1–12]. In: Gay C (ed): Historia física y política de Chile, Paris, 511 pp Thomson CG (1858) Försök till uppställning af Sveriges Staphyliner. Öfversi Kongli Vetensk-Akad Förhandl 15(1):27–40

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Tribe Oxypodini C.G. Thomson, 1859

Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom. Berlingska Boktryckeriet, Lund, [3] + 290 pp Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian coleoptera II. ZooKeys [Special Issue] 22:171–248 Webster RP, Klimaszewski J, Sweeney JD, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:83–118 Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016) Further contribution to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including description of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216

Tribe Tachyusini C. G. Thomson, 1859

Tribe Tachyusini C.G. Thomson, 1859 (Figs. 12.105–12.116) (Lohse 1989, Ashe 2001, Paśnik 2006, Osswald et al. 2013, Klimaszewski et al. 2018) Diagnosis. Body form diverse, from robust, to slender; head with neck more than one-third as wide as head or without distinct neck; pronotum without median sulcus, median elongate depression, or basal foveae; abdomen slightly narrower than elytra; or constricted at base; maxillae with galea and lacinia moderately elongate, length of galea usually shorter than, or subequal to, distance between base of galea to cardo; ligula Y-shaped, split at apex, with lobes diverging; mesocoxal cavities in most species moderately to narrowly separated or contiguous in some; process of metaventrite no longer than that of mesoventrite; tarsal formula 4-5-5.

12

2. Pronotal pubescence on disc directed straight posteriad and appearing velvet-like (Fig. 12.106a) ..............................Brachyusa Mulsant and Rey – Pronotal pubescence on disc directed anteriad, laterad or obliquely laterad from midline and not velvet-like, except slightly so in Paradilacra (Fig. 12.113a).....................................................3 3. Body in most strongly glossy and bluish metallic (Figs. 12.107a–12.112a); pubescence of forebody sparse (Figs. 12.107a–12.112a); head as wide as or wider than small pronotum; elytra markedly broader than pronotum (Figs. 12.107a– 12.112a) ....................... .Gnypeta C.G. Thomson – Body dull (Fig. 12.113a); pubescence of forebody dense (Fig. 12.113a); head about as wide as pronotum; elytra slightly wider than pronotum (Fig. 12.113a) ...................................... Paradilacra Bernhauer

Key to Genera of Tachyusini 1. Body robust (Figs. 12.106a and 12.107a– 12.113a); abdomen moderately narrower than elytra, not clavate and constricted at base; abdominal basal impressions without longitudinal ridges (Figs. 12.106a and 12.107a–12.113a)................2 – Body slender (Figs. 12.114a and 12.116a); abdomen clavate, constricted at base, and much narrower than elytra; abdominal basal impressions with longitudinal ridges (Figs. 12.114a and 12.116a) ................................................................. Tachyusa Erichson

Genus Brachyusa Mulsant and Rey, 1873 (Figs. 12.105–12.106) (Casey 1911 [as Tetralina], Seevers, 1978, Ashe 2001, Bousquet et al. 2013, Klimaszewski et al. 2011, 2016, 2018) Diagnosis. Body robust, elongate, arcuate laterally; integument covered with dense, fine, velvet-like pubescence, surface slightly glossy; head with genae short in dorsal view (only narrow strip visible behind eyes), eyes large, frons

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_12

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abruptly deflexed; clypeus strongly deflexed; antennomeres V–X elongate, subquadrate or transverse; terminal palpomere of maxillary palpus short and needle-shaped; pronotum trapezoidal in shape, slightly transverse; elytra slightly transverse, about as long as pronotum, posterior margin truncate and not emarginate laterally; abdomen moderately narrower than elytra, with first three visible tergites with basal impressions shallow; hind tarsus long, almost as long as hind tibia, basal tarsomere at least as long as next two combined; male sternite VIII extremely elongate; median lobe of aedeagus with tubus triangularly elongate, bulbus swollen, large, with large crista apicalis; spermatheca L- or S-shaped, capsule club-shaped, stem short and thick. Tarsal formula 4-5-5. Key to Species of Brachyusa 1. Tubus of median lobe of aedeagus in lateral view straight basally and bent ventrad apically, acutely tapering to apex (Fig. 12.105a)........ Brachyusa americana (Fenyes) – Tubus of median lobe of aedeagus in lateral view bent acutely ventrad basally, acutely tapering to apex (Fig. 12.106a)......................... Brachyusa helenae (Casey)

71. Brachyusa 12.105a, b) Brachyusa Seevers 1978.

americana

americana

(Fenyes) (Fig.

(Fenyes,

1921).

Diagnosis. Only one damaged specimen of this species is known. It is on loan to V. Gusarov and despite many efforts to examine it, it was not available for study. MALE [based on Seevers’ drawing]. Tubus of median lobe of aedeagus in lateral view short, straight basally and bent ventrad apically, acutely tapering to apex (Fig. 12.105a); paramere as illustrated (Fig. 12.105b).

Tribe Tachyusini C. G. Thomson, 1859

DISTRIBUTION. Origin: CANADA: BC. USA: not recorded.

Nearctic.

COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. The status of this species remains uncertain and the shape of the median lobe of aedeagus provided by Seevers (1978) is not typical for this genus. 72. Brachyusa helenae (Casey) (Fig. 12.106a– g) Tetralina helenae Casey, 1911. Klimaszewski et al. 2011, 2012, 2016, Brunke et al. 2012, Webster et al. 2012. Diagnosis. Body oval, slightly flattened, length 2.2–2.5 mm; colour predominantly black but sometimes brown, with appendages light brown; integument moderately glossy with pubescence short, silky (Fig. 12.106a); head narrower than elytra, with eyes large; antennomeres I–VII elongate, VIII–X quadrate to transverse; hind tarsus with basal article about as long as next two articles combined; pronotum wider than head but much narrower than elytra; elytra at suture longer than pronotum; abdomen strongly narrowed posteriorly. MALE. Tergite VIII truncate apically (Fig 12.106c); sternite VIII distinctly elongate, with apical margin broadly shallowly emarginate, lateral angles rounded (Fig. 12.106d); tubus of median lobe of aedeagus in lateral view short, bent acutely ventrad basally, acutely tapering to apex, apical triangular part varying in width (Fig. 12.106b); in dorsal view, basal half of median lobe broadly rounded/ swollen. FEMALE. Tergite VIII with apical margin very obtusely angulate (Fig. 12.106e); sternite VIII with apical margin broadly rounded on either side of shallow medial emargination (Fig.

12

Tribe Tachyusini C. G. Thomson, 1859

12.106f); spermatheca loosely S-shaped, with capsule elongate, bent at acute angle to sinuate stem (Fig. 12.106h). DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], LB, NB, NF, NT, ON, SK, YT. USA: AK, MT. We report this species from BC for the first time here based on the following: British Columbia, 8 mi W Creston, 10.VI.1968, A. Smetana (CNC) 1 female; BC, 12 mi E Hope, 2.VI.1968, ex river debris, Campbell and Smetana (CNC) 1 male; BC, Valemont, Yellowjacket ck, 3-8. VI.2014, pitfall, Kinbasket # 1050 &1051, 52.702-119.05, 758 m, C. wood removal, coll. Charlene Wood (BGC, LFC) 2 males; BC, Wynndel, Kootenay Lake, 14.VIII.1982, leg. R. Baranowski BWRS (ZMLU) 1 female; BCol., Toboggan Creek at Johnson Rd., N of Tobbogan Lk., 54
53.320 N, 127
15.790 W, 27. VII.2012, 480 m, in sedges along creek, A. Davies (CNC) 2 males, 7 females. New locality data: Northwest Territories, Martin Riv., 10 mi NW Ft. Simpson, 14.VI.1972, A. Smetana (CNC) 2 females; NWT, Inuvik, 24. VI.1972, A. Smetana (CNC) 1 sex?, ONT, Parry Sound Distr., 3 mi S Trout Cr., 18.IX.1974, I.M. Smith (CNC) 1 male. COLLECTION AND HABITAT DATA. Habitat: lives along shores of rivers and streams near flowing water. Adults in pine/spruce litter near stream, on sand and gravel on the banks of a river, near lake and river shorelines; on clay, sand, and gravel beaches, on sandy and silty river margins, in moss near the splash zone of a waterfall, in moss on rocks in the middle of a river, in gravel on a gravel bar along a shaded brook, and in gravel on a gravel/clay margin of a partially shaded brook, among cobblestones along a large shaded brook, in flood debris resting on a sand/ clay mix along an upper river margin, treading emergent Carex and grasses on the margin of a lake near the outflow of a stream and in debris on the margin of a vernal pond in a silver maple (Acer saccharinum L.) swamp near a river, and debris on a American beaver (Castor canadensis Kuhl) dam near the outflow area (flowing water)

237

from the dam; one specimen from debris within wall of beaver lodge (Klimaszewski et al. 2018). In BC, specimens were collected in sedges along creek, from river debris and coarse woody debris. Collecting period: IV–IX. Collecting method: pitfall traps, sifting litter near bodies of water, splashing water on moss, gravel, and sand or turning gravel and aspirating beetles. It took 5–10 min before adults appeared after repeated splashing of moss near splash zone of a waterfall.

Genus Gnypeta C.G. Thomson, 1859 (Figs. 12.107–12.112) (Klimaszewski et al. 2008, 2018) Diagnosis. Body robust, length 2.3–3.6 mm, integument moderately to highly glossy (often bluish metallic), finely punctate and pubescent, pubescence moderately long and silky in appearance in some; head and pronotum distinctly narrower than elytra; infraorbital carina partial or absent; ligula divided to base, or to near base, lobes parallel-sided; galea subequal in length to lacinia; pronotum broadest in apical one-third or at middle of disc, with sides converging apically and basally, pubescence on disc directed lateroposteriad from midline; elytra much broader than either head or pronotum, pubescence on disc often forming wavy pattern on each side; process of mesoventrite moderately broad, reaching middle of mesocoxae, its apex truncate; process of metaventrite broadly rounded; legs long and slender; tarsal formula 4-5-5; hind tarsus with basal article moderately elongate, usually shorter than two following ones combined; abdomen as broad as elytra at base or moderately narrower, subparallel, or broadening apically, first three visible tergites with deep basal impressions with coarse punctures but without pronounced longitudinal ridges; median lobe of aedeagus with tubus short, triangularly produced, bulbus enlarged, swollen, with crista apicalis large, subtriangular in shape laterally; paramere broad with short apical lobe bearing three long subapical and one short apical macrosetae; spermatheca of four types: S-shaped, C-shaped, hatchet-shaped, or club-shaped; capsule tubular with apical part roughly spherical, tubular, or funnel-shaped;

238

stem tubular, elongate, more-or-less sinuate and moderately swollen basally; sternite VIII of male and female with basal margin broadly separate from antecostal suture. Comments. Adults are associated with riparian habitats and debris along the margins of marshes, ponds, lakes, and streams. Some Canadian species were found in cold wet moss alongside streams. They may also occur in vegetation and litter along edges of streams, river, and lakes, in grass tussocks on mud flats, in gravel, wooded bogs, beaver lodges, and decaying fungi (Klimaszewski et al. 2008). This genus consists of mainly northern, cold-loving species, many of which have Holarctic distributions. Key to Species of Gnypeta 1. Elytra at suture approximately as long as pronotum (Figs. 12.110a, 12.111a and 12.112a) ............................................................................2 – Elytra at suture distinctly longer than pronotum (Figs. 12.107a, 12.108a and 12.109a) ............................................................................4 2. Antennomeres IV strongly transverse (Fig. 12.112a); median lobe of aedeagus and spermathecal as illustrated (Fig. 12.112b, c, h) ... Gnypeta brevicornis Casey – Antennomere IV elongate (Figs. 12.110a and 12.112a); genital characters not as above ..........3 3. Abdomen at base distinctly narrower than elytra at base (Fig. 12.111a); female sternite VIII with deep, narrow median emargination at apical margin of disc (Fig. 12.111g); spermatheca club-shaped (Fig. 12.111h); median lobe of aedeagus with sinuate apical part in lateral view (Fig. 12.110b).................... Gnypeta baranowskii Klimaszewski, sp. n. – Abdomen at base almost as wide as elytra at base (Fig. 12.110a); female sternite VIII with shallow, wide median emargination at apical margin of disc (Fig. 12.110g); spermathecal S-shaped (Fig. 12.110h); median lobe of aedeagus with straight apical part in lateral view

12

Tribe Tachyusini C. G. Thomson, 1859

(Fig. 12.110b).........................Gnypeta helenae Casey 4. Pronotum about two-fourth (0.6) as wide as elytra (Fig. 12.109a); median lobe of aedeagus in lateral view with apical part of tubus narrowly tapering apically (Fig. 12.109b); spermatheca with wide, U-shaped stem (Fig. 12.109h)....... Gnypeta crebrepunctata (Casey) – Pronotum about two-thirds (0.8) as wide as elytra (Figs. 12.107a and 12.108a); median lobe of aedeagus in lateral view with apical part of tubus broad and gradually tapering apically (Figs. 12.107b and 12.108b); spermatheca with thin and sinuate stem (Figs. 12.107h and 12.108h) ............................................................................5 5. Body surface with bluish metallic shine (Fig. 12.107a); antennomeres VIII–X subquadrate or slightly elongate (Fig. 12.107a); median lobe of aedeagus with broadly triangular and straight ventrally apex in lateral view (Fig. 12.107b); in dorsal view median lateral projections small (Fig. 12.107c); stem of spermatheca moderately sinuate (Fig. 12.107h).............. Gnypeta caerulea (C.R. Sahlberg) – Body surface without bluish metallic shine (Fig. 12.108a); antennomeres VIII–X slightly transverse (Fig. 12.108a); median lobe of aedeagus with narrowly triangular and sinuate ventrally apex in lateral view (Fig. 12.108b); in dorsal view median lateral projections large (Fig. 12.108c); stem of spermatheca strongly sinuate (Fig. 12.108h)............................................... Gnypeta lohsei Klimaszewski

Caerulea Species Group (Klimaszewski et al. 2008) Diagnosis. Male tergite VIII with two small lateral teeth at apical margin; spermatheca S-shaped, apical part of capsule mushroom-shaped, stem narrow and sinuate, slightly swollen basally; median lobe of aedeagus with short tubus broadly triangular apically in lateral view.

12

Tribe Tachyusini C. G. Thomson, 1859

73. Gnypeta caerulea (C.R. Sahlberg) (Fig. 12.107a–h) Aleochara caerulea C.R. Sahlberg 1831. Majka and Klimaszewski 2008, Klimaszewski et al. 2008, 2011, 2015, 2018. Diagnosis. Body length 3.0–3.5 mm; colour uniformly dark brown to black, sometimes middle of elytra, tarsi and sometimes entire legs rust brown; integument highly glossy with metallic reflection; pubescence yellowish-grey, moderately long and dense (Fig. 12.107a); antennomere IV distinctly elongate, V–X moderately to distinctly elongate, increasingly less so toward apex of antenna; head and pronotum of about same width; elytra and abdomen wider than either head or pronotum; head rounded posteriorly; pronotum about two-thirds (0.8) as wide as elytra, broadest in apical third, pubescence on disc directed anteriad along midline and laterad elsewhere; elytra at suture slightly longer than pronotum, pubescence on disc directed obliquely posterolaterad, forming wavy pattern medially on each side; abdomen subparallel, narrower than elytra at base, first three visible tergites with basal impressions deep, with large punctures; hind tarsus with two basal tarsomeres of about same length, third one slightly shorter. MALE. Tergite VIII transverse and truncate apically with two minute apical teeth (Fig. 12.107d); sternite VIII about as long as wide, rounded apically, a little more narrowly so at apex (Fig. 12.107e); tubus of median lobe of aedeagus in lateral view very short, broadly triangular apically, with ventral margin curved obliquely ventrad in basal half (Fig. 12.107b); in dorsal view bulbus moderately large with two small lateral projections (Fig. 12.107c); internal sac with structures as illustrated (Fig. 12.107b, c). FEMALE. Tergite VIII truncate apically (Fig. 12.107f); sternite VIII broadly rounded apically (Fig. 12.107g); spermatheca with capsule elongate, mushroom-shaped, stem sinuately club-shaped, gradually widening posteriad (Fig. 12.107h).

239

DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, LB, MB, NF, NS, NB, NT, ON, PE, QC, SK, YT. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: riparian and wetland habitats. In NB, adults collected from drift material along river margins, from gravel and under cobblestones on margins of cool, clear shaded brooks, treading emergent Carex and grasses on lake margin, sifted from leaf litter on beaver pond margin and from sticks and debris from outflow area of beaver dam, sifted from leaves and sedges from pond margin, and sifted from sphagnum from swamp edges. Elsewhere, collected from margins of streams, ponds, and lakes in moss, among rocks, gravel, sand, and driftwood, sweeping marsh vegetation, and sifted from vegetation at beaver pond. Collecting period: V–IX. Collecting method: splashing, treading, sifting litter, and turning gravel and cobblestones, and aspirating specimens, sweeping vegetation. Comments. The illustrations for G. caerulea (Klimaszewski et al. 2008: 44) and G. carbonaria (Klimaszewski et al. 2008: 57) were cited as “brincki” in the captions. 74. Gnypeta 12.108a–h)

lohsei

Klimaszewski (Fig.

Gnypeta lohsei Klimaszewski, Klimaszewski et al. 2008.

in

Diagnosis. Body length 2.7–3.0 mm; uniformly dark brown to black, sometimes central part of elytra and tarsi rust brown; integument moderately glossy, without bluish metallic reflection; pubescence yellowish grey and moderately long and dense (Fig. 12.108a); antennal article IV moderately elongate, V–VII subquadrate, VIII– X slightly transverse (Fig. 12.108a); head and pronotum of about the same width; elytra and

240

abdomen wider than either head or pronotum; head rounded posteriorly; pronotum broadest in apical third, about two-thirds (0.8) as wide as elytra, pubescence directed anteriad along midline and laterad elsewhere; elytra at suture slightly longer than pronotum, pubescence directed obliquely posterolaterad, in wavy pattern on medial parts of disc; abdomen subparallel, almost as broad as elytra at base; metatarsus with two basal tarsomeres of about the same length and the third one slightly shorter. MALE. Tergite VIII transverse and truncate apically with two minute apical teeth (Fig. 12.108d); sternite VIII as broad as long and parabolic apically, slightly pointed medially (Fig. 12.108e); median lobe of aedeagus with triangularly produced apical part of tubus and edges sinuate in lateral view (Fig. 12.108b); bulbus moderately enlarged with two anterior projections in dorsal view (Fig. 12.108c); internal sac with structures as illustrated (Fig. 12.108b, c). FEMALE. Tergite VIII truncate apically (Fig. 12.108f); sternite VIII broadly rounded posteriorly (Fig. 12.108g); spermatheca with capsule mushroom-shaped, elongate, stem sinuate and slightly swollen basally (Fig. 12.108h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: AK, WA. COLLECTION AND HABITAT DATA. Habitat: not recorded except high altitude collecting sites, up to 2300 m. Collecting period: VI–VIII. Collecting method: not recorded. Crebrepunctata Species Group (Klimaszewski et al. 2008) Diagnosis. Male tergite VIII truncate; spermatheca C-shaped, capsule and stem short, broad, and tubular or club-shaped; median lobe of aedeagus with bulbus enlarged. 75. Gnypeta crebrepunctata (Casey) (Fig. 12.109a–h) Tachyusa crebrepunctata Casey 1886. Casey 1906, Moore and Legner 1975, Klimaszewski et al. 2008.

12

Tribe Tachyusini C. G. Thomson, 1859

Diagnosis. Body length 2.7–3.0 mm; uniformly dark brown to black, with tarsi rust brown; integument moderately glossy; pubescence yellowish grey and moderately long and dense (Fig. 12.109a); antennal articles IV–VII moderately elongate, VIII–X slightly transverse; head and pronotum of about the same width; elytra and abdomen wider than either head or pronotum; head rounded posteriorly; pronotum broadest in apical third, about two-fourth (0.6) as wide as elytra, pubescence directed anteriad along midline and laterad elsewhere; elytra at suture slightly longer than pronotum, pubescence directed obliquely postero-laterad, in wavy pattern medially on each side of disc; abdomen subparallel, slightly narrower than elytra at base; metatarsus with two basal tarsomere of about the same length and the third one slightly shorter. MALE. Tergite VIII transverse and truncate apically with slight median emargination (Fig. 12.109d); sternite VIII slightly elongate and broadly rounded apically (Fig. 12.109e); median lobe of aedeagus with narrowly produced apical part of tubus in lateral view (Fig. 12.109b), in dorsal view tubus broadly triangular basally with apex narrow (Fig. 12.109c); internal sac with structures as illustrated (Fig. 12.109b, c). FEMALE. Tergite VIII truncate apically (Fig. 12.109f); sternite VIII broadly rounded apically (Fig. 12.109g); spermatheca C-shaped, capsule tubular and short, stem short and broadly tubular (Fig. 12.109h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, OR. COLLECTION AND HABITAT DATA. Habitat: found in gravel along an edge of a creek, and in rotting Boletus mushrooms. Collecting period: VII, VIII. Collecting method: not recorded. Helenae Species Group (Klimaszewski et al. 2008)

12

Tribe Tachyusini C. G. Thomson, 1859

Diagnosis. Male tergite VIII with apical margin truncate, slightly emarginate medially; median lobe of aedeagus in lateral view with tubus narrowly triangular apically. Female sternite VIII with apical margin moderately deeply emarginate medially; spermatheca S-shaped, capsule pipeshaped, with apical part small, cylindrical, connected at an angle with stem, which is narrow, tubular and sinuate posteriad. 76. Gnypeta helenae Casey (Fig. 12.110a–h) Gnypeta helenae Casey, 1906. Klimaszewski et al. 2008, Brunke et al. 2012. Diagnosis. Body length 2.7–3.0 mm; colour dark brown to black, elytra and legs and/or tarsi often rust brown; integument highly glossy (Fig. 12.110a); pubescence yellowish grey, moderately long and dense; antennomere IV distinctly elongate but distinctly less than half length of antennomere III, V–X subquadrate to slightly transverse; head and pronotum of about same width; elytra markedly wider than either head or pronotum; head rounded posteriorly; pronotum broadest in apical third, pubescence on disc directed anteriad along midline and laterad elsewhere; elytra at suture as long as pronotum, about 1/6 wider than maximum width of pronotum, pubescence on disc directed obliquely posterolaterad, forming wavy pattern medially on each side; abdomen subparallel, distinctly narrower than elytra at base; hind tarsus with basal tarsomere slightly longer than following one. MALE. Tergite VIII transverse, broadly arcuately truncate apically with very shallow emargination medially (Fig. 12.110d); sternite VIII slightly transverse, broadly rounded apically (Fig. 12.110e); median lobe of aedeagus in lateral view with tubus narrowly triangular apically, with ventral margin straight and curved obliquely ventrad apically (Fig. 12.110b); in dorsal view bulbus moderately enlarged and tubus basally as broad as bulbus, apex narrow (Fig. 12.110c); internal sac with structures as illustrated (Fig. 12.110b, c). FEMALE. Tergite VIII broadly arcuately truncate apically (Fig. 12.110f); sternite

241

VIII with apical margin broadly rounded on each side of broad, shallow median emargination (Fig. 12.110g); spermatheca S-shaped, capsule pipe-shaped, with apical part small, stem narrow, tubular, with single narrow curl posteriad (Fig. 12.110h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, ON. USA: AZ, CA, CO, MT, NV, NM, ND, OR. New locality data: British Columbia, Wynndel, Kootenay Lake, 14.VIII.1982, leg. R. Baranowski (ZMLU) 4 male, 3 females. COLLECTION AND HABITAT DATA. Habitat: banks of rivers and lakes and from a eutrophic pond edge; some from silt around Typha (cattails) on a river bank. Collecting period: VI–VIII. Collecting method: sifting litter. Nigrella Species Group (Klimaszewski et al. 2008) Diagnosis. Median lobe of aedeagus with moderately large bulbus which is streamlined with tubus in dorsal view, the internal sac with two pronounced subapical sclerites; female sternite VIII emarginated medially; spermatheca clubshaped, capsule approximately spherical and constricted to a narrow duct connected to a narrow, tubular and sinuate stem. 77. Gnypeta baranowski Klimaszewski, sp.n. (Fig. 12.111a–h) Holotype (male): Canada, British Columbia, 15 km W of Creston, 18.VIII.1982, leg. R. Baranowski (ZMLU). Paratypes: Canada, British Columbia, 12 km W of Cherryville, 20.VIII.1988, leg. R. Danielsson (ZMLU) 1 male; between Lumby and Mabel Lake, 4.VIII.1982, leg. R. Baranowski (ZMLU) 1 female.

242

Etymology. Named for Swedish explorer and insect collector, Richard Baranowski, who provided the type series. Diagnosis. Body length 2.8–3.2 mm; colour dark brown to black, tarsi rust brown; integument highly glossy (Fig. 12.111a); pubescence yellowish grey, moderately long and moderately dense; antennomere IV strongly elongate and as long as half length of antennomere III, V–VI moderately elongate, VII–X subquadrate to slightly transverse; head and pronotum of about same width; head rounded posteriorly; pronotum broadest in apical third, pubescence on disc directed anteriad along midline and laterad elsewhere; elytra at suture as long as pronotum, about 1/3 wider than maximum width of pronotum, pubescence on disc directed obliquely posterolaterad; abdomen subparallel, distinctly narrower than elytra at base; hind tarsus with basal tarsomere as long as following one. MALE. Tergite VIII transverse, broadly arcuately truncate apically with very shallow emargination medially (Fig. 12.111d); sternite VIII slightly transverse, broadly rounded apically (Fig. 12.111e); median lobe of aedeagus in lateral view with tubus narrowly triangular apically and sinuate, with ventral margin arcuately curved obliquely ventrad (Fig. 12.111b), in dorsal view bulbus broadly subparallel, tubus short, broad at base and triangular apically (Fig. 12.111c); internal sac with structures as illustrated (Fig. 12.111b, c). FEMALE. Tergite VIII broadly arcuately truncate apically with shallow median emargination (Fig. 12.111f); sternite VIII with apical margin broadly rounded on each side of broad, deep V-shaped median emargination (Fig. 12.111g); spermatheca club-shaped, capsule rounded, stem slightly sinuate, swelled posteriad (Fig. 12.111h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: VIII. Collecting method: sifting litter.

12

Tribe Tachyusini C. G. Thomson, 1859

Comments. This species is similar to G. saccharina Klimaszewski and Webster, known from New Brunswick. It may be readily distinguished from that species by narrower pronotum, different structures of internal sac of aedeagus, and by deep emargination on female sternite VIII. The two species are apparently closely related. Sellmani Species Group (Klimaszewski et al. 2008) Diagnosis. Median lobe of aedeagus in lateral view with tubus short, narrowly or broadly triangular apically, crista apicalis prominent, angular; spermatheca with capsule widely funnel-shaped to elongate bell-shaped, with broad invagination, stem club-shaped. 78. Gnypeta brevicornis Casey (Fig. 12.112a–h) Gnypeta brevicornis Casey, 1906. Moore and Legner 1975, Klimaszewski et al. 2008. Lectotype (male): Br. C. [British Columbia]; brevicornis Casey; Type USNM 38865; Casey bequest 1925; Lectotype des. Klimaszewski 2007 (USNM). Diagnosis. Body length 2.7–2.9 mm; rust brown with darker head and tip of abdomen (Fig. 12.112a); integument moderately glossy; pubescence yellowish grey and sparse; antennal article IV elongate but distinctly shorter than antennomere III, antennomeres V–VI slightly and VII–IX strongly transverse; head and pronotum of about the same width; elytra and abdomen slightly wider than either head or pronotum; elytra small and moderately elongate; head rounded posteriorly; pronotum broadest in apical third, pubescence directed postero-laterad from midline of disc; elytra at suture approximately as long as pronotum, 1/5 broader than maximum width of pronotum, pubescence directed obliquely posterolaterad from midline of disc; abdomen swollen posteriorly, as broad as elytra at base; metatarsus with basal article of about the same length as the two following

12

Tribe Tachyusini C. G. Thomson, 1859

243

articles combined. MALE. Tergite VIII transverse and truncate apically, slightly concave medially (Fig. 12.112d); sternite VIII elongate and parabolic posteriorly (Fig. 12.112e); median lobe of aedeagus with narrowly triangular apical part of tubus in lateral view (Fig. 12.112b), in dorsal view base of tubus with two lateral projections (Fig. 12.112c); internal sac with structures as illustrated (Fig. 12.112b, c). FEMALE. Tergite VIII truncate apically (Fig. 12.112f); sternite VIII broadly rounded posteriorly (Fig. 12.112g); spermatheca with capsule transverse, funnelshaped with sides arcuate, stem slightly sinuate and swollen posteriorly (Fig. 12.112h). DISTRIBUTION. Origin: CANADA: BC. USA: not recorded.

Nearctic.

COLLECTION AND HABITAT DATA. Habitat: banks of rivers and lakes and from a eutrophic pond edge; some from silt around Typha (cattails) on a river bank. Collecting period: VI–VIII. Collecting method: sifting litter.

Genus Paradilacra Bernhauer, 1909 (Fig. 12.113a–i) (Bernhauer 1909, Casey 1910, 1911, Fenyes 1918, Seevers 1978, Ashe 2001, Gusarov 2003) Diagnosis. Body subparallel, without macrosetae or bristles, integument with dense, minute asperulate punctules and fine microreticulation; pubescence pale, dense and short giving body a dull appearance; midline pubescence of pronotum directed posteriad in apical ½ to 1/3 and anteriorly in posterior ½ to 1/3, on sides of disc pubescence directed lateroposteriad and laterad at base of disc; elytral pubescence longitudinally subparallel except directed laterad basally and apically; head about as wide as pronotum; antennomeres I–II strongly elongate, IV–VI slightly elongate, and VII–X subquadrate; pronotum is two-fifths broader than long; elytra slightly wider than pronotum, at suture about as long as pronotum along midline. MALE. Tergite and sternite VIII

densely pubescent; median lobe of aedeagus of Gnypeta type, with triangular crista apicalis, and large bulbus and narrow triangular tubus in lateral view. FEMALE. Tergite and sternite VIII densely pubescent; spermatheca with globular capsule bearing wide but shallow apical invagination; stem narrowly elongate, slightly sinuate. 79. Paradilacra (Fig. 12.113a–i)

densissima

(Bernhauer)

Atheta densissima Bernhauer, 1909. Casey 1910, Seevers 1978, Ashe 2001, Gusarov 2003 [synonymies]. Lectotype (male): Cal. [ifornia], S. Anselmo; densissima Bernh. Typus Fenyes; ambigua Er.? Fenyes, det Bernh.; not ambigua Erichson det. Lohse; lectotypus Atheta densissima Bernhauer, V.I. Gusarov des. 1999; Paradilacra densissima (Bernh). det. V.I.Gusarov (FMNH). Examined. Paralectotype (male): Willist’n, ND, Jun 8–9, Wickham; 7614; densissima Bernh. Fenyes; paralectotypus Atheta densissima Bernhauer, V.I. Gusarov des. 1999; Paradilacra densissima (Bernh). det. V.I.Gusarov (FMNH). Examined. Diagnosis. Body subparallel, length 2.6–3.0 mm, dark brown with slightly reddish central part of elytra, tarsi and antennae slightly paler; integument with dense, minute asperulate punctules and fine microreticulation; pubescence pale, dense and short (Fig. 12.113a); head slightly narrower than elytra, eyes moderately large; antennomeres VII–X subquadrate; pronotum rounded laterally, pubescence pattern described under genus diagnosis; elytra distinctly wider than pronotum, at suture about as long as pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically (Fig. 12.113d); sternite VIII parabolic apically (Fig. 12.113e); bulbus of median lobe of aedeagus large and swollen, tubus short, about arcuately curved ventrad, narrowly triangular at apex in lateral view (Fig. 12.113b), in dorsal view bulbus broad, and tubus short, narrowly triangular apically (Fig. 12.113c). FEMALE. Tergite VIII arcuate apically and truncate medially (Fig. 12.113f); sternite VIII with small median apical emargination

244

(Fig. 12.113g); spermatheca with globular capsule bearing wide but shallow apical invagination, stem narrowly elongate, slightly sinuate (Fig. 12.113h, i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: CA, MT, NV, ND, OR, UT. COLLECTION AND HABITAT DATA. Habitat: banks lakes, rivers, and reservoir, banks of beaver pond. Collecting period: VI, VIII. Collecting method: treading vegetation in wet habitats.

Genus Tachyusa Erichson, 1837 (Figs. 12.114–12.116) (Paśnik 2006, Klimaszewski et al. 2008) Diagnosis. Body elongate with slender and clavate abdomen, convex and glossy, length 2.3–3.9 mm; head without infraorbital carina; pronotal hypomera fully visible in lateral view; elytra laterally distinctly arcuate, widest at middle, posterolateral angles distinctly emarginate or broadly rounded; abdomen clavate, distinctly constricted at base, base narrower than width of elytra, strongly swollen posteriorly, tergite V (third visible) moderately transverse, 1.2–1.8 times as wide as long; tergites III–V (first to third visible) or III–VI (first to fourth visible) distinctly transversely impressed at base, impressions coarsely punctate, interstices between punctures forming longitudinal ridges, some species with distinct median carina; pubescence on tergites III–V (first to third visible) directed inward. Comments. Species affiliated with wetland and riparian habitats along margins of ponds, lakes, streams, and marshes. Key to Species of Tachyusa 1. Pronotum subquadrate (Fig. 12.114a); head, pronotum and elytra uniformly dark brown, two

12

Tribe Tachyusini C. G. Thomson, 1859

basal abdominal tergites dark red brown (Fig. 12.114a); male sternite VIII rounded apically (Fig. 12.114g); female sternite VIII with shallow, scarcely visible apical emargination (Fig. 12.114i); tubus of median lobe of aedeagus as illustrated (Fig. 12.114b, c) ............. Tachyusa americanoides Paśnik – Pronotum distinctly transverse (Figs. 12.114a and 12.115a); forebody bicoloured, two basal abdominal tergites yellow or pale red brown (Figs. 12.114a and 12.115a); male sternite VIII rounded apically or produced medially and sharply pointed (Fig. 12.115f); female sternite VIII with deep, V-shaped apical emargination (Figs. 12.115i and 12.116h); tubus of median lobe of aedeagus as illustrated (Figs. 12.115b, c and 12.116b) ......................................................2 2. Pronotum evenly yellow light brown (Fig. 12.115a); male sternite VIII produced apically and pointed medially (Fig. 12.115f); tubus of median lobe of aedeagus sinuate medially (Fig. 12.115b)....................................................... Tachyusa cavicollis LeConte – Pronotum evenly dark brown or blackish with paler base of disc (Fig. 12.116a); male sternite VIII rounded apically (Fig. 12.116e); tubus of median lobe of aedeagus arcuate medially (Fig. 12.116b)............ Tachyusa obsoleta Casey

Coarctata Species Group (Paśnik 2006) Diagnosis. Elytral punctation dense, interstices between punctures equal to puncture diameter, punctures distinctly asperate; first three visible abdominal tergites with moderately dense punctation, interstices between punctures 1.5–2.0 times their diameter, punctures small but clearly visible, punctation on next two tergites VI–VII slightly denser than that on first three; abdomen moderately glossy, with pubescence short, dense and subrecumbent.

12

Tribe Tachyusini C. G. Thomson, 1859

80. Tachyusa 12.114a–k)

americanoides

Paśnik (Fig.

Tachyusa americanoides Paśnik, 2006. Klimaszewski et al. 2011, 2018, Webster et al. 2012. Diagnosis. Body convex, parallel-sided, length 2.5–3.2 mm, integument glossy, colour dark brown to black, abdomen black with first three visible tergites reddish brown, legs reddish brown, antennae brown with antennomeres I–III red (Fig. 12.114a); head circular in outline, convex, glossy, narrower than pronotum; eyes relatively small, moderately protruding from lateral contours of head, slightly shorter than genae in dorsal view; surface of head without microsculpture, punctation fine and moderately dense, pubescence on disc short and moderately dense, directed inward, antennae short, slightly incrassate apically, antennomeres II and III subequal in length, antennomeres IV–VII subquadrate to slightly elongate, antennomeres VIII–X sexually dimorphic, quadrate in male and slightly transverse in female, antennomere XI nearly conical; pronotum quadrate, convex, glossy, widest in apical third, lateral sides rounded in front, gradually narrowed in straight line to obtuse hind angles, with small and shallow transverse basal impression, surface lacking microsculpture, punctation fine, dense and asperate, pubescence at midline directed posteriad; elytra longer than wide, slightly wider than pronotum, lateral sides arcuate, at suture as long as pronotal length at midline; surface lacking microsculpture; punctation fine, dense and asperate; abdomen parallel-sided, first three visible tergites with deep transverse basal impression, impressions with 7–8 longitudinal ridges and distinct median carina, tergal punctation fine and moderately dense, punctures moderately small and well visible, surface without microsculpture. MALE. Tergite broadly arcuate apically (Fig. 12.114f); sternite VIII rounded apically

245

(Fig. 12.114g); tubus of median lobe of aedeagus arcuately curved slightly ventrad in basal half, then straight to narrowly triangular and narrowly rounded apex in lateral view (Fig. 12.114b, c), in dorsal view bulbus ovoid in shale, tubus short, triangular (Figs. 12.114d, e). FEMALE. Tergite arcuate apically (Fig. 12.114h); sternite VIII truncate apically with minute median emargination (Fig. 12.114i); spermatheca with stem thick, slightly sinuate (Fig. 12.114j, k). Externally, this species is extremely similar to T. americana Casey, but in T. americanoides antennomeres II and III are subequal in length, the elytra are longer than wide, the elytral length at the suture is as long as the pronotal length at the midline, and the shape of the tubus of the median lobe of the aedeagus is distinctive. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NF, NS, NT, ON, QC. USA: IL, MA, MT, NH, NY, PA, VT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: under/in debris in riparian habitats and along pond margins; under debris on muddy soil near a small pool, under debris on clay and sand at a river margin, in grassy debris near the overflow area of a beaver dam, among sticks, debris, and clay on a beaver dam, under debris on muddy soil along a slow-flowing river in a silver maple swamp, in debris along the margin of a rocky river, under debris on muddy soil near a small pool in a silver maple forest, and among gravel on gravel bar along a small shaded brook in a northern coniferous forest. Collecting period: V–VII. Collecting method: sifting debris and, aspirating adults from debris on muddy soils or from soil surface. 81. Tachyusa cavicollis LeConte (Fig. 12.115a–i) Tachyusa cavicollis LeConte, 1863. Casey, 1906 [as T. carolinae]; Paśnik 2006.

246

Diagnosis. Body length 2.7–3.0 mm, convex, parallel-sided, glossy; head pitchy brown to black, pronotum brownish red, elytra brown with posterior margin yellow, abdomen black with three basal tergites red, legs and antennae yellow to red (Fig. 12.115a); head quadrate, flattened dorsally, broadly and moderately deeply impressed medially in male and without impression in female; eyes large, moderately protruding from lateral contours of head, length of each seen from above subequal to that of postocular region; surface of head with distinct isodiametric mesh microsculpture in male and without microsculpture in female; punctation fine and moderately dense; antennae long, incrassate, antennomere III shorter than II, antennomeres IV–IX longer than wide, antennomere X quadrate, antennomere XI nearly conical; pronotum transverse, lateral sides gradually narrowed in straight line to obtuse hind angles; before base with small and shallow transverse impression; broadly and deeply impressed medially in male and without impression in female; surface with strong isodiametric mesh microsculpture in male and without microsculpture in female; punctation fine, moderately dense and asperate; pubescence at midline directed posteriorly; elytra quadrate, at suture as long as pronotal length at midline; surface lacking microsculpture; punctation fine, dense and aspirate; abdomen parallel-sided, bases of tergites III–V each with deep transverse impression, impressions with 7–8 longitudinal ridges and distinct median carina, tergal punctation fine and dense, punctures small and well visible, surface without microsculpture. MALE. Tergite VIII truncate apically (Fig. 12.115e); sternite VIII produced apically and pointed medially (Fig. 12.115f); tubus of median lobe of aedeagus slightly sinuate ventrally and narrowly subparallel at apex, apex rounded in lateral view (Fig. 12.115b, c), median lobe in dorsal view with ovoid bulbus and triangular tubus (Fig. 12.115d). FEMALE. Tergite truncate apically (Fig. 12.115g); sternite VIII with narrow V-shaped apical emargination (Fig. 12.115h); spermatheca with stem sinuate, slightly swollen posteriorly (Fig. 12.115i).

12

Tribe Tachyusini C. G. Thomson, 1859

Tachyusa cavicollis is similar to T. obsoleta from which it can be distinguished by the stouter body, the paler pronotum, the male pronotum more strongly impressed medially, and particularly by the different shape of genitalia. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AR, FL, IL, IN, KS, LA, MA, ME, MD, NE, NC, NH, NJ, NM, OH, PA, SD, TX, VA, VT, WY. COLLECTION AND HABITAT DATA. Habitat: in unspecified forests, along edge of drying pools, along mud shore (Paśnik 2006). Collecting period: IV–VIII. Collecting method: not recorded. Comments. We have examined additional specimens from IL and OH and confirm their identity (FMNH). 82. Tachyusa obsoleta Casey (Fig. 12.116a–h) Tachyusa obsoleta Casey, 1906. Paśnik 2006, Webster et al. 2012, Klimaszewski et al. 2018. Diagnosis. Body convex, parallel-sided, length 2.5–2.8 mm, integument glossy, head pitchy brown to black, pronotum brown, elytra brownish red to light brown, abdomen black with three visible tergites red to reddish brown, legs yellow, antennae brown with antennomeres I–III yellowish red (Fig. 12.116a); head quadrate, broadly and shallowly impressed medially in male and without impression in female; eyes large, subequal in length to postocular region; temples parallel near eyes; surface of head with isodiametric mesh microsculpture in male and without microsculpture in female; punctation fine and moderately dense; antennae long, slightly incrassate, antennomere III shorter than II, antennomeres IV–VII elongate, VIII–X subquadrate, and XI nearly conical; pronotum transverse, lateral sides gradually narrowed in about straight line to obtuse hind angles, with small and shallow basal transverse impression broadly and shallowly impressed medially in male and without medial impression in female,

12

Tribe Tachyusini C. G. Thomson, 1859

247

surface with isodiametric mesh microsculpture in male and without microsculpture in female, punctation fine, dense and asperate, pubescence at midline directed posteriad; elytra slightly transverse, at suture as long as pronotal length at midline, surface lacking microsculpture, punctation fine, dense and asperate; abdomen parallelsided, bases of first three visible tergites each with deep transverse impression, impressions with 7–8 longitudinal ridges and distinct median carina, tergal punctation fine and dense, punctures small and well visible, surface without microsculpture. MALE. Tergite truncate apically (Fig. 12.116d); sternite VIII rounded apically (Fig. 12.116e); tubus of median lobe of aedeagus in lateral view arcuate from tip of large fin-shaped crista apicalis in basal half, then almost straight, subparallel in apical half to narrowly rounded apex (Fig. 12.116b), median lobe in dorsal view as illustrated (Fig. 12.116c). FEMALE. Tergite truncate apically (Fig. 12.116f); sternite VIII V-shaped emarginated medially at apex (Fig. 12.116g); spermatheca with stem thin, S-shaped, swollen posteriorly (Fig. 12.116h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, SK. USA: CA, CO, DC, IL, IN, KS, MT, NE, NH, NV, OK, SD, UT, WI, WY. COLLECTION AND HABITAT DATA. Habitat: riparian habitats usually on moist sand or clay and in debris. Most specimens from NB, were collected from sand near a river margin. Others from moist clay and from grass and debris along river margins and creeks. Collecting period: V–VIII. Collecting method: lightly splashing water on fine sand near a river margin and aspirating specimens, sifting grass and debris.

Fig. 12.105 (a–b) Brachyusa americana (Fenyes) [after Seevers (1978)]: (a) median lobe of aedeagus of in lateral view; (b) paramere. Scale bar for habitus ¼ 0.2 mm

248

Fig. 12.106 (a–g) Images of Brachyusa helenae (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

12

Tribe Tachyusini C. G. Thomson, 1859

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Tachyusini C. G. Thomson, 1859

Fig. 12.107 (a–h) Images of Gnypeta caerulea (C.R. Sahlberg): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

249

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

250

Fig. 12.108 (a–h) Images of Gnypeta lohsei Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; d, male tergite VIII; (e) male

12

Tribe Tachyusini C. G. Thomson, 1859

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

12

Tribe Tachyusini C. G. Thomson, 1859

Fig. 12.109 (a–h) Images of Gnypeta crebrepunctata Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

251

female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

252

Fig. 12.110 (a–h) Images of Gnypeta helenae Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

12

Tribe Tachyusini C. G. Thomson, 1859

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

12

Tribe Tachyusini C. G. Thomson, 1859

Fig. 12.111 (a–h) Images of Gnypeta baranowskii Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

253

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–d), holotype; (e–g), paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

254

Fig. 12.112 (a–h) Images of Gnypeta brevicornis Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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Tribe Tachyusini C. G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a, b, d, e) lectotype, front legs reconstructed. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Tachyusini C. G. Thomson, 1859

Fig. 12.113 (a–i) Images of Paradilacra densissima (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

255

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (b) holotype; (a, c–d) paralectotype; (f–i) after Gusarov (2003). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

256

Fig. 12.114 (a–k) Images of Tachyusa americanoides Paśnik: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d, e) median lobe of aedeagus in dorsal view; (f) male tergite VIII; (g) male sternite VIII;

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Tribe Tachyusini C. G. Thomson, 1859

(h) female tergite VIII; (i) female sternite VIII; (j, k) spermatheca. (b, d, k) after Paśnik (2006). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Tachyusini C. G. Thomson, 1859

Fig. 12.115 (a–i) Images of Tachyusa cavicollis LeConte: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male

257

tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. Figure (c, d) after Paśnik (2006). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

258

Fig. 12.116 (a–h) Images of Tachyusa obsoleta Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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Tribe Tachyusini C. G. Thomson, 1859

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Figure (c) after Paśnik (2006). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Tachyusini C. G. Thomson, 1859

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Bernhauer M (1909) Neue Aleocharini aus Nordamerika. (Col.) (4. Stück.). Deutsch Entomol Z 1909 (4):515–528 Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013) Checklist of beetles (Coleoptera) of Canada and Alaska, 2nd edn. Pensoft, Series Faunistica No. 109. Sofia-Moscow, 402 pp Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II Zookeys [Special Issue] 186:119–206 Casey TL (1886) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1:285–336 Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St. Louis 16(6):125–434 Casey TL (1910) New species of the staphylinid tribe Myrmedoniini [pp 1–183]. In: Memoirs on the Coleoptera. I. New Era Printing Co., Lancaster, PA, 205 pp Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, PA, 259 pp Erichson WF (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera Insectorum Vol. XXVII, Fascicule 173A. M. Nijhoff, Den Haag. 453 pp + 7 pls Fenyes A (1921) New genera and species of Aleocharinae with a polytomic synopsis of the tribes. Bull Mus Comp Zool 65(2):17–36 Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353:1–134 Klimaszewski J, Savard K, Pelletier G, Webster R (2008) Species review of the genus Gnypeta Thomson from Canada, Alaska and Greenland (Coleoptera, Staphylinidae, Aleocharinae): systematics, bionomics and distribution. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys 2 [Special Issue]:11–84 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera,

259 Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Bourdon C (2012) Further contributions to the aleocharine fauna of the Yukon Territory, Canada (Coleoptera, Staphylinidae). In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. Zookeys [Special Issue] 186:207–237 Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp LeConte JL (1863) New species of North American Coleoptera. Prepared for the Smithsonian Institution. Part I. Smithson Miscell Collect 6(167):1–86 Lohse GA (1989) Familie: Staphylinidae. In: Lohse GA, Lucht WH (eds) Die Käfer Mitteleuropas, Band 12, 1 Suplementband mit Katalogteil. Goecke and Evers, Krefeld, pp 185–240 Majka CG, Klimaszewski J (2008) Introduced Staphylinidae (Coleoptera) in the Maritime Provinces of Canada. Can Entomol 140(1):48–72 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Mulsant E, Rey C (1873) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens (Suite) [Tableau... des Aléochariens + Septième Branche, Myrmédoniaires, part 1]. Deyrolle, Paris, [2] + 695 pp, pls 1–5 Osswald J, Bachmann L, Gusarov VI (2013) Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae). Syst Entomol 38 (3):507–522 Paśnik G (2006) A revision of the World species of the genus Tachyusa Erichson, 1837 (Coleoptera, Staphylinidae: Aleocharinae). Zootaxa 1146(1):1–152 Sahlberg CR (1831) [pp 361–408]. In: Insecta Fennica, dissertationibus academicis, A. 1817–1834 editis. Pars I: A. J.C. Frenckel, Helsingfors, VIII + 519 pp Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana Zool 71:vi + 289 pp

260 Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom. Berlingska Boktryckeriet, Lund [3] + 290 pp Webster RP, Klimaszewski J, Sweeney J D, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with

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new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:83–118

Tribe Hypocyphtini Laporte, 1835

Tribe Hypocyphtini Laporte, 1835 (Figs. 13.117–13.119) (Ashe 2001) Diagnosis. Body minute and compact, length 0.8–1.6 mm; antennae with ten antennomeres, antennomeres VIII–X enlarged and forming loose club; hind coxae with large ventral lamella covering base of femur; tarsal formula 4-4-4. Key to Genera of Hypocyphtini 1. Pronotum strongly transverse, maximum width of pronotum at least as maximum width of elytra, elytra at suture shorter than pronotum along midline, habitus as in Fig. 13.117a ...................................Cypha Leach – Pronotum moderately transverse, maximum width of pronotum distinctly less than maximum width of elytra, elytra at suture as long as pronotum along midline, habitus as in Figs. 13.118a and 13.119a .................................2 2. Body broadly ovoid, robust (Fig. 13.118a); pronotal hypomeron not visible in lateral view; median lobe of aedeagus with very long flagellum, several times coiled in bulbus (Fig. 13.118b, c) ....................................................... Holobus Solier – Body narrowly ovoid or parallel-sided (Fig. 13.119a); pronotal hypomeron at least narrowly visible in lateral view; median lobe of aedeagus with flagellum shorter than length of

13

median lobe of aedeagus (Fig. 13.119b).............. ...........................................Oligota Mannerheim

Genus Cypha Leach, 1819 (Fig. 13.117a–d) (Ashe 2001; Klimaszewski et al. 2008, 2018) Diagnosis. Body subovate, length 0.8–1.25 mm, colour brownish to black, integument moderately glossy, with coarse sculpture and inconspicuous pubescence; head hypognathous (strongly deflexed), narrower than elytra; eyes large; antennae slender, antennomeres VIII–X forming club, terminal antennomere as long as two preceding ones combined; pronotum narrow and distinctly transverse, twice as wide as long, apex narrower than base and broadly emarginate adjacent to head, sides rounded, hind angles obtusely rounded, posterior margin slightly bisinuate, maximum width of pronotum at least as maximum width of elytra; pronotal hypomera not visible in lateral view; elytra at suture distinctly shorter then pronotum along midline; mesocoxae narrowly separated; process of mesoventrite long and obtusely pointed, extending to three-quarters length of mesocoxae and overlapping apex of short and broader process of metaventrite; abdomen acuminate, tergites not impressed; hind tarsi with tarsomere I subequal in length to II and III

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_13

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combined; median lobe of aedeagus very long, narrow, gradually tapering apicad, paramere evenly narrowly elongate with small apical lobe bearing short apical setae; spermatheca not found. 83. Cypha crotchii (Horn) (Fig. 13.117a–d) Hypocyphtus crotchii Horn, Klimaszewski et al. 2008, 2016.

Tribe Hypocyphtini Laporte, 1835

DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, SK. USA: ID, OR, WA. COLLECTION AND HABITAT DATA. Habitat: decaying mushrooms. Collecting period: IX. Collecting method: sifting decaying mushrooms.

1877.

Genus Holobus Solier, 1849 Diagnosis. Body length 0.8–1.0 mm, small, subovate, colour light brown to dark brown (Fig. 13.117a); pubescence long, sparse; head broad, retracted under pronotum; pronotum distinctly transverse, markedly narrowed apically, wider than elytra; elytra distinctly transverse, shorter at suture than pronotum; abdomen shortly triangular, usually about as long as pronotum (individual specimens may exhibit different degree of abdominal contraction). MALE. Tergite VIII distinctly transverse, with apical margin broadly truncate, arcuate laterally, microsetae long and elevated on microtubercules (Fig. 13.117c); sternite VIII transverse with lateral margins obliquely converging posteriad to narrowly truncate apex, antecostal suture broadly separated from basal margin, pubescence as on tergite VIII (Fig. 13.117d); median lobe of aedeagus in lateral view with bulbus arched abruptly dorsad apically so that base of tubus situated on dorsoapical margin of bulbus, tubus bent obliquely ventrad at base, straight medially and projecting ventrad at apex (Fig. 13.117b), in dorsal view tubus narrow basally, very long, evenly tapering to narrowly rounded apex, internal sac with two small lunular-shaped sclerites, visible as one in lateral view (Fig. 13.117b). FEMALE. Undescribed.

(Fig. 13.118a–f) (Ashe 2001; Klimaszewski and Winchester 2002)

Cypha inexpectata Klimaszewski and Godin is extremely similar externally to the western C. crotchii (Horn), the only other recorded species of this genus from Canada. It differs from C. crotchii in having the median lobe of the aedeagus with the tubus broadly arcuate, with the apex very fine in lateral view and the internal sac lacking sclerites.

Diagnosis. Body minute, length 1.1 mm, broadly ovoid, compact and glossy, sparsely pubescent, reddish brown with darker some abdominal tergites and paratergites, basal antennomeres and legs yellowish (Fig. 13.118a); head transverse, narrowed behind strongly deflexed, without distinct neck; eyes large and slightly protruding; basal antennomere thick,

Diagnosis. Body broadly ovoid, robust, and glossy, length 1.0–1.4 mm; head, large, inflexed under pronotum, eyes large and slightly protruding; pronotum narrower than maximum width of elytra, pronotal hypomera not visible in lateral view; elytra at suture about as long as pronotum; abdomen conical, strongly tapering apicad. Male median lobe of aedeagus with very long flagellum, several times longer than median lobe, in repose position coiled in the bulbus. Distinguished from Oligota by pronotal hypomera not visible in lateral view and morphology of median lobe of aedeagus. Some species are found on vegetation and are predators of phytophagous mites (Ashe 2001). One species is recorded from British Columbia (Klimaszewski and Winchester 2002), and six from eastern North America and California (Ashe 2001). 84. Holobus vancouveri (Fig. 13.118a–f)

Klimaszewski

Holobus vancouveri Klimaszewski, Klimaszewski and Winchester 2002.

in

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Tribe Hypocyphtini Laporte, 1835

263

next five slender and last three enlarged and forming loose club; pronotum about twice as wide as long; elytra at suture about as long as pronotum; abdomen conical strongly tapering apically. MALE. Tergite VIII broadly arcuate apically (Fig. 13.118e); sternite VIII truncate apically (Fig. 13.118f); tubus of median lobe of aedeagus straight ventrally and narrowly produced ventrad in subapical part, apex pointed in lateral view (Fig. 13.118b, c); flagellum longer than median lobe and several times coiled in bulbus (Fig. 13.118b, c); in dorsal view tubus narrowly elongate and asymmetrical apically pointing sideways (Fig. 13.118d). FEMALE. Undescribed.

tergites impressed basally; median lobe of aedeagus with large bulbus and moderately long tubus, internal sac structures complex and distinct; spermatheca simple, capsule and stem sac-shaped.

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

85. Oligota parva Kraatz { (Fig. 13.119a–f)

COLLECTION AND HABITAT DATA. Habitat: clear-cut site of Sitka spruce forest in Carmanah Valley, Vancouver Island. Collecting period: VII–VIII. Collecting method: Malaise trap.

Genus Oligota Mannerheim, 1830 (Fig. 13.119a–f) (Frank 1972; Frank et al. 1992; Ashe 2001; Klimaszewski et al. 2016, 2018; Webster et al. 2016) Diagnosis. Body narrowly oval to subparallel, short, compact and robust, length 0.9–1.6 mm, integument with coarse microsculpture; colour brown to black, often with appendages lighter; head deflexed, about one-fourth wider than long; eyes large; antennomeres I–II distinctly elongate, III slightly elongate or subquadrate, IV–VI small and slender, VII–X enlarged, distinctly transverse and forming loose club; pronotum up to one-half broader than long, dorsum convex, pronotal hypomera at least narrowly visible in lateral view, pubescence on disc directed obliquely posteriad; elytra as long as pronotum or slightly longer; mesocoxae widely separated, process of mesoventrite broad; process of metaventrite moderately arcuate; first three visible abdominal

Comments. Widespread genus, with many cosmopolitan species. Adults often found on flowers, in vegetation, fungi, and decaying organic materials where they feed on mites. There is an additional adventive species, Oligota inflata (Mannerheim), recently recorded from Saskatchewan but so far undetected in BC (Klimaszewski et al. 2016). Several species are recorded in eastern Canada (Klimaszewski et al. 2018).

Oligota parva Kraatz, 1862. Horion 1967, Moore and Legner 1975, Williams 1979, Smetana 2004, Majka et al. 2008, Klimaszewski et al. 2013, 2018, Webster et al. 2016. Diagnosis. Body narrowly elongate, moderately convex, length 0.9–1.2 mm, colour dark brown to almost black, with posterior section of abdomen, legs and bases of antennae light brown, head slightly deflexed (Fig. 13.119a); antennomeres VII–X distinctly transverse, forming a loose club; pronotum convex, transverse, punctate, with sculpture of minute tubercles; elytra transverse, at suture at most as long as pronotum; abdomen with deep basal impressions on first three visible tergites, without carinae. MALE. Tergite VIII truncate apically (Fig. 13.119c); sternite VIII broadly rounded apically (Fig. 13.119d); median lobe of aedeagus moderately broad in dorsal view, in lateral view tubus arcuately curved ventrad in apical 2/3, apex acute, with distinct sclerites as illustrated (Fig. 13.119b). FEMALE. Tergite VIII truncate apically (Fig. 13.119e); sternite VIII broadly rounded apically (Fig. 13.119f); spermatheca not found, possibly not sclerotized or lacking. DISTRIBUTION. Origin: Cosmopolitan, adventive in Canada and USA. CANADA: BC [new

264

generic and specific provincial record], NB, PE. USA: CA, MA, MO, NV, TX. We report this species from BC for the first time here based on the following: British Columbia, Victoria, Oak Bay, 25.V.2015, 48.4207–123.3033, 3 m, compost, soil sifting, coll. B. Godin and D. Horwood (BGC, LFC) 2 males. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in NB, it was common in a pile of decaying and moldy

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Tribe Hypocyphtini Laporte, 1835

corncobs and cornhusks near a composter in a residential area adjacent to a mixed forest. Mites were abundant in the moldy corncobs and cornhusks where the specimens were collected. In PE, adults were found under coastline drift at the top of the littoral zone. In BC, two males were captured in compost. Elsewhere, associated with anthropogenic habitats and found in compost, on dung, in fermenting materials, in old hay and grass, and in other decomposing materials. Collecting period: VIII–X. Collecting method: sifting organic litter.

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Tribe Hypocyphtini Laporte, 1835

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Fig. 13.117 (a–d) Images of Cypha crotchii (Horn): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 13.118 (a–f) Images of Holobus vancouveri Klimaszewski (holotype): (a) habitus in dorsal view; (b, c), median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f)

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Tribe Hypocyphtini Laporte, 1835

male sternite VIII. (b, d) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Hypocyphtini Laporte, 1835

Fig. 13.119 (a–f) Images of Oligota parva Kraatz: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

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(e) female tergite VIII; (f) female sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

268

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Frank JH (1972) The genus Oligota Mannerheim in the Caribbean region (Coleoptera: Staphylinidae). Coleopt Bull 26(4):125–146 Frank JH, Bennett FD, Cromroy HL (1992) Distribution and prey records for Oligota minuta (Coleoptera: Staphylinidae), a predator of mites. Florida Entomol 75(3):376–380 Horion A (1967) Faunistik der Mitteleuropȁischen Kȁfer. Bd. XI: Staphylinidae. 3. Teil: Habrocerinae bis Aleocharinae (ohne Subtribus Athetae). P.C.V. Schmidt, Űberlingen – Bodensee, xxiv + 419 pp Horn GH (1877) Synopsis of the genera and species of the staphylinid tribe Tachyporini of the U.S. Trans Am Entomol Soc 6:81–128 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belg Entomol 40:3–126 Klimaszewski J, Godin B, Pelletier G, Savard K (2008) Six new species and records of aleocharine beetles from the Yukon and Alaska (Coleoptera: Staphylinidae: Aleocharinae). Can Entomol 140 (3):265–291 Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013) Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia-Moscow, 360 pp Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp

13

Tribe Hypocyphtini Laporte, 1835

Kraatz G (1862) Synonymische Bemerkungen. Ueber Coleopteren. Berliner Entomol Z 6:298–300 Laporte FL (1835) Études entomologiques, ou description d’insectes nouveaux: et observations sur leur synonymie. Méquignon-Marvis Père et Fils. Paris, 159 pp + 4 pls Leach WE (1819) New genera. In: Samouelle (ed) The Entomologist’s Useful Compendium; or an introduction to the knowledge of British insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; etc. Thomas Boys, London, 496 pp + 12 pls Majka CG, Klimaszewski J, Lauff RF (2008) The coastal rove beetles (Coleoptera, Staphylinidae) of Atlantic Canada: a survey and new records. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:115–150 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp [separate, re-issued in Mémoires Présentées à l’Académie Impériale des Sciences de St.-Pétersbourg 1:415–501, in 1831] Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Smetana A (2004) Family Staphylinidae Latreille, 1802 New nomenclatorial and taxonomic acts, and comments [pp 29–35], subfamilies OmaliinaeDasycerinae [pp 237–272], subfamilies Phloeocharinae-Apateticinae [pp 329–495], subfamilies Piestinae-Staphylininae [pp 504–698]. In: Löbl I, Smetana A (eds) Catalogue of Palearctic Coleoptera, vol 2. Hydrophiloidea – Histeroidea – Staphylinoidea. Apollo Books, Stenstrup, 942 pp Solier AJ (1849) Coleoptera. In: Historia fisica y politica de Chile. Insecta 4(2):145–312, 673–824 Webster RP, Klimaszewski J, Bourdon C, Sweeney JD, Hughes CC, Labrecque M (2016) Further contribution to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including description of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216 Williams SA (1979) The genus Oligota Mannerheim (Col., Staphylinidae) in the Ethiopian region. Entomol Mon Mag 114[1978]:177–190

Tribe Myllaenini Ganglbauer, 1895

Tribe Myllaenini Ganglbauer, 1895 (Figs. 14.120–14.123) (Klimaszewski 1982; Ashe 2001) Diagnosis. Body fusiform, pronotal hypomera not visible in lateral view, abdomen tapering apically, with coarse macrosetae; labial and maxillary palpi very long and stylate; galeae very slender with setae at apex; laciniae long and stylate with widely scattered teeth internally; mentum with anterolateral margins produced into spinose processes; tarsal formula 4-4-5, tarsal claws slender. Species affiliated with riparian habitats. Comments. Includes species.

riparian

and

wetland

14

reduced to narrow, transverse plate in front of procoxae; procoxae conical; tarsal formula 4-4-5; elytra transverse, laterally subparallel or rounded, sinuate at base; scutellum small, covered by pronotum; abdomen tapering from base, with coarse macrosetae; median lobe of aedeagus narrow, bulbus moderately large, internal sac with simple or complex sclerites, paramere with apical lobe narrow and very long, with two long basal and two short apical setae; spermatheca variable in shape, capsule spherical or sac-shaped with apical invagination small to large, stem short and not coiled or long and tightly multi-coiled. Comments. Adults are found along margins of fresh water streams, ponds, lakes, and bogs. Key to Species of Myllaena

Genus Myllaena Erichson, 1837 (Figs. 14.120–14.123) (Klimaszewski 1982) Diagnosis. Body narrowly ovoid or fusiform, length 0.6–3.4 mm; integument covered with short and dense pubescence; head hypognathous; maxillary palpi with four palpomeres, palpomeres I and IV vestigial; glossae styliform; labial palpi and paraglossae reduced; mandibles with inner margins smooth; pronotum broadly rounded anteriad, basal margin arcuate or sinuately truncate, posterior lateral angles well defined, hypomera not visible in lateral view; prosternum

1. Antennomeres VII–X about two times longer than wide (Figs. 14.120a and 14.122a); elytra at suture slightly shorter than pronotum along midline (Figs. 14.120a and 14.122a) .......................2 – Antennomeres VII–X about one and a half times longer than wide (Figs. 14.121a and 14.123a); elytra at suture at least as long as pronotum along midline (Figs. 14.121a and 14.123a) .............................................................3 2. Spermathecal capsule tubular, without apical invagination, instead with a small apical projection (Fig. 14.120h, i); median lobe of aedeagus

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_14

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270

with characteristic arrangement and shape of internal sac sclerites, as illustrated (Fig. 14.120b, c) ............................................ Myllaena audax Casey – Spermathecal capsule spherical, with broad apical invagination (Fig. 14.122i); median lobe of aedeagus with different arrangement and shape of internal sac sclerites, as illustrated (Fig. 14.122b–d)........ Myllaena insomnis Casey 3. Pronotum convex, rounded apically, width/ length ratio 1.5 (Fig. 14.121a); median lobe of aedeagus and spermatheca as illustrated (Fig. 14.121b, c, h); male and female tergite VIII strongly produced apically (Fig. 14.121d, f) ...................................... Myllaena decreta Casey – Pronotum flattened, truncate apically, width/ length ratio 1.6 (Fig. 14.123a); median lobe of aedeagus and spermatheca as illustrated (Fig. 14.123b–d, i); male and female tergite VIII rounded apically (Fig. 14.123e, g)....................... ...............................Myllaena fenyesi Bernhauer

Insomnis Species Group (Klimaszewski 1982) Diagnosis. Body length 1.9–2.6 mm; antennomeres VII–X at least twice as long as maximum width; tergite X of both sexes nearly completely divided; sternite VIII of both sexes with 5–7 pairs of setae; median lobe of aedeagus in lateral view with apex narrowly rounded; stem of spermatheca with many intertwined coils.

14

Tribe Myllaenini Ganglbauer, 1895

about equal to that of elytra; elytra at suture slightly shorter than pronotum along midline of disc; abdomen tapering apically. MALE. Tergite VIII slightly obtusely triangularly produced apically (Fig. 14.120d); sternite VIII broadly parabolically rounded apically (Fig. 14.120e); median lobe of aedeagus in dorsal view with tubus elongate, subparallel, truncate apically (Fig. 14.120c), in lateral view tubus with ventral margin arcuate basally, straight to narrow and rounded apex (Fig. 14.120b). FEMALE. Tergite and sternite VIII similar to those of male (Fig. 14.120f, g); spermatheca with capsule pipe-shaped, with small apical projection, stem long, with about 24 tight intertwined coils (Fig. 14.120h, i). DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NF, NT, ON, QC. USA: AK, CA, GA, IL, LA, MA, NH, NJ, NY, OR, RI, UT, WA. COLLECTION AND HABITAT DATA. Habitat: various wetland habitats; in leaf litter and grass on hummocks in a wet alder swamp, sedge marshes in leaf litter and moss at base of trees, tamarack bog in sphagnum hummocks with dense leatherleaf, in debris near creeks, rivers, and lakes, in beaver lodge debris, in leaf litter on beach, on a pond margin, and in black spruce forest (no specific habitat data). Collecting period: III–XII (mainly V–VII). Collecting method: sifting leaf litter, moss, sphagnum, and other organic debris.

86. Myllaena audax Casey (Fig. 14.120a–i) 87. Myllaena decreta Casey (Fig. 14.121a–h) Myllaena audax Casey, 1911. Klimaszewski 1982, Klimaszewski and Génier 1986, Klimaszewski and Frank 1992, Klimaszewski et al. 2011, 2018. Diagnosis. Body moderately broadly fusiform, length 1.9–2.6 mm, colour dark brown with appendages and tip of abdomen light brown (Fig. 14.120a); head small and deflexed, about half as wide as maximum width of pronotum; antennomeres elongate, VII–X at least twice as long as wide; pronotum broad, maximum width

Myllaena decreta Casey, 1911. Klimaszewski 1982 [list of synonymies]. Diagnosis. Body moderately broadly fusiform, length 1.9–2.6 mm, colour dark brown with appendages and tip of abdomen light brown or yellowish; pubescence uniformly brown; head small and deflexed, about one-third as wide as maximum width of pronotum (Fig. 14.121a); all antennomeres elongate, VII–X about one and a half as long as wide; pronotum convex, rounded

14

Tribe Myllaenini Ganglbauer, 1895

apically, broad, width to length ratio 1.5, maximum width about equal to that of elytra; elytra at suture as long as pronotum along midline of disc; abdomen tapering apically. MALE. Tergite VIII with apical margin slightly acutely triangularly produced, with apex pointed (Fig. 14.121d); sternite VIII broadly parabolically rounded apically and slightly pointed medially (Fig. 14.121e); median lobe of aedeagus in dorsal view with tubus elongate, subparallel, broadly rounded apically (Fig. 14.121c), in lateral view tubus slightly arcuate ventrally with apex broad and rounded (Fig. 14.121b). FEMALE. Tergite and sternite VIII similar to those of male (Fig. 14.121f, g); spermatheca with capsule ovoid, apical invagination large, stem long with about 14–15 tight intertwined coils (Fig. 14.121g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: AZ, CA, NM, OR, UT, WA. COLLECTION AND HABITAT DATA. Habitat: various wetland montaneous habitats; stream and river banks at altitudes 540–10,600 feet. Collecting period: V–VIII. Collecting method: not recorded. 88. Myllaena insomnis Casey (Fig. 14.122a–i) Myllaena insomnis Casey, 1911. Klimaszewski 1982, Klimaszewski and Frank 1992, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2015. Diagnosis. Body moderately broadly fusiform, length 1.9–2.6 mm, colour dark brown with appendages and tip of abdomen light brown; head small and deflexed, about one-third as wide as maximum width of pronotum (Fig. 14.122a); all antennomeres elongate, VII–X at least twice as long as wide; pronotum broad, maximum width about equal to that of elytra; elytra at suture slightly shorter than pronotum along midline of disc; abdomen tapering apically. MALE. Tergite VIII with apical margin slightly acutely

271

triangularly produced, with apex narrowly rounded (Fig. 14.122e); sternite VIII broadly parabolically rounded apically (Fig. 14.122f); median lobe of aedeagus in dorsal view with tubus elongate, subparallel, broadly rounded apically (Fig. 14.122d), in lateral view tubus slightly arcuate ventrally with apex narrow, rounded, slightly curved dorsad (Fig. 14.122b, c). FEMALE. Tergite and sternite VIII similar to those of male (Fig. 14.122g, h); spermatheca with capsule pipe-shaped, apical invagination large, stem long with about 16–24 tight intertwined coils (Fig. 14.122i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK, YT. USA: AK, ID, MA, MI, MN, MT, NY, WI. COLLECTION AND HABITAT DATA. Habitat: various wetland habitats; in NB, along lake margin/dune in flood debris, treading emergent Carex and grasses along a lake margin, on bare mud along margin of slow flowing stream, in moist leaves along vernal pond margin in a mixed forest, calcareous fen in sphagnum moss and litter. Elsewhere, in debris near margins of ponds, streams, rivers, and lakes, in marsh refuse, and treading in a in bog. One from a Malaise trap. Collecting period: III–XI. Collecting method: treading marsh and bog vegetation, sifting various kinds of plant debris, malaise trap. Potawatomi 1982)

Species

Group (Klimaszewski

Diagnosis. Body length 1.6–3.0 mm; antennomeres VII–X elongate, from 1.5 to 2 times as long as maximum width; tergite X of both sexes deeply emarginated, emargination reaching about half length of disc; sternite VIII of both sexes with four pairs of setae; median lobe of aedeagus in lateral view with apex truncate; stem of spermatheca with a single posterior coil or multicoiled. 89. Myllaena fenyesi Bernhauer (Fig. 14.123a–i)

272

Myllaena fenyesi Bernhauer, 1907. Klimaszewski 1982. Lectotype (Female): Cal. [ifornia], Pasadena; fenyesi Bernh., typus; Lectotype M. fenyes Bernh. des. J. Klimaszewski; lectotype teste A. Westrich 2015 GDI Imaging Project (FMNH). Diagnosis. Body moderately broadly fusiform, length 1.9–2.7 mm, colour dark brown with appendages and tip of abdomen light brown or yellowish; pubescence uniformly brownish gray; head small and deflexed, about one-third as wide as maximum width of pronotum (Fig. 14.123a); all antennomeres elongate, VII–X about one and a half as long as wide; pronotum flattened, broad, truncate apically, maximum width about equal to that of elytra, width to length ratio 1.6; elytra at suture as long as pronotum along midline of disc; abdomen tapering apically. MALE. Tergite VIII with apical margin arcuate (Fig. 14.123e); sternite VIII apical margin rounded and slightly produced

14

Tribe Myllaenini Ganglbauer, 1895

medially (Fig. 14.123f); median lobe of aedeagus in dorsal view with tubus elongate, subparallel, broadly rounded apically (Fig. 14.123d), in lateral view tubus slightly arcuate ventrally with apex truncate (Fig. 14.123b, c). FEMALE. Tergite and sternite VIII similar to those of male (Fig. 14.123g, h); spermatheca with capsule spherical, apical invagination not apparent, stem broad and twisted apically (Fig. 14.123i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AZ, CA, NV, OR, UT, WA. COLLECTION AND HABITAT DATA. Habitat: various wetland montaneous habitats; stream and river banks at altitudes 165–3280 m. Collecting period: V–VIII. Collecting method: not recorded. Comments. We have examined 27 specimens from CA and confirm their identity (FMNH).

14

Tribe Myllaenini Ganglbauer, 1895

Fig. 14.120 (a–i) Images of Myllaena audax Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

273

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (b, h) after Klimaszewski (1982). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

274

Fig. 14.121 (a–h) Images of Myllaena decreta Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

14

Tribe Myllaenini Ganglbauer, 1895

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h), after Klimaszewski (1982). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

14

Tribe Myllaenini Ganglbauer, 1895

Fig. 14.122 (a–i) Images of Myllaena insomnis Casey: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e), male tergite VIII; (f) male sternite VIII; (g)

275

female tergite VIII; (h) female sternite VIII; (i) spermatheca. (b) after Klimaszewski (1982). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

276

Fig. 14.123 (a–i) Images of Myllaena fenyesi Bernhauer: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male sternite VIII; (g)

14

Tribe Myllaenini Ganglbauer, 1895

female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c, d) after Klimaszewski (1982). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

14

Tribe Myllaenini Ganglbauer, 1895

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Bernhauer M (1907) Neue Aleocharini aus Nordamerika. (Col.) (3. Stück.). Dtsch Entomol Z 1907(4):381–405 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing, Lancaster, PA, 259 pp Erichson W F (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin. viii + 384 pp Ganglbauer L (1895) Die Käfer von Mitteleuropa. Die Käfer der österreichisch-ungarischen Monarchie, Deutschlands, der Schweiz, sowie des französischen und italienischen Alpengebietes, Zweiter Band, Familienreihe Staphylinoidea, I. Theil: Staphylinidae, Pselaphidae. Carl Gerold’s Sohn, Vienna. vi + 881 pp Klimaszewski J (1982) Studies of Myllaenini (Coleoptera: Staphylinidae, Aleocharinae). 1. Systematics, phylogeny, and zoogeography of Nearctic Myllaena Erichson. Can Entomol 114(3):181–242

277 Klimaszewski J, Frank JH (1992) New distributional data for Nearctic Myllaena Erichson (Coleoptera: Staphylinidae). Supplement 2. Coleopt Bull 46 (4):397–402 Klimaszewski J, Génier F (1986) New distribution and collection data for Nearctic Myllaena Erichson: studies of Myllaenini (Coleoptera: Staphylinidae: Aleocharinae). Supplement 1. Coleopt Bull 40 (1):33–36 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39

Tribe Diglottini Jakobson, 1909

Tribe Diglottini Jakobson, 1909 (Fig. 15.124a–h) (Moore et al. 1973; Ashe 2004; Klimaszewski et al. 2008, 2018) Diagnosis. Body form distinctive with head large, broadly rounded, eyes small, head at least as broad as pronotum, pronotum strongly narrowed posteriorly, elytra shorter than pronotum, abdomen broadly oval, at maximum width broader than elytra; labial palpi very long and stylate; maxillary lobes very long and slender with widely scattered teeth on laciniae; tarsal formula 4-4-5. Comments. Occurs in the intertidal zone of sea beaches where they occur on sand, under stones, burrowed in sand, or among rocks and sand (Moore et al. 1973, Ahn and Jeon 1994, Ahn 1998, Ashe 2004).

Genus Bryothinusa Casey, 1904 (Fig. 15.124a–h) (Casey 1904; Moore 1956; Moore et al. 1973; Moore and Orth 1979; Ahn and Jeon 1994; Ahn 1998; Ashe 2004) Diagnosis. Small beetles, length 2.0–2.5 mm, body shape more or less parallel-sided; colour brown, dark brown, or black; labrum about as long as wide, broadly and evenly rounded in apical outline; mandibles elongate; lacinial setae interdigitating with spinose scales, galea narrower

15

and shorter than lacinia; maxillary palp with three palpomeres, terminal one needle-shaped; cardo very elongate; ligula small and entire; labial palps stylate and elongate; pronotum subquadrate, narrower at base; hypomera visible in lateral view; mesocoxal cavities contiguous, not margined posteriorly; mesosternal process sharply pointed; postero-lateral margins of elytra sinuate; tarsal formula 4-4-5; empodial setae spatulate; small gland opening on tergite VII; and apical lobe of paramere elongate. Members of Bryothinusa live exclusively on sandy beaches (Ahn 1998). Bryothinusa may be readily distinguished from its sister genus Diglotta Champion by having third antennomere very short (almost as long as second one in Diglotta), and shape of the median lobe of aedeagus and spermatheca. Comments. The tribal placement of the genus Bryothinusa is not settled. Liu et al. (2020) included this genus in Myllaenini Ganglbauer. 90. Bryothinusa catalinae Casey (Fig. 15.124a–h) Bryothinusa catalinae Casey 1904. Moore 1956, Moore et al. 1973, Moore and Orth 1979, Ahn 1998, Ahn and Jeon 1994. Diagnosis. Body length 1.9–2.0 mm, colour dark brown to almost black, with paler (yellow brown) elytra, tibiae and tarsi, basal

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_15

279

280

antennomeres and visible mouth parts (Fig. 15.124a); head large, as broad as pronotum, labrum elongate; pronotum trapezoidal, narrowest at base; elytra subquadrate as long as pronotum; hind tarsus with four tarsomeres; forebody with pubescence short and dense; antennae with scape elongate, as long as pedicel, antennomere III–VII subquadrate, VIII–X slightly transverse, XI twice as long as preceding one. MALE. Tergite VIII transverse, broadly and shallowly emarginated apically (Fig. 15.124d); sternite VIII elongate, parabolic apically (Fig. 15.124e); tubus of median lobe of aedeagus sinuate ventrally in lateral view (Fig. 15.124b). FEMALE. Tergite VIII transverse, broadly and shallowly emarginated apically (Fig. 15.124f); sternite VIII rounded apically and truncate medially (Fig. 15.124g); spermatheca with small and spherical capsule attached to tubular neck, stem narrowly tubular, long and irregularly twisted (Fig. 15.124h).

15

Tribe Diglottini Jakobson, 1909

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new generic and specific records]. USA: CA. We report this species from BC for the first time here based on the following: British Columbia: BCol., Vancouver Island, Rathtrevor Provincial Park, 49
190 1800 N, 124
150 4800 W, 25.IV.2016, running on wet sand between rocks, sea beach, A. Davies (CNC) 1 male, 1 female, 7 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: intertidal zone of sea beaches. In BC, adults were found on beach running on wet sand between rocks. In CA, adults and larvae were found beneath and on stones in an association with dense worm tubes, chitons, limpets, small abalones, flat worms, small crabs and brittle stars (Moore and Orth 1979). Collecting period: V, VII. Collecting method: collecting from or between rocks and stones on sea beaches at low tide.

15

Tribe Diglottini Jakobson, 1909

Fig. 15.124 (a–h) Images of Bryothinusa catalinae Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

281

female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

282

References Ahn K-J (1998) Bryothinusa chengae, a new species of the intertidal genus Bryothinusa Casey (Coleoptera: Staphylinidae: Aleocharinae) from the Caroline Islands. Coleopt Bull 52(4):333–338 Ahn K-J, Jeon M-J (1994) The Korean species of the genus Bryothinusa Casey with descriptions of two new species (Coleoptera: Staphylinidae: Aleocharinae). Coleopt Bull 58(1):29–35 Ashe JS (2004) A new species of the intertidal staphylinid genus Bryothinusa Casey from Malaysia, with an overview of geographic distribution and an annotated catalogue of Bryothinusa (Staphylinidae: Aleocharinae: Myllaenini). Coleopt Bull 58(4):581–596 Casey TL (1904) On some new Coleoptera including four new genera. Can Entomol 36:312–324 Jakobson GG (1909) Zhuki Rossii i Zapadnoi Evropy. Part 8, pp 561–640, pls. 62–68. A. F. Devrien, St. Petersburg [in Russian] Klimaszewski J, Webster R, Assing V, Savard K (2008) Diglotta mersa (Haliday) and Halobrecta flavipes Thomson, two new species for the Canadian fauna (Coleoptera, Staphylinidae, Aleocharinae). In: Majka

15

Tribe Diglottini Jakobson, 1909

CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:175–188 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Liu T, Ono H, Maruyama M (2020) A new species and new record of the intertidal rove beetle genus Bryothinusa Casey (Staphylinidae: Aleocharinae: Myllaenini) from Japan. Zootaxa 4743(4):589–593 Moore I (1956) A revision of the Pacific coast Phytosi with a review of the foreign genera (Coleoptera: Staphylinidae). Trans San Diego Soc Nat Hist XII (7):103–152 Moore I, Orth RE (1979) Notes on Bryothinusa with description of the larva of B. catalinae Casey (Coleoptera: Staphylinidae). Psyche 85:183–189 Moore I, Legner E, Chan T (1973) A review of the genus Bryothinusa with description of three new species. Entomol News 84:73–81

Tribe Liparocephalini Fenyes, 1918

Comments. Includes only seashore species.

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– Body slender, weakly to moderately sclerotized, length 1.9–3.4 (Figs. 16.125a and 16.126a) .............................................................3

Tribe Liparocephalini Fenyes, 1918 (Figs. 16.125–16.129) (Ahn and Ashe 1996b; Ahn et al. 2010; Maruyama and Ahn 2000; Ahn 1997; Ashe 2001; Frank and Ahn 2011) Diagnosis. Body densely covered with fine and short setae; elytra very short, at suture distinctly shorter than pronotum, flight wings absent; labial palpus elongate with two palpomeres; ligula long, slender and entire at apex; prementum with one median seta, and with narrow medial pseudopore field; mandibles with denticles in ventral molar area; metaventrite greatly shortened; middle coxae contiguous; tarsal formula 4-4-5 or 4-4-4 in some Diaulota, tarsal claws robust. Includes genera with submarine species found exclusively in the intertidal zones of Pacific shores. Comments. See comments under Bryobiota bicolor (Casey) of the tribe Falagriini about the new genus Ashella Klimaszewski, with Ashella giulanii (Moore) [new combination], not recorded from BC. Key to Genera of Liparocephalini 1. Body robust, heavily sclerotized, length 2.5–5.2 mm (Figs. 16.127a, 16.128a and 16.129a) .............................................................2

2. Head subquadrate, broader than maximum width of pronotum (Figs. 16.128a and 16.129a); infraorbital region of head about as long as 2–3 times width of an eye (Figs. 16.128a and 16.129a); antennomers V–X subquadrate to moderately transverse (Figs. 16.128a and 16.129a); basal abdominal tergite with shallow basal impressions....................Liparocephalus Mäklin – Head elongate, about as wide as maximum width of pronotum (Fig. 16.127a); infraorbital region of head about as long as 3–4 times with of an eye (Fig. 16.127a); antennomers V–X slightly to moderately transverse (Fig. 16.127a); basal abdominal tergite with deep basal impressions................................. Diaulota Casey 3. Eyes vestigial scarcely visible, each consisting of 4–5 ommatidia; forebody pale, yellow brown or red brown and abdomen dark brown (Fig. 16.125a); pronotum narrow, maximum width of pronotum slightly less that maximum width of head, apical corners rounded (Fig. 16.125a); elytra with posterior margin almost straight and slightly sinuate laterally (Fig. 16.125a); body length 1.9–2.2 mm ......................................... Amblopusa Casey – Eyes small but clearly visible, each consisting of multiple ommatidia (Fig. 16.126a); forebody

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_16

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284

reddish with abdomen dark brown or entire body dark brown (Fig. 16.126a); pronotum wide, maximum width of pronotum equals to maximum width of head, apical corners slightly pointed (Fig. 16.126a); elytra with posterior margin arcuate and slightly sinuate laterally (Fig. 16.126a); body length 2.5–3.4 mm ...................................... .............................Paramblopusa Ahn and Ashe

Genus Amblopusa Casey, 1893 (Fig. 16.125a–g) Amblopusa Casey 1893. Casey 1906, 1911, Fenyes 1918, Ahn and Ashe 1996a, Frank and Ahn 2011. Diagnosis. Body small, narrow, subparallel, flattened, length 1.9–2.2 mm; forebody pale, yellow brown and abdomen dark brown except for its paler apex in most; integument with evenly distributed short pubescence, and scattered macrosetae; head elongate, slightly deflexed; eyes vestigial, scarcely visible, each consisting of 4–5 ommatidia; labrum transverse; mandibles with apices acute and slightly turned downward, prosteca membranous, well developed; lacinia and galea elongate; lacinia with a large subapical spine, a row of short subapical spines; maxillary palps with four palpomeres, subapical one swollen apically; labial palps styliform and elongate; ligula elongate, thin, entire at apex; mentum trapezoidal with broadly arcuate apical emargination, apical corners produced, spine-like; pronotum narrow, maximum width of pronotum slightly less that maximum width of head, apical corners rounded; elytra with posterior margin almost straight and slightly sinuate laterally, hind wings absent, abdomen broad at base and sides uniformly converging to rounded apex, three basal tergites strongly impressed at base; tarsal formula 4-4-5. Species confined to the rocky seashores of Pacific coast of North America. 91. Amblopusa brevipes Casey (Fig. 16.125a–g)

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Tribe Liparocephalini Fenyes, 1918

Amblopusa brevipes Casey, 1893. Ahn and Ashe 1996a. Lectotype: USA, Ft. Wrangel, Alaska, Wickham; Amblopusa brevipes Casey; Type USNM 38771; Lectotype Amblopusa brevipes Casey, design. K.J. Ahn and J.S. Ashe 1994; Zerche rev. 1997 (USNM). Images examined. Diagnosis. Body length 1.9–2.2 mm, forebody pale, yellow brown and abdomen dark brown (Fig. 16.125a); antennomeres I and II strongly elongate and of about the same length, IV–V subquadrate, VI–X moderately to strongly transverse, antennae incrassate apically; head elongate, slightly wider than maximum width of pronotum; pronotum subquadrate, narrowest at base and widest in apical third; elytra about one third wider than long along suture, and 0.8 times as long as pronotum, at suture shorter than pronotum; abdomen slightly broadening apically, at base almost as wide as elytra. MALE. Tergite VIII truncate apically, antecostal suture broadly separated from base of disc (Fig. 16.125c); sternite VIII triangularly produced apically, apex rounded (Fig. 16.125d); bulbus of median lobe of aedeagus ovoid, tubus strongly sinuate and bent ventrad apically in lateral view, apex triangular (Fig. 16.125b). FEMALE. Tergite VIII truncate apically, similar to that of male (Fig. 16.125e); sternite VIII slightly produced apically with rounded apex (Fig. 16.125f); spermatheca small, capsule clubshaped, apical invagination not apparent, stem narrow, short, and slightly hooked posteriorly (Fig. 16.125g). DISTRIBUTION. Origin: Nearctic. CANADA: BC [Massett]. USA: AK, CA. COLLECTION AND HABITAT DATA. Habitat: confined to the rocky seashores of Pacific coast of North America. Collecting period: III. Collecting method: not recorded. New locality data: Canada, British Columbia, Port Edward, 28.VI.1968, Campbell and Smetana (CNC) 2 males, 2 females. USA: Alaska, Kenai

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Tribe Liparocephalini Fenyes, 1918

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Pen., 1 mi W Anchor Pt., 1.VI.1978, Smetana and Becker (CNC) 1 male, 3 females. Comments. Ahn and Ashe (1996a) recorded this species from AK and CA in the USA, and in Canada from BC, Queen Charlotte Is., Massett (CAS). Illustration of spermatheca (Ahn and Ashe 1996a) does not correspond to the shape of spermatheca of specimens studied by us from BC and AK.

Genus Paramblopusa Ahn and Ashe, 1996a (Fig. 16.126a–g) (Ahn and Ashe 1996a, b; Ahn and Maruyama 2000; Frank and Ahn 2011) Diagnosis. Body narrowly parallel-sided, flattened; length 2.5–3.4 mm; forebody reddish or dark brown and abdomen dark brown in most; integument with short microsetae about uniformly distributed and macrosetae scattered; head slightly deflexed, broader than remaining parts of body; eyes small, consisting of multiple ommatidia; mandibles with small teeth between apex and median tooth; lacinian setae only on mesal surface; maxillary palpus with four palpomeres; labial palpus styliform; ligula elongate, entire at apex; pronotum wide, maximum width of pronotum equals to maximum width of head, apical corners slightly pointed; elytra with posterior margin arcuate and slightly sinuate laterally; hind wings absent; abdomen broadening apically, basal three abdominal tergites strongly impressed at base, anterior margins of abdominal tergites III–VI straight, not emarginate; tarsal formula 4-4-5. Species confined to intertidal zones of Pacific. 92. Paramblopusa (Fig. 16.126a–g)

borealis

(Casey)

Amblopusa borealis (Casey, 1906). Moore and Legner 1975, Ahn and Ashe 1996a, b, Ahn and Maruyama 2000. Holotype (male): Canada, British Columbia: QCI [Queen Charlotte Is., Massett in orig. desc.];

borealis Csy.; Type USNM 38773; Casey bequest 1925; Lectotype Amblopusa borealis Casey, des. K.J. Ahn and J.S. Ashe, 1994 (USNM). Examined. Diagnosis. Body narrowly linear, length 2.5–3.4 mm, colour variable, reddish brown with head and most of abdomen dark brown in most; pubescence uniformly yellow brown; head large, slightly deflexed, at least as broad as maximum width of pronotum (Fig. 16.126a); antennomeres I–II strongly elongate, IV subquadrate, V–X subquadrate to slightly transverse; head large, transverse; eyes small but distinct with multiple ommatidia; pronotum trapezoidal in shape slightly elongate, maximum width of pronotum equals to maximum width of head, apical corners slightly pointed; elytra with posterior margin arcuate and slightly sinuate laterally; abdomen broadening apically. MALE. Tergite VIII with apical margin broadly arcuate (Fig. 16.126c); sternite VIII with apical margin broadly triangularly produced medially, apex pointed (Fig. 16.126d); bulbus narrowly ovoid, tubus of median lobe of aedeagus strongly bent ventrad in lateral view, apex broad (lancetshaped) and sharply pointed ventrad (Figs. 16.126b). FEMALE. Tergite VIII truncate apically (Fig. 16.126e); sternite VIII triangularly produced apically (Fig. 16.126f); spermatheca small and slender, capsule spherical connected to straight neck, stem thin, broadly coiled and twisted posteriorly (Fig. 16.126g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, OR, WA. New locality data: British Columbia, Vancouver Is., Long Beach 12 mi S Tofino, 22/23.V.1968, Campbell and Smetana (CNC) 1 male, 1 female. COLLECTION AND HABITAT DATA. Habitat: confined to beaches of Pacific coast; found on flats under rocks. Collecting period: V, VIII, X. Collecting method: hand collecting from organic debris on beaches.

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Comments. Casey (1906) specified in the original description that he described this species on a single specimen with broken antennae and therefore it constitutes a holotype. Ahn and Ashe (1996a) recorded this species from BC: Massett (CAS) 4 specimens; Caspaco, Skeen River estuary, 11.VIII.1973, G. Schulte and Beaver (SEMKU) 2 specimens; and Harbor, 13. IX.1970, W.G. Evans (SEMKU) 2 specimens.

Genus Diaulota Casey, 1893 (Fig. 16.127a–h) Diaulota Casey, 1893. Ahn and Ashe 1996b, Frank and Ahn 2011. Diagnosis. Body robust, narrowly elongate, length 2.5–2.9 mm; colour variable, black, dark brown, reddish brown, or brown; integument with relatively long and evenly distributed microsetae; head slightly deflexed, 0.8–1.2 times as long as wide; eyes small, 0.2 times length of head, tempora very long, neck absent; antennomeres V–X slightly to moderately transverse; labrum semicircular; mandibles asymmetrical or nearly symmetrical, apex acute and turned downward, right mandible with median tooth, internal teeth absent, prostheca well developed; maxillary palps with 4 palpomeres; labial palpus with 2–3 palpomeres; ligula simple, elongate; mentum without V seta; elytra shorter than pronotum; hind wings absent; abdomen enlarged, sides arcuate, broader than maximum width of head, tergites III–VI impressed at base; sternite not constricted at base; tarsal formula 4-4-5, 4-4-4, or 3-3-4. Species confined to the intertidal zone of rocky shores of Pacific. 93. Diaulota densissima Casey (Fig. 16.127a–h) Diaulota densissima Casey, 1893. Ahn and Ashe 1996b.

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Tribe Liparocephalini Fenyes, 1918

Diagnosis. Body robust, heavily sclerotized, length 2.5–2.9 mm, colour black or dark brown with appendages and tip of abdomen often reddish brown (Fig. 16.127a); pubescence uniformly brownish gray; head large, elongate, as long as pronotum, at base about as broad as maximum width of pronotum; antennomeres VII–X strongly transverse; pronotum elongate, trapezoidal in shape, broadest apically, about as long as head; elytra 1.1 times as long as wide, 0.7 times as long as pronotum; abdomen large, broader than forebody, tergite impression shallow, paratergites evenly narrow. MALE. Tergite VIII with apical margin truncate, slightly emarginate medially (Fig. 16.127d); sternite VIII with apical margin strongly sinuate laterally and broadly triangularly produced medially, apex rounded (Fig. 16.127e); median lobe of aedeagus with bulbus narrowly ovoid, tubus moderately produced ventrad in lateral view (Fig. 16.127b), in dorsal view median lobe as illustrated (Fig. 16.127c). FEMALE. Tergite VIII truncate apically and shallowly emarginated medially (Fig. 16.127f); sternite VIII rounded apically (Fig. 16.127g); spermatheca small, capsule spherical, apical invagination not apparent, stem narrowly elongate and sinuate (Fig. 16.127h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, OR, WA. New locality data: British Columbia, Queen Charlotte Is., Hot Springs Is., 11.VIII.1983, 83-73, ex intertidal rocks, J.M. Campbell (CNC) 30 sex undetermined; BC, Queen Charlotte Is., Tanu Is., Tanu, 9.VIII.1983, ex intertidal rocks, J.M. campbell (CNC) 15 sex undetermined; BC, Queen Charlotte Is., Moresby Is., Kaisun, 1. VIII.1983, 83-48, 100 , intertidal barnacles, J.M. Campbell (CNC) 51 sex undetermined, 6 larvae; BC, Long Beach, 12 mi S Tofino, 22/23. V.1968, Campbell and Smetana (CNC) 1 female, 7 sex undetermined.

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Tribe Liparocephalini Fenyes, 1918

COLLECTION AND HABITAT DATA. Habitat: confined to the rocky seashores of Pacific coast of North America, found in rock crevices of intertidal zones; ex Fucus zone; in wrack, in rocky shore with mud and sand, between green algae in oligohaline water; in rocky shore, euhaline water, between Fucus, barnacles and lichens; in vertical crevices of rocks in splash zone (Ahn and Ashe 1996b). Collecting period: I–IX. Collecting method: hand collecting from rock crevices in intertidal zones.

Genus Liparocephalus Mäklin, 1853 (Figs. 16.128 and 16.129) (Ahn and Ashe 1996b; Maruyama and Ahn 2000; Ahn 1997; Ahn et al. 2010; Frank and Ahn 2011) Diagnosis. Body length 3.8–5.2 mm, robust, heavily sclerotized, integument with moderately long setae densely and uniformly distributed; head slightly deflexed, large and robust; antennomeres V–X subquadrate to slightly transverse; eyes small, 0.2 times length of head, and without setae between facets; labrum transverse; mandibles almost symmetrical with 4–5 teeth between apex and median tooth; maxillary palpi with four palpomeres; labial palpi with two palpomeres; ligula simple, elongate; pronotum trapezoidal in shape, broadest apically, about size of head; elytra reduced to small rounded lobes; membranous wings absent; abdomen broad and robust, sides arcuate, abdominal tergites without deep transverse impressions; tarsi with large spatulate setae; tarsi with spatulate setae, claws narrow, long and sickle-shaped, tarsal formula 4-4-5. Male sternite VIII prolonged apically as broad triangular projection. Includes submarine species confined to the rocky intertidal zones of Pacific shores. Key to Species of Liparocephalus 1. Maximum width of head about equal to maximum width of pronotum (Fig. 16.128a); antennomeres VII–X subquadrate to slightly transverse (Fig. 16.128a); pronotum subquadrate to slightly transverse (Fig. 16.128a); abdominal paratergites narrow anteriorly and broadening

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posteriorly forming broadly serrate lateral margin of abdomen (Fig. 16.128a); apical projection of male tergite VIII broadly triangular and rounded at apex (Fig. 16.128d) .......................................... ...................Liparocephalus brevipennis Mäklin – Maximum width of head distinctly greater than maximum width of pronotum (Fig. 16.129a); antennomeres VII–X subquadrate to slightly elongate (Fig. 16.129a); pronotum transverse (Fig. 16.129a); abdominal paratergites evenly narrow and lateral margin of abdomen subparallel (Fig. 16.129a); apical projection of male tergite VIII narrowly triangular and slightly pointed at apex (Fig. 16.129f).............................. ...................Liparocephalus cordicollis LeConte

94. Liparocephalus (Fig. 16.128a–g)

brevipennis

Mäklin

Liparocephalus brevipennis Mäklin, 1853. Ahn 1997. Diagnosis. Body robust, heavily sclerotized, length 4.2–4.4 mm, colour black or dark brown with appendages and tip of abdomen often reddish brown (Fig. 16.128a); pubescence uniformly brownish gray; head large, slightly deflexed, about as wide as maximum width of pronotum; antennomeres VII–X slightly transverse; pronotum trapezoidal in shape, broadest apically, about as long as wide; elytra reduced to two small lobes, at suture much shorter than pronotum; abdomen large, tergite impression scarcely visible, paratergites narrow anteriorly and broadening posteriorly creating distinct broadly serrate margin. MALE. Tergite VIII with apical margin arcuate and truncate medially (Fig. 16.128c); sternite VIII with apical margin strongly sinuate laterally and broadly triangularly produced medially (Fig. 16.128d); median lobe of aedeagus with bulbus large and ovoid, tubus narrow and strongly bent ventrad in lateral view (Fig. 16.128b). FEMALE. Tergite VIII similar to that of male (Fig. 16.128e); sternite VIII slightly produced and pointed at apex (Fig. 16.128f); spermatheca with capsule spherical, apical invagination not apparent, stem narrowly elongate and slightly sinuate (Fig. 16.128g).

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DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: confined to the rocky intertidal zones of Pacific shores, found under rocks, in rock crevices on mud flat at low tide; in rocky shore, among Enteromorpha, Fucus and barnacles. Collecting period: V–IX. Collecting method: hand collecting from rock crevices in intertidal zones. 95. Liparocephalus (Fig. 16.129a–i)

cordicollis

LeConte

Liparocephalus cordicollis LeConte, 1880. Moore and Legner 1975, Ahn 1997. Holotype (male): USA, California, Mendocino, Mr. A. Agassiz [in orig. description]: Mendocino; L. cordicollis; Type 6583; MCZ-ENT 00006583 (MCZ). Examined. Diagnosis. Body robust, heavily sclerotized, length 3.6–5.2 mm, colour black or dark brown with appendages and tip of abdomen often reddish brown, some females have reddish brown pronotum or head and pronotum; pubescence uniformly brownish gray (Fig. 16.129a, b); head large (largest in male than in female), slightly deflexed, distinctly broader than maximum width of pronotum; antennomeres VII–X subquadrate to slightly elongate; pronotum trapezoidal in shape and constricted basally, broadest apically, transverse (more transverse in male than in female); elytra reduced to two small lobes, at suture much shorter than pronotum; abdomen large, tergite impression scarcely visible, paratergites evenly narrow. MALE. Tergite VIII with apical margin arcuate (Fig. 16.129e); sternite VIII with apical margin strongly sinuate laterally and narrowly triangularly produced medially, apex pointed (Fig. 16.129f); median lobe of aedeagus with bulbus large and ovoid, tubus narrow, arcuate, and strongly bent ventrad in lateral view (Fig. 16.129c), in dorsal view median lobe as illustrated (Fig. 16.129d). FEMALE. Tergite VIII broadly arcuate apically (Fig. 16.129g); sternite VIII triangularly produced and rounded at apex (Fig. 16.129h);

Tribe Liparocephalini Fenyes, 1918

spermatheca very small and usually found inside pygidium, capsule spherical, apical invagination present, stem narrowly elongate and slightly sinuate (Fig. 16.129i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, OR, WA. New locality data: Canada, British Columbia: Queen Charlotte Is., Liebeck coll., barcode 00681844 (MCZ) 1 female; Queen Charlotte Is., J. Fletcher, brevipennis, barcode 00681839 (MCZ) 1 female; Queen Charlotte Is., brevipennis, J. Fletcher, barcodes 00681838, 00681840, 00681837 (MCZ) 1 male, 2 sex undetermined; Queen Charlotte Is., Fredrick Blanchard coll., barcode 00681841 (MCZ) 1 sex undetermined; Queen Charlotte Is., Massett, 33, Liebeck coll., barcode 00681843 (MCZ) 1 sex undetrmined; Queen Charlotte Is., Graham Is., Massett, Keen leg. (FMNH) 2 males, 3 females, 8 sex undetermined; Queen Charlotte Is., Moresby Is., Kaisun, 1.VIII.1983, 83-48, 100 , ex intertidal barnacles, J.M. Campbell (CNC) 29 sex undetermined. USA: Oregon, Charleston, 7-2-47, 6-23-47, intertidal, G.H. Nelson coll., C.A. Frost coll. 1962, barcodes 00681833, 00681834, 00681835, 00681836 (MCZ) 1 male, 1 female, 2 sex undetermined; Oregon, Squaw Is., Coos Co., 19.VI.1947, I.M. Newell, CNHM Borys Malkin coll. 1960 (FMNH) 2 females; Oregon, Winchester Bay, Lane Co., 13.IV.1947, B. Malkin and I.M. Newell, CNHM Borys Malkin coll. 1960 (FMNH) 1 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: confined to intertidal zones of Pacific shores; found on Egregia, captured from just above Porphyra, in mid-tide crevices, on low-tide intertidal rocks, among barnacles and mussels, wrack in intertidal zones. Collecting period: II, V–VIII. Collecting method: hand collecting from rocks in intertidal zones. Comments. The spermatheca illustrated by Ahn (1997) does not belong to this species.

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Tribe Liparocephalini Fenyes, 1918

Fig. 16.125 (a–g) Images of Amblopusa brevipes Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

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(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

290

Fig. 16.126 (a–g) Images of Paramblopusa borealis (Casey): (a) habitus in dorsal view [holotype, broken antennae reconstructed]; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

16

Tribe Liparocephalini Fenyes, 1918

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. (a–d) holotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

16

Tribe Liparocephalini Fenyes, 1918

Fig. 16.127 (a–h) Images of Diaulota densissima Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

291

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

292

Fig. 16.128 (a–g) Images of Liparocephalus brevipennis Mäklin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

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Tribe Liparocephalini Fenyes, 1918

male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Liparocephalini Fenyes, 1918

Fig. 16.129 (a–i) Images of Liparocephalus cordicollis LeConte: (a), habitus in dorsal view; (b) color variation of pronotum and elytra; (c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male

293

tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (a, c, e, f) holotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

294

References Ahn K-J (1997) A review of Liparocephalus Mäklin (Coleoptera: Staphylinidae: Aleocharinae) with descriptions of larvae. Pan-Pac Entomol 73(2):79–92 Ahn K-J, Ashe JS (1996a) Revision of the intertidal aleocharine genus Amblopusa Casey and description of the new genus Paramblopusa (Coleoptera: Staphylinidae). J NY Entomol Soc [1995] 103 (2):138–154 Ahn K-J, Ashe JS (1996b) Phylogeny of the intertidal aleocharine tribe Liparocephalini (Coleoptera: Staphylinidae). Syst Entomol 21:99–114 Ahn K-J, Maruyama M (2000) Paramblopusa eoa, a new intertidal species (Coleoptera: Staphylinidae: Aleocharinae) from Kuril Islands. Coleopt Bull 54 (3):359–364 Ahn K-J, Jeon M-J, Branham MA (2010) Phylogeny, biogeography and the stepwise evolutionary colonization of intertidal habitat in the Liparocephalini based on morphological and molecular characters (Coleoptera: Staphylinidae: Aleocharinae). Cladistics 26:344–358 Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC: https://www.crcpress.com/Ameri can-Beetles-Volume-I-Archostemata-MyxophagaAdephaga-Polyphaga/Jr-Thomas/p/book/ 9780849319259]

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Tribe Liparocephalini Fenyes, 1918

Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7:281–606 [see reference in Chapter 7 [1892–1894] for explanation of date] Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St. Louis 16(6):125–434 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp. 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, PA, 259 pp Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera insectorum, vol XXVII, Fascicule 173A. M. Nijhoff, Den Haag, 453 pp + 7 pls Frank JH, Ahn K-J (2011) Coastal Staphylinidae (Coleoptera): a worldwide checklist, biogeography and natural history. ZooKeys 107:1–98 LeConte JL (1880) Short studies of North American Coleoptera. Trans Am Entomol Soc 12:163–218 Mäklin FW (1853) [New species and notes]. In: Mannerheim CG: Dritter Nachtrag zur KaeferFauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Impériale Nat Moscou 26(3):95–273 Maruyama M, Ahn K-J (2000) Redescription of Liparocephalus litoralis, and key to the species of Liparocephalus (Coleoptera: Staphylinidae: Aleocharinae). Can Entomol 132(5):567–571 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514

Tribe Autaliini C. G. Thomson, 1859

Tribe Autaliini C. G. Thomson, 1859 (Figs. 17.130 and 17.131) (Ashe 2001) Diagnosis. Body somewhat ant-shaped, length 1.7–2.1 mm; head with neck narrow, less than one-third width of head; pronotum with four ante-basal foveae; elytra about two times as broad as pronotum, with two basal foveae; abdomen narrowly elongate.

Genus Autalia Leach, 1819 (Figs. 17.130 and 17.131) (Hoebeke 1985, 1988; Assing 1997) Diagnosis. Body somewhat ant-shaped, length 1.3–3.7 mm; head with neck narrow, less than one-third width of head; ligula dichotomously branched into four divergent and pointed processes; pronotum with four ante-basal foveae; elytra almost twice as wide as pronotum, with two basal foveae; abdomen narrowly elongate, first three visible tergites with deep transverse impressions, each with large medial carina and smaller lateral carinae. Adults are capable of flight, associated with fungi and various decaying matter, at least some species are known as predators.

17

median furrow of pronotum long, deep, reaching behind to basal transverse groove (Fig. 17.131a); tubus of median lobe of aedeagus and spermatheca as illustrated (Fig. 17.131b, c, h)........................... ........................... Autalia rivularis (Gravenhorst) – Body dark brown, often with reddish tinge and paler appendages, length 2.7–3.0 mm (Figs. 17.130a and 17.132a); median furrow of pronotum short, distinct apically but obsolete basally, not reaching behind to basal transverse groove, or furrow obsolete (Figs. 17.130a and 17.132a); median lobe of aedeagus and spermatheca differently shaped .....................................2 2. Punctures of pronotum coarse, dense, noticeable asperate (Fig. 17.130a); apex of tubus of median lobe of aedeagus long and gradually and gently turned ventrad in lateral view (Fig. 17.130b); spermathecal with stem slightly bent (Fig. 17.130h)... Autalia puncticollis Sharp – Punctures of pronotum less coarse, sparse, not asperate (Fig. 17.132a); apex of tubus of median lobe of aedeagus short and strongly angularly turned ventrad in lateral view (Fig. 17.132b); spermatheca with stem strongly bent (Fig. 17.132h)............. Autalia truncatula Casey

Key to Species of Autalia

96. Autalia { (Fig. 17.130a–h)

puncticollis

1. Body black, length 1.7–2.1 mm; pronotum with very fine and sparse punctation (Fig. 17.131a);

Autalia puncticollis Hoebeke 1988.

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_17

Sharp,

Sharp

1864.

295

296

Diagnosis. Body form distinctive, somewhat ant-shaped (Fig. 17.130a), length 2.8–3.0 mm; colour dark brown, central part of elytra and basal tergites with reddish tinge, appendages yellowish red or light brown; coarse, dense, noticeable asperate; head distinctly wider than pronotum, semicircularly rounded behind eyes; pronotum widest at apical third, distinctly narrower than elytra, disc with median furrow of pronotum short, distinct apically but obsolete basally, not reaching behind to basal transverse groove, or furrow obsolete; elytra at suture distinctly longer than pronotum; abdomen wide, slightly swollen at apical third, about as wide as elytra, three first visible tergites with impressions deep, with longitudinal carinae; antennomeres IV–X slightly transverse. MALE. Tergite VIII truncate apically and concave medially, antecostal suture broadly separated from basal margin (Fig. 17.130d); sternite VIII broadly triangularly produced at apex, apex rounded (Fig. 17.130e); median lobe of aedeagus in dorsal view with tubus gradually broadening to near apex (Fig. 17.130c); in lateral view tubus curved obliquely ventrad from base and triangular apex pointing ventrad, bulbus with base elongate, sharply curled toward apex of tubus (Fig. 17.130b). FEMALE. Tergite VIII with apical margin truncate and shallowly emarginate medially (Fig. 17.130f); sternite VIII with apical margin broadly arcuate (Fig. 17.130g); spermatheca with capsule spherical, stem short and slightly bent near capsule, tapering to base (Fig. 17.130h). DISTRIBUTION. Origin: Palearctic, adventive in Canada and USA. CANADA: BC. USA: CA, WA. COLLECTION AND HABITAT DATA. Habitat: habitat is similar to that of A. rivularis; it has been collected in various decaying organic matter, predominantly in droppings of large mammals but also found on fungi and carrion (Assing 1997). Collecting period: not recorded in Nearctic region. Collecting method: sifting organic matter, pitfalls.

17

Tribe Autaliini C. G. Thomson, 1859

97. Autalia rivularis { (Fig. 17.131a–h)

(Gravenhorst)

Aleochara rivularis Gravenhorst, 1802. Assing 1997, Smetana 2004, Hoebeke 1988, Klimaszewski et al. 2005, 2011, 2013, 2016, 2018, Majka and Klimaszewski 2008. Diagnosis. Body form distinctive, somewhat ant-shaped (Fig. 17.131a), length 1.7–2.1 mm, colour dark brown to black; head slightly wider than pronotum, semicircularly rounded behind eyes; pronotum widest at apical third, narrower than elytra, median furrow of pronotum long, deep, reaching behind to basal transverse groove, punctation fine and sparse; elytra at suture longer than pronotum; abdomen wide, slightly swollen at apical third, about as wide as elytra, three first visible tergites with impressions deep, with longitudinal carinae; antennomeres IV–X quadrate to slightly transverse. MALE. Tergite VIII truncate apically, slightly concave medially, antecostal suture broadly separated from basal margin (Fig. 17.131d); sternite VIII triangularly produced at apex, apex pointed (Fig. 17.131e); median lobe of aedeagus in dorsal view with tubus gradually broadening to near apex (Fig. 17.131c); in lateral view tubus curved obliquely ventrad from base and angularly bent at apical third, so that triangular apex actually pointing obliquely posteriad, bulbus with base elongate, sharply curled toward apex of tubus (Fig. 17.131b). FEMALE. Tergite and sternite VIII with apical margin broadly shallowly emarginate medially (Fig. 17.131f, g); spermatheca with capsule spherical, stem short and slightly curved, tapering to base (Fig. 17.131h). DISTRIBUTION. Origin: Palearctic, adventive in Canada and USA. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK. USA: CA, ME, MI, NH, NJ, NY, OR, PA, VT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mixed forests, hardwood woodland, red spruce regenerating forest, coastal shrubby barrens, and

17

Tribe Autaliini C. G. Thomson, 1859

pastures. In NB, adults abundant in horse dung in a pasture; also found in decaying mushrooms, and in drift material on lakeshore. Flying adults collected late in the afternoon by sweeping with butterfly net. Elsewhere, found in leaf litter, compost piles, cow droppings, and other decomposing organic matter. Collecting period: V–IX. Collecting method: sifting dung, leaf litter, decaying mushrooms, sweeping with butterfly net, flight intercept traps, pitfall traps. 98. Autalia truncatula Casey, 1911 (Fig. 17.132a–h) Autalia truncatula Casey, 1911. Hatch 1957, Moore and Legner 1975, Hoebeke 1988, Klimaszewski and Winchester 2002. Diagnosis. Body form distinctive, somewhat ant-shaped (Fig. 17.132a), length 2.7–3.0 mm; colour dark brown with rust tinge, and often paler, yellowish brown lateral margins of pronotum, central part of elytra, and appendages; punctation fine, sparse and not asperate; head slightly wider than pronotum, semicircularly rounded behind eyes; pronotum widest at apical third, narrower than elytra, disc with two basolateral sulci, with slightly depressed areas near posterior angles, median furrow of pronotum short, distinct apically but obsolete basally, not reaching behind to basal transverse groove; elytra

297

at suture longer than pronotum; abdomen wide, slightly swollen at apical third, about as wide as elytra, three first visible tergites with impressions deep, with longitudinal carinae; antennomeres IV–X slightly transverse. MALE. Tergite VIII truncate apically and concave medially, antecostal suture broadly separated from basal margin (Fig. 17.132d); sternite VIII broadly triangularly produced at apex, apex slightly pointed (Fig. 17.132e); median lobe of aedeagus in dorsal view with tubus gradually broadening to near apex (Fig. 17.132c); in lateral view tubus curved obliquely ventrad from base and angularly bent at apical third, so that triangular apex pointing ventrad, bulbus with base elongate, sharply curled toward apex of tubus (Fig. 17.132b). FEMALE. Tergite VIII with apical margin truncate (Fig. 17.132f); sternite VIII with apical margin broadly arcuate (Fig. 17.132g); spermatheca with capsule spherical, stem short and slightly curved, tapering to base (Fig. 17.132h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA. COLLECTION AND HABITAT DATA. Habitat: Sitka spruce forest in Carmanah Valley, Vancouver Island; most specimens were captured in transition zone followed by forest interior, during fall season. Collecting period: VI–IX. Collecting method: pitfall traps.

298

Fig. 17.130 (a–h) Images of Autalia puncticollis Sharp: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

17

Tribe Autaliini C. G. Thomson, 1859

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

17

Tribe Autaliini C. G. Thomson, 1859

Fig. 17.131 (a–h) Images of Autalia rivularis (Gravenhorst): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

299

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

300

Fig. 17.132 (a–h) Images of Autalia truncatula Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

17

Tribe Autaliini C. G. Thomson, 1859

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

17

Tribe Autaliini C. G. Thomson, 1859

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 acccording to CRC] Assing V (1997) Review of the Palearctic species of Autalia Leach in Samouelle, 1819 (Coleoptera, Staphylinidae, Aleocharinae). Entomol Bl Biol Syst Käfer 93(1):69–85 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp. 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing Co., Lancaster, Pennsylvania, 259 pp Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Hatch MH (1957) The beetles of the Pacific Northwest. Part II: Staphyliniformia. University of Washington Publications in Biology 16:IX + 384 pp Hoebeke ER (1985) A revision of the rove beetle tribe Falagriini of America north of Mexico (Coleoptera: Staphylinidae: Aleocharinae). J N Y Entomol Soc 9 (2):913–1018 Hoebeke ER (1988) A new species of rove beetle, Autalia phricotrichosa (Coleoptera: Staphylinidae: Aleocharinae) from Mexico, with a key to the new world species of Autalia. Coleopt Bull 42(1):87–93 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belge Entomol 40:3–126 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp

301 Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013) Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia-Moscow, 360 pp Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Leach WE (1819) New genera. In: Samouelle (ed) The entomologist’s useful compendium; or an introduction to the knowledge of British insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; etc. Thomas Boys, London, 496 pp + 12 pls Majka CG, Klimaszewski J (2008) Introduced Staphylinidae (Coleoptera) in the Maritime Provinces of Canada. Can Entomol 140(1):48–72 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Sharp D (1864) New taxon. J Proc Entomol Soc Lond 1864:45 Smetana A (2004) Family Staphylinidae Latreille, 1802 New nomenclatorial and taxonomic acts, and comments [pp 29–35], subfamilies OmaliinaeDasycerinae [pp 237–272], subfamilies Phloeocharinae-Apateticinae [pp 329–495], subfamilies Piestinae-Staphylininae [pp 504–698]. In: Löbl I, Smetana A (eds) Catalogue of Palearctic Coleoptera, vol 2. Hydrophiloidea – Histeroidea – Staphylinoidea. Apollo Books, Stenstrup, 942 pp Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom. Berlingska Boktryckeriet, Lund, [3] + 290 pp

Tribe Homalotini Heer, 1839

Tribe Homalotini Heer, 1839 (Figs. 18.133–18.148) Klimaszewski et al. 2018)

(Ashe

2001;

Diagnosis. Body of diverse forms, pronotum and elytra without fovea and grooves; tarsal formula 4-4-5; mandibles with minute teeth in ventral molar region; prementum with bases of medial setae very close together, medial pseudopore field very narrow; ligula broadly rounded, completely split (divided from apex to base) or bifid (divided only apically); mesocoxae narrowly to broadly separated.

18

3. Eyes small in most and not protruding from outline of head (Fig. 18.133a); median lobe of aedeagus with distinct wide and rigid flagellum (Fig. 18.133b, c)......................... Leptusa Kraatz – Eyes medium-sized and protruding from outline of head (Fig. 18.135a); ligula slender and split apically (Fig. 18.134f); median lobe of aedeagus with thin and weakly sclerotized flagellum (Fig. 18.135b) ..................... Leptusomorpha Klimaszewski and Hoebeke, gen. n.

1. Labial palpi extremely elongate, extending beyond labrum, forming a V-shape structure (Fig. 18.148a)............................ Silusa Erichson – Labial palpi not as above ..............................2

4. Mesocoxae separated by less than width of mesocoxal cavity by moderately broad or narrow processes of meso- and metaventrite; pronotum usually moderately glossy, with punctation and pubescence moderately dense (Figs. 18.136a– 18.139a) ...................................... Stictalia Casey – Mesocoxae separated by about width of mesocoxal cavity by broad and broadly contiguous processes of meso- and metaventrite; pronotum moderately or highly glossy, with punctation and pubescence sparse and dispersed (Figs. 18.140a–18.145a) ....................................5

2. Body subparallel, moderately convex (Figs. 18.133a and 18.135a); integument usually asperate and often granulose on pronotum (Figs. 18.133a and 18.135a); pronotum widest in apical one-third (Figs. 18.133a and 18.135a); mesocoxae narrowly separated ..........................3 – Characters not as above or in different combinations ......................................................4

5. Body small, length 1.9–2.0 mm, broadly oval in dorsal outline, robust, very compact, abdominal tergites short and tight (Fig. 18.140a); anterolateral margin of pronotum strongly deflexed ............................ Encephalus Stephens – Body medium sized, length over 2 mm in most, ovoid or elongate in outline, moderately compact (Figs. 18.141a–18.145a), other

Comments. Structurally diverse group with many species associated with fungi. Key to Genera of Homalotini

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_18

303

304

characters not as above ........................................ .................................. Gyrophaena Mannerheim

Subtribe Bolitocharina C. G. Thomson, 1859 Diagnosis. Integument often coarsely asperate; body usually convex, sometimes subcylindrical; labial palpi with three palpomeres, not styliform, basal two palpomeres indistinctly separated in Leptusa; ligula slender and elongate; mandibles with large, dense patch of denticles in ventral molar region; prementum with medial setae arranged one behind the other, medial “pseudopore field” of prementum narrow, without pseudopores; male tergites VII and VIII often with secondary sexual structures consisting of carinae or knobs, tergite VIII often emarginate apically and bearing teeth or denticles (Seevers 1978; Ashe 2001). Comments. Adults and larvae occur in association with macrofungi, especially the Polyporaceae and lignicolous Agaricales (Ashe 2001).

Genus Leptusa Kraatz, 1856 (Fig. 18.133a–h) (Pace 1989; Klimaszewski et al. 2004, 2018) Diagnosis. Body convex, subparallel, integument usually with asperate punctation; pronotum often granulose, broadest at apical one-third; elytra variable in length, sometimes shorter than pronotum; abdomen subparallel or broadening apically; eyes small not protruding from outline of head; antenna with three basal antennomeres elongate, remaining ones quadrate to transverse; ligula slender and entire; maxillary palpi with four palpomeres, apical palpomere narrow, about one-third length of penultimate one; labial palpi with three palpomeres but basal two indistinctly separated, last one narrow, glossae slender and entire; mandibles robust with small molar areas; labrum transverse; mesocoxae narrowly separated by narrow and pointed process of

18

Tribe Homalotini Heer, 1839

mesoventrite which extends about two-thirds length of mesocoxae; tarsal formula 4-4-5, hind tarsus with basal article about as long as two following ones together or slightly longer; median lobe of aedeagus with flagellum broadly tubular, predominantly shorter than length of median lobe, rarely long and coiled within bulbus (subgenus Adoxopisalia), internal sac with two well-defined pairs of basal and apical sclerites, tubus subparallel and moderately long; spermatheca with capsule spherical or ovoid, stem usually narrow and short. Comments. Adults occur on fungi or under bark (sometimes in association with bark beetles), and in forest litter. We have seen the guts of some Canadian species filled with fungal flesh. At least some species overwinter in the adult stage in bark beetle galleries or under bark. Species are known from coniferous and hardwood forests. Subgenus Gatineauleptusa Klimaszewski, 2018 (Fig. 18.133a–h) (Klimaszewski et al. 2018) Diagnosis. Body narrowly subparallel; forebody sparsely pubescent, with integument deeply coarsely punctate; antennomeres IV–X transverse and incrassate, VII–X distinctly transverse, each antennomere at least three times as wide as long; male tergite VIII truncate and crenulate apically; median lobe of aedeagus with apical sclerites of internal sac not apparent, basal sclerites in form of two elongate structures, apical part of membrane of internal sac with minute denticles, flagellum short and thick, forming long ovoid base; spermathecal of a unique mushroom-shaped, capsule kidney-shaped, without apparent invagination, stem long, thin, with one posterior loop. Evenly narrow body (head, pronotum, elytra and abdomen of about the same width), pronounced body microsculpture, elytra with setae on short, transverse line-like elevated structures, and the unique shape of spermatheca define this subgenus.

18

Tribe Homalotini Heer, 1839

99. Leptusa (Gatineauleptusa) gatineauensis Klimaszewski and Pelletier (Fig. 18.133a–h) Leptusa gatineauensis Klimaszewski and Pelletier, in Klimaszewski et al. 2004, 2011, 2016, 2018, Bishop et al. 2009, McLean et al. 2009, Majka and Klimaszewski 2008, Webster et al. 2012. Diagnosis. Body length 1.5–1.9 mm, colour brown, head and abdomen darker than remainder of body, elytra and pronotum with dark reddish tinge, antennae dark yellowish-brown (Fig. 18.133a); integument with punctation moderate in size, with distinct isodiametric microsculpture, pronotum and elytra slightly granulose; antenna with antennomeres I–III distinctly elongate, III a little shorter and slimmer than II, IV subquadrate, V–X transverse and VII– X distinctly so, each about three times as broad as long; pronotum transverse, about 1.4 times as wide as long, with sides gradually converging basad and broadest about middle of its length, shorter than elytra at suture, punctation moderate in size, distance between punctures greater than diameter of a puncture; elytra slightly transverse, at suture about 1.2 times as long as pronotum; abdomen subparallel, with first three visible tergites deeply impressed basally and punctate. MALE. Tergite VIII with apical margin broadly truncate medially, minutely crenulate (Fig. 18.133d); sternite VIII with apical margin very obtusely produced, broadly arcuate medially (Fig. 18.133e); median lobe of aedeagus in lateral view with crista apicalis of bulbus narrowly oval, tubus moderately elongate, ventral margin curved ventrad in basal half, distinctly sinuate to narrow and narrowly rounded apex (Fig. 18.133b, c); internal sac with flagellum shorter than median lobe, lamina basalis in form of two elongate structures, lamina apicalis inconspicuous, with minute denticles (Fig. 18.133b, c). FEMALE. Tergite VIII truncate apically, antecostal suture sinuate, moderately narrowly separated from basal margin (Fig. 18.133f); sternite VIII broadly rounded apically, antecostal suture narrowly separated from basal margin (Fig. 18.133g);

305

spermatheca mushroom-shaped, capsule kidneyshaped, without apparent invagination, stem long, thin, with one posterior loop (Fig. 18.133h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, NF, NS, ON, QC, SK. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: adults in NB collected from Lindgren funnel traps in an old red pine forest, a mature (110-year-old) red spruce forest, an old red oak forest, a rich Appalachian hardwood forest, a trembling aspen stand with a few conifers, and a jack pine forest. Also reported from deciduous and mature red sprucehemlock and mixed forests in other areas. Some collected from forest litter, in/on Polyporus betulinus (Bull.) Fr. Collecting period: V–VI. Collecting method: Lindgren funnel traps, flight intercept traps, pitfall traps, sifting litter, and fungi.

Genus Leptusomorpha Klimaszewski and Hoebeke, gen. n. (Figs. 18.134 and 18.135) Type species: Leptusomorpha Klimaszewski, sp. n.

claudiae

Etymology. The generic name derives from the generic name Leptusa and the ending morpha (body), in allusion to the external similarity to the genus Leptusa. Gender feminine. Diagnosis. Body narrowly subparallel, moderately flattened, integument of forebody with distinct microsculpture consisting of hexagonal sculpticells, punctation and pubescence sparse (Fig. 18.135a); head as wide as pronotum, pronotum widest in basal third and distinctly narrower than base of elytra, elytra at suture about as long as pronotum along midline; head with postocular region subparallel and strongly constricted basally, as long as 1.5 times diameter of eye visible from above, eyes medium-sized and protruding; antennomere IV slightly transverse,

306

VII–X strongly transverse (Fig. 18.135a); labrum truncate apically (Fig. 18.134a); ligula slender and split apically (bifid) (Fig. 18.134f); labial palpi with moderately elongate three palpomeres (Fig. 18.134f); maxillary palpi with four palpomeres, terminal one narrow, needle-shaped (Fig. 18.134b, c); galea with soft apical part; lacinia with numerous apical teeth (Fig. 18.134b, c); right mandible with minute subapical tooth; labium truncate apically (Fig. 18.134f); pronotum trapezoidal in shape, narrowest basally and widest in apical third; elytra with angular shoulders, subparallel, posterior angles weakly sinuate; abdomen subparallel; mesocoxae narrowly separate; tarsal formula 4-45; basal 4 tarsomeres of hind tarsus of about same length. 100. Leptusomorpha claudiae Klimaszewski, sp. n. (Figs. 18.134a–f and 18.135a–d) Holotype (male): Canada, British Columbia, Mt. Cain, 50.13N, 126.21W, 23.V-9.VI.1997, 16.6 LIN7, N. Winchester (RBCM). Paratypes: BC, Monashee Mtn., near Cheriville, 1400–1600 m, 12.VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 male; BCol., Driftwood Canyon Provincial Park, 54
49.650 N, 127
01.410 W, 752 m, 27.VII.2012, alder, poplar litter, and moss along creek, A. Davies (CNC) 1 male. Etymology. This species is dedicated to Claudia Copley of the RBCM, Victoria, British Columbia, who collected many specimens examined in this study. Diagnosis. Body length 2.6–2.7 mm; head, pronotum, abdomen and antennae dark brown, elytra and legs yellow light brown (Fig. 18.135a); forebody with strong microsculpture, sculpticells hexagonal; punctation and pubescence sparse; head strongly narrowed basally, as wide as maximum width of pronotum; eyes protruding; elytra moderately transverse, shoulders angular and rounded, at suture as long as pronotum along midline, pubescence directed laterad from midline of

18

Tribe Homalotini Heer, 1839

disc; abdomen subparallel, widest in apical third. MALE. Tergite VIII truncate and slightly emarginated medially (Fig. 18.135c); sternite VIII triangularly produced apically, apex rounded (Fig. 18.135d); tubus of median lobe of aedeagus arcuate basally and strongly produced ventrally in apical half, apex thin and slightly pointed in lateral view (Fig. 18.135b); flagellum thin and inconspicuous. FEMALE. Unknown. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: montane habitats, elevation 752–1600 m; found in alder, poplar litter, and moss along creek. Collecting period: VI, VIII. Collecting method: Lindgren funnel trap, sifting litter.

Genus Stictalia Casey, 1906 (Figs. 18.136–18.139) (Casey 1885, 1893, 1906, Fenyes 1918, Seevers 1978, Ashe 1992, Klimaszewski and Winchester 2002) Diagnosis. Body moderately narrow, subparallel; integument usually moderately glossy, with or without microsculpture, punctation moderately dense, asperate or not; head slightly narrowed behind eyes, in some species forming an inconspicuous neck; eyes small, 1.2–0.5 times length of temple; maxillary palpus with four palpomeres, labial palpus with three palpomeres; ligula elongate and shallowly split apically (bifid); pronotum slightly transverse, at most 1.3 times as wide as long, slightly to distinctly narrower than base of elytra; hypomeron visible in lateral view; mesocoxal cavities narrowly separated by intercoxal process; mesoventral process narrow and acutely pointed apically; tergites III–IV or III–V with moderate to deep transverse impressions; male sternite VII with numerous basal triangular sensory pores; apical projection of male sternite VIII with asetose posterior margin; internal sac of median lobe of aedeagus

18

Tribe Homalotini Heer, 1839

without distinct sclerites; flagellum broadly tubular and rigid; spermathecal duct sclerotized and complexly looped. Comments. Stictalia notata (Mäklin), described as Bolitochara notata Mäklin, 1852, from Alaska, was not examined because the type specimen/s is/are presumably lost in the collection of Zoological Museum, Helsinki, Finland. We were informed about this fact by the collection manager who could not locate the type(s). Key to Species of Stictalia 1. Elytra as wide as pronotum (Fig. 18.139a); forebody with very coarse punctation (Fig. 18.139a)....................................................... Stictalia kranabetteri Klimaszewski and Godin – Elytra distinctly broader than pronotum (Figs. 18.136a–18.138a); forebody with fine to coarse punctation (Figs. 18.136a–18.138a) .......2 2. Pronotum with fine punctation (Fig. 18.138a), female tergite and sternite VIII and spermathecal as illustrated (Fig. 18.138b–d) ............................. ......................................Stictalia carlottae Casey – Pronotum with coarse punctation (Figs. 18.136a and 18.137a) ..............................3 3. Body robust, predominantly yellow brown (Fig. 18.137a); median lobe of aedeagus arcuate ventrally and straight apically in lateral view (Fig. 18.137b).........Stictalia californica (Casey) – Body less robust, predominantly red brown (Fig. 18.136a); median lobe of aedeagus arcuate ventrally and strongly sinuate apically in lateral view (Fig. 18.136b).............................................. ..................................Stictalia brevicornis Casey

101. Stictalia brevicornis Casey (Fig. 18.136a–g) Stictalia brevicornis Casey, 1906. Ashe 1992, Klimaszewski and Winchester 2002. Stictalia arcuata Casey, 1906. New synonymy.

307

Lectotype: British Columbia, Victoria, Vancouver Is.; arcuata Casey; Type USNM 39502; Casey bequest 1925 (USNM) 1 female. Present designation. Diagnosis. Body length 2.8–3.0 mm, moderately narrow, subparallel, convex, moderately glossy, bicoloured and variable, head, often pronotum, posterior angles of elytra and posterior part of abdomen light to dark brown, remainder of the body yellowish, antenna dark brown apically and yellow brown basally (Fig. 18.136a); forebody densely and moderately coarsely punctate, with minute punctation between setae, pubescence moderately long, microsculpture present; head elongate, narrowed behind eyes; antenna slender, antennomeres I–V strongly to slightly elongate, VII–X moderately to strongly transverse; pronotum slightly transverse, 1.2–1.3 times as wide as long; elytra elongate, moderately convex, about 1.5 times as long as pronotum along midline; abdomen subparallel, tergal impressions with coarse punctation. MALE. Tergite VIII transverse, with very small apical emargination, apical margin with or without small crenulation (Fig. 18.136c); sternite VIII transverse, triangularly produced apically, apex pointed (Fig. 18.136d); tubus of median lobe of aedeagus broadly arched ventrally and strongly sinuate subapically, apex narrowly triangular in lateral view (Fig. 18.136b). FEMALE. Tergite VIII transverse, truncate apically (Fig. 18.136e); sternite VIII rounded apically (Fig. 18.136f); spermatheca with relatively large hemispherical capsule and short U-shaped stem, posterior duct thin and coiled several times (Fig. 18.136g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA. New locality data: Vancouver, Stanley Park, Merilees Trail, CWH, 49.1840
N, 123.0902
W, Funnel Trap 1, 14.IV-22.V.2008, J.A. McLean and A. Li (LFC) 1 male, 1 sex undetermined; same data except: South Creek Trail, 49.1803
N, 123.0825
W, 22.V-27-VI.2008 (LFC) 1 sex undetermined. DNA voucher

308

18

specimen, CCDB-23526-H03, BOLD Proc.ID: LFCST087-14: BC, Upper Carmanah Valley, Vancouver Is., UTM: 10U CJ 802998, 17. X.1991, N. Winchester, FF.MT4 (LFC) 1 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: Sitka spruce forest in Carmanah Valley, Vancouver Island; most specimens were captured in transition zone and forest interior, during fall season. Majority of specimens were captured in October. Collecting period: VI–X. Collecting method: pitfall traps, funnel traps. 102. Stictalia 18.137a–h)

californica

(Casey) (Fig.

Bolitochara californica Casey 1885. Casey 1893, 1906, Moore and Legner 1975, Klimaszewski and Winchester 2002. Stictalia densicollis Casey, 1906. New synonymy. Lectotype: Br. C. [British Columbia]; densicollis Casey; Type USNM 39498; Casey bequest 1925 (USNM). Present designation. Diagnosis. Body length 3.0–3.3 mm, narrow, subparallel, moderately convex, moderately glossy, bicoloured, head, pronotum, and posterior part of abdomen light to dark brown, remainder of the body yellowish or reddish yellow (Fig. 18.137a); forebody densely and moderately coarsely punctate, with minute punctation between setae, pubescence moderately long, microsculpture present; head elongate, narrowed behind eyes to form inconspicuous neck, about as wide as pronotum; antenna slender, antennomeres I–III strongly elongate, V–X moderately to strongly transverse; pronotum slightly transverse, 1.2–1.3 times as wide as long; elytra distinctly broader than pronotum, slightly transverse, moderately convex, at suture as long as pronotum along midline; abdomen subparallel, tergal impressions with coarse punctation. MALE. Tergite VIII transverse, with small apical emargination, apical margin with or without small crenulation (Fig. 18.137d); sternite VIII

Tribe Homalotini Heer, 1839

transverse, triangularly produced apically and rounded (Fig. 18.137e); tubus of median lobe of aedeagus broadly arched ventrally and apex narrowly triangular in lateral view (Fig. 18.137b), median lobe in dorsal view as illustrated (Fig. 18.137c). FEMALE. Tergite VIII transverse, truncate apically (Fig. 18.137f); sternite VIII rounded apically (Fig. 18.137g); spermatheca with relatively large hemispherical capsule slightly pointed apically and short U-shaped stem, posterior duct thin and coiled several times (Fig. 18.137h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK [new state record], CA. We report this species from AK for the first time here based on the following: AK, Seward/ Kenai Peninsula, 15.VII.2008, Lowell Point Waldrand, K. Renner (LFC) 2 males. COLLECTION AND HABITAT DATA. Habitat: Sitka spruce forest in Carmanah Valley, Vancouver Island; most specimens were captured in transition zone and forest interior, during fall season. Majority of specimens were captured in October. Collecting period: VI–X. Collecting method: pitfall traps. 103. Stictalia carlottae Casey (Fig. 18.138a–d) Stictalia carlottae Casey, 1911. Lectotype: QCI [in original description Queen Charlotte Islands, Massett]; carlottae Casey; Type USNM 39503; Casey bequest 1925 (USNM) 1 female. Present designation. Diagnosis (based on female lectotype). Body length 3.2 mm, narrow, subparallel, moderately convex, moderately glossy, bicoloured, head, irregular areas of scutellar and lateral posterior angles of elytra, and most of abdomen dark brown, remainder of the body reddish light brown (Fig. 18.138a); elytra densely and moderately coarsely punctate but head and pronotum finely punctate, with minute punctation between setae, pubescence moderately long,

18

Tribe Homalotini Heer, 1839

microsculpture present; head elongate, narrowed behind eyes and rounded, about as wide as pronotum; antenna slender, antennomeres I–III strongly elongate, IV subquadrate, V–X moderately to strongly transverse; pronotum slightly transverse, ca. 1.2 times as wide as long; elytra elongate, distinctly wider and longer than pronotum, moderately convex; abdomen subparallel, tergal impressions with coarse punctation. MALE. Unknown. FEMALE. Tergite VIII transverse, arcuate laterally and truncate apically (Fig. 18.138b); sternite VIII rounded apically and slightly produced medially (Fig. 18.138c); spermatheca with relatively large hemispherical capsule and short U-shaped stem, posterior duct thin and coiled several times (Fig. 18.138d). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. New locality data: British Columbia, Vancouver Island, 3 km SW Brentwood Bay, 48
330 1200 N, 123
290 3600 W, 6.X.2011, moss, fern and Amelanchier litter, edge of upland pond in forest, A. Davies (CNC) 1 female. COLLECTION AND HABITAT DATA. Habitat: moss, fern and Amelanchier litter at the edge of upland pond in forest. Collecting period: X. Collecting method: sifting litter. Comments. We tentatively consider Stictalia carlottae as a valid species but additional specimens and particularly males from the type locality are needed to validate the status of this species. 104. Stictalia kranabetteri Klimaszewski and Godin, sp. n. (Fig. 18.139a–g) Holotype (male): Canada, British Columbia, 65 km N of Prince George, 54.2N, 122.37W, 780 m, VIII–IX.2012, pitfall, OM1C2, SP#137, coll. M. Kranabetter (RBCM). Paratypes: Canada, British Columbia, 65 km N of Prince George, 54.2N, 122.37W, 780 m, VIII-IX.2012, pitfall, OM1C2, SP#137, coll. M. Kranabetter

309

(RBCM) 2 females; BC, Mt. Cain, 50.13N, 126.21W, 7.VII-20.VII.1997, 16.6 PIT 2, N. Winchester; Barcode of life, DNA voucher specimen, CCDB 23526–H04, LFCST088-14 (LFC) 1 male [formerly identified as S. californica]. Etymology. This species is named for Dr. John Marty Kranabetter of the Ministry of Forests of British Columbia, Vancouver Island, who collected the original series of specimens. Diagnosis. Body length 3.0–3.2 mm, narrow, subparallel, moderately convex, moderately glossy, bicoloured, head, pronotum, and posterior part of abdomen light to dark brown, remainder of the body red light brown (Fig. 18.139a); forebody densely and very coarsely punctate, with minute punctation between setae, pubescence moderately long, microsculpture present; head elongate, narrowed behind eyes to form inconspicuous neck, about as wide as pronotum; antenna slender, antennomeres I–III strongly elongate, IV slightly elongate, V–X moderately to strongly transverse; pronotum about as wide as elytra, slightly transverse, 1.2 times as wide as long; elytra slightly transverse, moderately convex, at suture slightly longer than pronotum along midline; abdomen subparallel, tergal impressions with coarse punctation. MALE. Tergite VIII transverse, with small apical emargination, apical margin with small crenulation (Fig. 18.139c); sternite VIII transverse, triangularly produced apically, apex slightly pointed (Fig. 18.139d); tubus of median lobe of aedeagus broadly arched ventrally and apex narrowly triangular in lateral view (Fig. 18.139b). FEMALE. Tergite VIII transverse, truncate apically (Fig. 18.139e); sternite VIII rounded apically and slightly produced medially (Fig. 18.139f); spermatheca with relatively large hemispherical capsule slightly pointed apically and short U-shaped stem, posterior duct thin and coiled several times (Fig. 18.139g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

310

18

COLLECTION AND HABITAT DATA. Habitat: coniferous forests. Collecting period: VIII, IX. Collecting method: pitfall traps. Comments. This species may be readily distinguished from the remaining Canadian species by narrow elytra, which are about as wide as pronotum and by very coarse body punctation. Subtribe Gyrophaenina Kraatz, 1856 (Ashe 1984, 2001; Klimaszewski et al. 2018) Diagnosis. Mesocoxae separated by width of mesocoxal cavity or slightly less by broad and broadly contiguous processes of meso- and metaventrite; laciniae distinctive, apex obliquely truncate and covered with numerous spinules, with setae and spines of inner surface reduced or absent (for illustrations see Ashe 2001). Comments. Associated with fresh fruiting bodies of basidiomycete fungi.

Genus Encephalus Stephens, 1832 (Fig. 18.140a–e) (Seevers 1951, 1978; Ashe 2001) Diagnosis. Body robust, compact, broadly oval in dorsal outline, broadest at abdominal level, body with antero-posterior outline convex in lateral view; head hypognathous, deflexed and incompletely visible from above, infraorbital carina present; eyes moderate in size, finely faceted; ligula entire; antennae short, antennomeres IV–X strongly incrassate; pronotum strongly convex, anterolateral margins of pronotum strongly deflexed, pronotal hypomera not visible in lateral view; mesoventrite nearly in a vertical plane; mesoventral process wide and long, extending to posterior limits of middle coxae; metaventral process absent; abdomen with tergites broad and short. 105. Encephalus (Fig. 18.140a–e)

americanus

Seevers

Tribe Homalotini Heer, 1839

Encephalus americanus Seevers, 1951. Fenyes 1918, Seevers 1978. Paratype: Creston, BC, 18.V.1951, G. Stace Smith, ex fungus (FMNH) 1 male. Examined. Diagnosis. Body length 1.9–2.0 mm, robust, compact, subovate in dorsal outline; head, pronotum and abdomen black, elytra rufous in central part of disc and blackish laterally and basally, appendages yellowish brown (Fig. 18.140a); head with about eight moderate punctures medial to each antenna; pronotum twice as broad as wide, sparsely punctate, with one coarse pair of punctures, subbasal impression of disc with confused punctation; elytra at suture about as long as pronotum, coarsely and sparsely punctate and pubescent except near suture; abdomen short, strongly narrowed posteriad. MALE. Tergite VIII with four slender teeth, the median two slightly shorter (Fig. 18.140d); sternite VIII parabolically rounded apically (Fig. 18.140e); median lobe of aedeagus with oval bulbus and triangular tubus in dorsal view (Fig. 18.140c); in lateral view tubus of median lobe strongly sinuate subapically and with apex sharply pointed ventrad (Fig. 18.140b). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC. USA: IL, MT, NH. Material examined: USA, Ill., Antioch, coll. F. Psota (FMNH) 1 male. COLLECTION AND HABITAT DATA. Habitat: found in fungus. Collecting period: VIII, IX. Collecting method: not recorded.

Genus Gyrophaena Mannerheim, 1830 (Figs. 18.141–18.145) Klimaszewski et al. 2018)

(Seevers

1951;

Diagnosis. Body broadly oval and dorsoventrally flattened, length 1.2–3.5 mm, moderately or highly glossy; color flavate, brown,

18

Tribe Homalotini Heer, 1839

piceous, or black, usually bicoloured but sometimes uniformly coloured; integument of forebody with large insertion pores, often elevated (asperities), forming distinct patterns on pronotum and head; isodiametric meshed microsculpture usually present; head with genae well developed; infraorbital carina present; eyes large and often prominent, finely faceted and broadly separated; labrum broadly oval or trapezoidal with sensory setae forming patterns; mandibles robust, the right mandible bearing one, usually well-developed internal tooth, and a welldeveloped internal membranous lobe (the prostheca), which bears rows of small teeth; maxillae with laciniae obliquely truncate and with numerous closely spaced teeth (spore brush), and rows of setae on the outer lobe, galeae provide a cup-like cap over the apex of the lacinial comb; ligula short, entire, produced and broadly rounded at apex; antennomere IV small, V slightly broader than VI, and IV–X usually incrassate apically; pronotum usually transverse with hypomera visible from the side; mesocoxae broadly separated by about width of mesocoxal cavity by broad and broadly contiguous processes of meso- and metaventrite; elytra usually broadshouldered; abdomen tapering apically; process of mesoventrite truncate apically, at least as long as that of metaventrite; spermatheca sclerotized, with lateral fin-like flange on the short stem, capsule spherical; median lobe of aedeagus variably shaped, often tubular or trough-shaped, often with ventral projections of various shapes and sizes forming a complex structure; apical portion of internal sac rigid and forming elongate projection from where flagellum is normally exerted, the internal sac lacks internal and external spines and sclerites (unlike most other aleocharine genera); paramere with broad apical lobe bearing four setae, usually two longer and two shorter ones; apical margin of male tergite VIII usually with two relatively large sublateral teeth bordering emargination with smaller teeth, rarely larger sublateral teeth reduced and smaller teeth lacking,

311

or sublateral teeth replaced with large lobes; species strictly associated with mushrooms. The species groups in Gyrophaena are mainly defined by the similarities of the median lobe of the aedeagus. Key to Species of Gyrophaena

Groups

and

Species

1. Elytral punctures broad and shallow (Fig. 18.141a); median lobe of aedeagus with tubus split into two arms, flagellum coiled externally (Fig. 18.141b); apical margin of male tergite VIII with two long and narrow sublateral teeth, with or without two minute teeth in-between (Fig. 18.141c)....................................................... ....................... Gyrophaena affinis Mannerheim – Elytral punctures and genital characters different ..............................................................2 2. Elytral punctures coarse, deep and dense (Fig. 18.144a); median lobe of aedeagus with tubus triangularly elongate, sharply pointed apically, with or without short vestigial narrow projection in lateral view (Fig. 18.144b)................... ............................... Gyrophaena nana (Paykull) – Elytral punctures and tubus of median lobe of aedeagus not as above........................................ 3 3. Head distinctly narrower than pronotum (Fig. 18.145a); male tergite VIII with two long and thin apical teeth and two minute ones between (Fig. 18.145c); tubus of median lobe of aedeagus straight medially and split at apex in lateral view (Fig. 18.145b); female sternite VIII rounded apically and slightly pointed medially (Fig. 18.145f) ....................................................... ........................... Gyrophaena californica Casey – Head at least as broad as pronotum (Figs. 18.142a and 18.143a); female sternite VIII triangularly produced apically and strongly pointed medially, other genital characters not as above ..................................................................4

312

4. Pronotum light to dark brown, male tergite VIII with two long, narrow lateral teeth and two shorter, finer medial teeth (Fig. 18.143c); tubus of median lobe of aedeagus short, sinuate ventrally in lateral view (Fig. 18.143b)............................... .................................... Gyrophaena keeni Casey – Pronotum dark rufo-testaceous to piceous (Fig. 18.142a), male tergite VIII with two long, narrow teeth and two, rarely three, minute median teeth which are sometime obsolete (Fig. 18.142c); tubus of median lobe of aedeagus long, strongly angular medially and sinuate subapically (Fig. 18.142b)........... Gyrophaena uteana Casey

Affinis Species Group (Seevers 1951) Diagnosis. Elytral punctures broad and shallow; median lobe of aedeagus with tubus subdivided into two arms, apical part of internal sac coiled and twisted, flagellum narrowly elongate. 106. Gyrophaena affinis Mannerheim { (Fig. 18.141a–g) Gyrophaena affinis Mannerheim, 1830. Casey 1906 [as G. lacustris and G. subpunctata], Seevers 1951, Klimaszewski et al. 2009, 2011, Brunke et al. 2012, Enushchenko and Semenov 2016, Klimaszewski et al. 2016, 2018. Diagnosis. Body narrowly oval, length 1.7–2.1 mm; head rufo-piceous to piceous, pronotum flavate to light brown, elytra testaceous to light brown (Fig. 18.141a); abdomen rufoflavate, subapical part of abdomen usually darker; vertex of head with about 10 large umbilicate punctures on each side, pronotum with a few scattered large punctures, elytra with numerous broad shallow punctures; forebody with microsculpture reticulate throughout; antennomeres VI–X subquadrate to slightly transverse; pronotum 1.4 times as wide as long. MALE. Apical margin of tergite VIII truncate, with two long, arcuate sublateral teeth, margin smooth between them (Fig. 18.141c); sternite

18

Tribe Homalotini Heer, 1839

VIII broadly rounded apically (Fig. 18.141d); median lobe of aedeagus in lateral view with tubus bent perpendicular to bulbus basally, narrowly triangular, split subapically, ventral margin sinuate, angulate basally, apex narrow and rounded, dorsal projection of internal sac moderately elongate, coiled with flagellum inside (Fig. 18.141b). FEMALE. Tergite VIII distinctly transverse, truncate apically, antecostal suture obtusely angular medially (Fig. 18.141e); sternite VIII with apical margin obtusely angulate, apex pointed (Fig. 18.141f); spermatheca as illustrated (Fig. 18.141g). DISTRIBUTION. Origin: Holarctic or Palearctic, possibly adventive in Canada and USA, but its distribution status in not clear. CANADA: BC, MB, NB, NF, NS, ON, QC, SK. USA: AZ, DC, IL, IN, IA, KY, MA, ME, MI, MN, MO, NC, NH, NJ, NM, NY, OH, PA, TN, WA, WI, WV. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mixed forest, 8.5-year-old regenerating mixed forest, fir forests, eastern white cedar swamp, mature red spruce forests, red maple forest, and a red oak forest. Adults on/in gilled mushrooms on forest floor, on log, and on a stump, on small gilled mushrooms on side of decayed log. This species has also been found in rotting mushrooms, in an orange bracket (polypore) fungus, on bracket fungus on white birch, and on Pleurotus sp. on dead standing Populus tremuloides Michx. Some reared from fungus in ON. Collecting period: VI–IX. Collecting method: sifting mushrooms, pitfall traps. Bihamata Species Group (Seevers 1951) Diagnosis. Antennomeres V–X slightly transverse; pronotum moderately transverse; apical margin of male tergite VIII with two long, slender, widely separated teeth (very small in G. flavicornis); flagellum protruding between three apical processes of tubus.

18

Tribe Homalotini Heer, 1839

107. Gyrophaena uteana Casey (Fig. 18.142a–g) Gyrophaena uteana Casey, 1906. Casey 1906 [as G. pacifica], Seevers 1951, Klimaszewski et al. 2009 [as G. gaudens], Webster et al. 2012, Brunke et al. 2012, Klimaszewski et al. 2015, 2018. Diagnosis. Body narrowly oval, length 1.4–1.7 mm (Fig. 18.142a); head about as broad as pronotum, piceous, pronotum dark rufotestaceous to piceous; elytra reddish-brown with sutural region and posterior angles piceous, abdomen yellowish- or reddish-brown, apical portion often darker, piceous; vertex of head with at least six moderate-sized umbilicate punctures on each side, pronotum with six punctures in two confused median rows and additional scattered punctures, elytra with punctures fine and sparse; forebody with microsculpture reticulate throughout, weaker on pronotum, most evident on head; antennomeres VI–X moderately transverse; pronotum 1.5 times as wide as long. MALE. Apical margin of tergite VIII with two long, narrow teeth (narrower and more sharply pointed than those of G. michigana) bordering broad, shallow emargination with two, rarely three, minute median teeth (Fig. 18.142c); sternite VIII very broadly parabolically rounded apically (Fig. 18.142d); tubus of median lobe of aedeagus elongate, narrow, ventral margin bent angularly ventrad near base and again before middle, without angular swelling at middle, dorsal margin with minute subapical tooth-like projection (Fig. 18.142b), apical projection of internal sac irregular in shape (Fig. 18.142b). FEMALE. Tergite VIII truncate apically (Fig. 18.142e); sternite VIII with apical margin obtusely triangularly produced, rounded at apex (Fig. 18.142f); spermatheca as illustrated (Fig. 18.142g). Gyrophaena uteana is externally similar to G. michigana but differs in the male genital features. Gyrophaena uteana is similar externally and has genitalia similar to those of the Palearctic G. bihamata Thomson, but according to Seevers (1951), G. bihamata is a distinct species, differing

313

from G. uteana in having the reticulation less pronounced, the antennae less incrassate, and the median lobe of the aedeagus with a slightly different shape. For illustrations of G. bihamata, see Enushchenko and Semenov (2016). These two species need to be re-examined because they appear to be difficult to separate. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, ON, QC, SK. USA: CA, CO, UT. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in a mixed forest, a silver maple swamp, and a maple-beech forest. Adults on/in polypore fungi, on bracket and gilled fungi, on a small (fresh) gilled mushroom on a log, and from a Pleurotus sp. on a dead, standing trembling aspen (Populus tremuloides Michx.). Collecting period: VI–IX. Collecting method: sifting mushrooms, Berlese extraction of leaf and log litter, malaise pans. Keeni Species Group (Seevers 1951) Diagnosis. Antennomeres V–X moderately to distinctly transverse; pronotum sparsely pubescent and highly glossy; male tergite VIII with two large lateral teeth and two small approximate median teeth; tubus of median lobe of aedeagus narrowly triangular and sinuate ventrally in lateral view, flagellum everted, coiled and twisted. 108. Gyrophaena keeni Casey (Fig. 18.143a–g) Gyrophaena keeni Casey, 1911. Seevers 1951, Klimaszewski and Winchester 2002, Klimaszewski et al. 2005, 2009, 2011, 2018. Diagnosis. Body roughly narrowly ovoid, length 1.6–1.9 mm; head about as broad as pronotum, rufo-piceous to piceous; pronotum reddish-brown or medium dark brown, elytra light reddishbrown or light brown with posterior angles dark brown, abdomen reddish-brown or light brown with posterior portion piceous (Fig. 18.143a). vertex of head with at least six small punctures on each side, disc of pronotum usually with

314

median rows of three punctures and a small cluster medially near base, elytra sparsely and irregularly punctate; microsculpture distinct, finely meshed on head and pronotum; antennomeres VI–X moderately transverse; pronotum 1.4 times as wide as long. MALE. Tergite VIII with apical margin truncate, with two long, narrow sublateral teeth and two shorter, finer medial teeth (Fig. 18.143c); sternite VIII with margins almost evenly arcuate from base to apex (Fig. 18.143d); median lobe of aedeagus in lateral view with tubus moderately long and broad, sinuately tapering to acute apex, curved obliquely ventrad basally, apical projection of internal sac tubular and short, flagellum everted and curled, compressor plate elevated (Fig. 18.143b). FEMALE. Tergite VIII truncate apically (Fig. 18.143e); sternite VIII with apical margin obtusely angulate (Fig. 18.143f); spermatheca as illustrated (Fig. 18.143g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, YT. USA: FL, MA, ME, MT, NH, NY, OR, TN, WA, WY. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mature mixed forest, 8.5-year-old regenerating mixed forest, red oak forest, eastern white cedar (Thuja occidentalis L.) swamps, red spruce (Picea rubens Sarg.) and red maple (Acer rubrum L.) forest (80–120 years old), black spruce forests, recently burned coniferous forest, and an eastern hemlock forest [Tsuga canadensis (L.)] 120+ years old. Adults in in gilled mushrooms, in gilled fungi on rotten log, in gilled mushrooms on stump, in moss near brook, on polypore fungi on dead standing Populus sp. Collecting period: VI–IX. Collecting method: sifting mushrooms, sifting moss, flight intercept trap, and pitfall trap. Comments (Klimaszewski et al. 2018). Strand (1938) described Gyrophaena orientalis from the Palearctic region, and it is known to occur in central and Northern Europe, Caucasus and Siberia (Enushchenko and Semenov 2016). This species is extremely similar to Gyrophaena keeni

18

Tribe Homalotini Heer, 1839

and very likely a synonym of the Nearctic species. We have compared images of the body, median lobe of the aedeagus and male tergite VIII (Enushchenko and Semenov 2016) of G. orientalis with those of G. keeni and found no significant differences between the two species. For formal synonymization of G. orientalis, the types of the two species need to be examined and compared. Nana Species Group (Seevers 1951) Diagnosis. Antennomeres V–X moderately to distinctly transverse; head with punctures strongly impressed; apical margin of male tergite VIII with two long, apically rounded sublateral teeth and two shorter, finer, apically rounded teeth medially; tubus of median lobe of aedeagus simple in lateral view, roughly triangular apically, dorsal margin usually with minute, subapical denticle; internal sac and flagellum everted and straight or sinuate. 109. Gyrophaena nana (Paykull)  (Fig. 18.144a–g) Staphylinus nana Paykull, 1800. Campbell and Davies 1991, Klimaszewski et al. 2009, 2011, 2012, Webster et al. 2012, Enushchenko and Semenov 2016. Diagnosis. Body widely oval, length 1.7–2.0 mm,; head piceous, pronotum reddishbrown with central part of disc darker, elytra piceous, paler anteriorly and along lateral margins, abdomen dark brown or lighter with some tergites dark brown (Fig. 18.144a); antennomeres VI–X transverse and incrassate apically; head slightly narrower than pronotum, pronotum distinctly transverse, widest at middle, distinctly narrower than elytra; elytra at suture longer than pronotum; abdomen wide, basally almost as wide as elytra, punctures coarse, deep and dense. MALE. Apical margin of tergite VIII with two sublateral, moderately long teeth and two finer, slightly shorter, approximate teeth medially (Fig. 18.144c); sternite VIII

18

Tribe Homalotini Heer, 1839

315

semicircularly rounded from base (Fig. 18.144d); median lobe of aedeagus in lateral view with tubus simple, curved very obliquely ventrad before middle of ventral margin, gradually tapering to acute apex, apical projection of internal sac curved dorsad (Fig. 18.144b). FEMALE. Tergite VIII virtually truncate apically (Fig. 18.144e); sternite VIII similar to that of male but slightly obtuse apically (Fig. 18.144f); spermatheca as illustrated (Fig. 18.144h). DISTRIBUTION. Origin: Holarctic, Europe, Caucasus, Far East, and North America. CANADA: AB, BC, MB, NB, NF, ON, YT. USA: AK, CA, MA, ME, MI, MT, WI, WY. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: one specimen from NB from mushroom on log in mixed forest; specimens from NL from rotting mushrooms in unspecified forest type. Little else is known about habitat associations. Collecting period: IV–IX. Collecting method: sifting mushrooms.

brown (Fig. 18.145a); integument strongly reticulate; head distinctly broader than pronotum, punctation of head shallow and weak; pronotum 1.4 times as wide as long, sparsely punctate and pubescent, median rows of punctures represented by one subbasal pair; antennae short, antennomeres V–X strongly transverse, incrassate; elytra with punctures sparse. MALE. Tergite VIII with two long and narrow lateral teeth and two minute median teeth (Fig. 18.145c); sternite VIII rounded apically (Fig. 18.145d); tubus of median lobe of aedeagus bent basally, straight medially and produced ventrad at apex, apex split in lateral view (Fig. 18.145b). FEMALE. Tergite VIII truncate apically (Fig. 18.145e); sternite VIII parabolic apically (Fig. 18.145f); spermatheca of a generalized shape, as illustrated (Fig. 18.145g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, CO, NM. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded.

Strictula Species Group (Seevers 1951) Diagnosis. Body colour more or less uniformly brown, integument with strong reticulation, punctuation of head and pronotum weak, elytra often densely asperate, antennae short and compact, antennomeres V–X strongly transverse, male tergite VIII with two larger lateral and two very small median teeth, and median lobe of aedeagus distinctive, often with split apex. 110. Gyrophaena 18.145a–g)

californica

Subtribe Silusina Fenyes, 1918 (Ashe 2001; Klimaszewski et al. 2003, 2018) Diagnosis. Labial palpi extremely elongate, filiform or stylate, appearing as with two palpomeres but borders between them often diffused; ligula elongate, entire apically; mesocoxae narrowly separated by narrow processes of meso- and metaventrite (Fig. 217, in Ashe 2001).

Casey (Fig.

Genus Silusa Erichson, 1837 1906.

(Figs. 18.146–18.148) (Klimaszewski et al. 2003, 2018)

Diagnosis. Body length 1.5–1.8 mm, head and posterior part of abdomen except for apex dark brown to almost black, the remainder of body reddish brown but pronotum and elytra with irregular blackish spots, appendages yellowish

Diagnosis. Body elongate and subparallel, robust, length 2.5–4.5 mm; integument usually coarsely punctate, pubescence fine and sparse to dense; head narrowly protruding in front of eyes, infraorbital carina strong and complete; antennomeres VII–X 1.3–2.0 times as wide as

Gyrophaena Seevers 1951.

californica

Casey,

316

long; ligula narrowly elongate and entire; labial palpi extremely elongate, filiform or stylate, forming an acute V structure; maxillary palpi with four palpomeres, terminal one narrow and pointed; mentum with anterolateral margins not produced as spiniform processes; pronotum transverse and broadest about middle or in apical one-third, pubescence on disc directed anteriad along midline apically and posteriad elsewhere, on each side directed obliquely lateroposteriad; pronotal hypomera visible in lateral view; elytra more-or-less elongate with posterolateral angles distinctly, sinuately emarginate, pubescence on disc directed slightly obliquely posteriad; first four visible abdominal tergites deeply impressed basally; hind tarsus with basal article slightly longer than following one; apical margin of male tergite VIII denticulate; median lobe of aedeagus strongly curved or bent ventrad at some point, internal sac with distinct subapical sclerites and with flagellum strongly sclerotized, with basal part swollen; spermatheca with capsule large, usually globose, lacking apical invagination, stem thin and short. Key to Species of Silusa 1. Pronotum broadest in apical third, disc flattened medially (Fig. 18.148a); body colour predominantly dark brown (Fig. 18.148a); male tergite VIII with small apical teeth of about same size (Fig. 18.148e); median lobe of aedeagus and spermathecal as illustrated (Fig. 18.148b–d)............... Silusa vesperis Casey – Pronotum broadest in basal third, disc convex medially (Figs. 18.146a, 18.147a); body colour predominantly yellowish or reddish brown (Figs. 18.146a and 18.147a); male tergite VIII with large teeth of different size (Figs. 18.146d and 18.147e); median lobe of aedeagus not as above (Figs. 18.146b, c and 18.147b–d) ...........2 2. Body surface highly glossy (Fig. 18.146a); colour dark reddish brown with margins of pronotum, elytra and abdominal tergites paler (Fig. 18.146a); in lateral view median lobe of aedeagus with tubus abruptly bent almost perpendicularly ventrad at middle of ventral margin

18

Tribe Homalotini Heer, 1839

(Fig. 18.146b); spermatheca with capsule spherical, stem long, thin, sinuate, with accessory sclerites at base of seminal channel (Fig. 18.146h) ...........................Silusa californica (Bernhauer) – Body surface moderately glossy (Fig. 18.147a); colour dark or light brown with margins of pronotum, elytra and abdominal tergites paler (Fig. 18.147a); in lateral view median lobe of aedeagus with tubus evenly arcuate, apex not so strongly produced ventrad (Fig. 18.147b, c); spermatheca with capsule spherical, stem moderately long, thin, sinuate, without accessory sclerites at base of seminal channel (Fig. 18.147i) ........ Silusa opaca Fenyes

Californica Species Group (Klimaszewski et al. 2003, 2018) Diagnosis. Median lobe of aedeagus in lateral view with tubus moderately narrow and elongate, about as long as bulbus, curved or abruptly bent ventrad at about middle of ventral margin; flagellum broad and moderately long, about as long as tubus, with basal part highly enlarged, funnelshaped. 111. Silusa californica (Fig. 18.146a–h)

(Bernhauer)

Stenusa (Silusa) californica Bernhauer, Klimaszewski and Winchester Klimaszewski et al. 2003, 2005, 2008, 2016, 2018, Majka and Klimaszewski 2010, Brunke et al. 2012.

1905. 2002, 2011, 2008,

Diagnosis. Body length 3.0–4.0 mm; colour dark reddish brown with margins of pronotum, elytra, abdominal tergites and basal antennomeres paler (Fig. 18.146a); body surface highly glossy, moderately densely punctate and pubescent; antennomere IV elongate, VIII–X distinctly transverse; pronotum distinctly transverse, pronotal punctures dense and coarse, usually separated by diameter of a puncture at least basally, the intervals with fine meshed microsculpture; elytra moderately transverse; first four visible abdominal tergites with deep transverse impressions

18

Tribe Homalotini Heer, 1839

bearing moderately coarse punctation. MALE. Tergite VIII broadly shallowly emarginate apically bearing distinct, rounded, slightly uneven teeth (Fig. 18.146d); sternite VIII with apical margin obtusely triangularly produced with apex rounded (Fig. 18.146e); median lobe of aedeagus in lateral view with tubus narrowly elongate, bent obtusely ventrad at base, abruptly bent again at middle of ventral margin so that apex directed slightly basad (Fig. 18.146b), in dorsal view median lobe as illustrated (Fig. 18.146c). FEMALE. Tergite VIII transverse, with apical margin broadly truncate, arcuately produced laterally (Fig. 18.146f); sternite VIII with apical margin obliquely produced laterally, apex slightly emarginate (Fig. 18.146g); spermatheca with capsule large, globose, stem sinuate, thin, duct with auxiliary posterior sclerites (Fig. 18.146h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, PE, QC, SK, YT. USA: AK, CA, MN. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: in many hardwood and conifer forest types; unspecified hardwood forest types, rich Appalachian hardwood forest, red oak forest, old mixed forest with red oak, mixed forests, northern hardwood forest, coastal red spruce and birch forest, mature red spruce forest with eastern white cedar, old red spruce forest, spruce forest, white spruce forest, yellow birch/balsam fir forest, trembling aspen forest with a small amount of eastern balsam poplar, white birch forest, white spruce, and willow forests. In NB, adults common in decaying mushrooms in late summer; in fleshy fungi, in decaying gilled mushrooms, baited with pile of decaying mushrooms, fleshy fungi in various stages of decay, in gilled mushrooms, in rotten fungi. Elsewhere, in wet moss on forest floor, marten dung on moss, and in mushrooms. Collecting period: VII–X. Collecting method: sifting mushrooms, baiting with mushrooms, pitfall traps, Luminoc pit-light traps, malaise traps, flight intercept traps, carrion baited traps, and sifting forest litter.

317

112. Silusa opaca Fenyes (Fig. 18.147a–i) Silusa opaca Fenyes, 1909. Moore and Legner 1975, Klimaszewski and Winchester 2002. Diagnosis. Body length 2.8–3.2 mm; colour dark or light brown with margins of pronotum, elytra and abdominal tergites paler, in some body mottled with reddish tinge (Fig. 18.147a); integument moderately glossy, densely punctate and pubescent; antennomeres VII–X strongly transverse; pronotum distinctly transverse, broadest in basal third, pronotal punctures dense and fine, usually separated by diameter of a puncture at least basally, the intervals with fine meshed microsculpture; elytra slightly transverse; first four visible abdominal tergites with deep transverse impressions bearing moderately coarse punctation. MALE. Tergite VIII truncate apically with two larger lateral teeth and four smaller ones medially (Fig. 18.147e); sternite VIII produced apically, apex rounded (Fig. 18.147f); median lobe of aedeagus with narrowly oval bulbus and broad, subparallel tubus, apex triangular in dorsal view (Fig. 18.147d); in lateral view tubus arcuate and apex produced ventrad (Fig. 18.147b, c); internal sac structures as illustrated (Fig. 18.147b, c). FEMALE. Tergite VIII arcuate apically and slightly emarginated medially (Fig. 18.147g); sternite VIII rounded apical and slightly produced medially (Fig. 18.147h); spermatheca with spherical capsule lacking apical invagination, stem thin and sinuate (Fig. 18.147i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002). Habitat: Sitka spruce forest in Carmanah Valley, Vancouver Island, adults captured in forest interior, transition zone and clearcut. Collecting period: VI–VIII. Collecting method: Malaise traps. Vesperis Species Group (Klimaszewski et al. 2003, 2018)

318

Diagnosis. Median lobe of aedeagus in lateral view with tubus moderately narrow and elongate, about as long as bulbus, ventral margin obliquely to perpendicularly curved ventrad near middle, apex acutely triangular; flagellum broad and moderately long, about as long as tubus, with basal part highly enlarged, funnel-shaped. 113. Silusa vesperis Casey (Fig. 18.148a–i) Silusa vesperis Casey, 1893. Moore and Legner 1975, Klimaszewski et al. 2003. Diagnosis. Body length 3.4–4.0 mm; predominantly dark brown to almost black, or dark brown with paler, rust-brown central part of elytra, tibiae, tarsi and antennae (Fig. 18.148a); integument of head and pronotum dull and sparsely punctate and pubescent; antennomeres VII–X strongly transverse; pronotum distinctly transverse, flattened, broadest in apical third, pronotal punctures sparse and fine, usually separated by diameter of a puncture at least basally, the intervals with fine meshed microsculpture; elytra slightly transverse, at suture as long as pronotum

18

Tribe Homalotini Heer, 1839

along midline; first four visible abdominal tergites with deep transverse impressions, bearing moderately coarse punctation. MALE. Tergite VIII emarginate medially with small and even dents (Fig. 18.148e); sternite VIII produced medially, apex broadly rounded (Fig. 18.148f); median lobe of aedeagus broad and with four elongate sclerites of internal sac in dorsal view (Fig. 18.148d); in lateral view tubus of median lobe arcuate and strongly bent ventrad apically in lateral view (Fig. 18.148b, c). FEMALE. Tergite VIII arcuate apically (Fig. 18.148g); sternite VIII rounded apically (Fig. 18.148h); spermatheca with capsule spherical, stem thin and sinuate with small posterior structure (Fig. 18.148i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, OR, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2003). Habitat: mountain habitats, at elevations 304–944 m. Collecting period: V–VIII. Collecting method: not recorded.

18

Tribe Homalotini Heer, 1839

Fig. 18.133 (a–h) Images of Leptusa gatineauensis Klimaszewski and Pelletier: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) male tergite VIII; (e) male

319

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) after Klimaszewski et al. (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

320

18

Tribe Homalotini Heer, 1839

Fig. 18.134 (a–f) Mouthparts of Leptusomorpha claudiae Klimaszewski, sp. n.: (a) labrum; (b, c) maxillae; (d, e) mandibles; (f) mentum, labial palps and ligula. Scale bar ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.135 (a–d) Images of Leptusomorpha claudiae Klimaszewski, sp. n.: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite

321

VIII; (d) male sternite VIII. (a–d) holotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

322

Fig. 18.136 (a–g) Images of Stictalia brevicornis Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

18

Tribe Homalotini Heer, 1839

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.137 (a–h) Images of Stictalia californica (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

323

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

324

Fig. 18.138 (a–d) Images of Stictalia carlottae Casey (lectotype): (a) habitus in dorsal view (apical part of abdomen removed); (b) female tergite VIII; (c) female sternite

18

Tribe Homalotini Heer, 1839

VIII; (d) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.139 (a–g) Images of Stictalia kranabetteri Klimaszewski and Godin, sp.n.: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

325

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. (a–d) holotype, (e–g) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

326

Fig. 18.140 (a–e). Images of Encephalus americanus Seevers (lectotype): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of

18

Tribe Homalotini Heer, 1839

aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.141 (a–g) Images of Gyrophaena affinis Mannerheim: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

327

female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

328

Fig. 18.142 (a–g) Images of Gyrophaena uteana Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

18

Tribe Homalotini Heer, 1839

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.143 (a–g) Images of Gyrophaena keeni Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

329

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

330

Fig. 18.144 (a–g) Images of Gyrophaena nana (Paykull): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

18

Tribe Homalotini Heer, 1839

female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.145 (a–g) Images of Gyrophaena californica Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

331

male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

332

Fig. 18.146 (a–h) Images of Silusa californica (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

18

Tribe Homalotini Heer, 1839

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca with associated structures. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

Fig. 18.147 (a–i) Images of Silusa opaca Fenyes: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f)

333

male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c, i), after Klimaszewski et al. (2003). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

334

Fig. 18.148 (a–i) Images of Silusa vesperis Casey: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male sternite VIII; (g) female

18

Tribe Homalotini Heer, 1839

tergite VIII; (h) female sternite VIII; (i) spermatheca. (b, d, i) after Klimaszewski et al. (2003). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

18

Tribe Homalotini Heer, 1839

References Ashe JS (1984) Generic revision of the subtribe Gyrophaenina (Coleoptera: Staphylinidae: Aleocharinae) with a review of the described subgenera and major features of evolution. Quaestiones Entomol 20:129–349 Ashe JS (1992) Phylogeny and revision of genera of the subtribe Bolitocharina (Coleoptera: Staphylinidae: Aleocharinae). Univ Kansas Sci Bull 54(10):335–406 Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Bernhauer M (1905) Neue Aleocharinen aus Nordamerika. Dtsch Entomol Z 1905(2):249–256 Bishop DJ, Majka CG, Bondrup-Nielsen S, Peck SB (2009) Deadwood and saproxylic beetle diversity in naturally disturbed and managed spruce forests in Nova Scotia. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:309–340 Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:119–206 Campbell JM, Davies A (1991) Family Staphylinidae rove beetles. pp 86–124. In: Bousquet Y (ed) Checklist of beetles of Canada and Alaska. Research Branch Agriculture Canada Publication 1861/E, vi + 430 pp Casey TL (1885) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1:285–336 Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7 [1892–1894]:281–606 [see reference in Chapter 7 for explanation of date] Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St Louis 16(6):125–434 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp. 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing, Lancaster, PA, 259 pp Enushchenko IV, Semenov VB (2016) A review of the genus Gyrophaena Mannerheim 1830 (Coleoptera: Staphylinidae: Aleocharinae: Gyrophaenina) of the Caucasus and adjacent territories. Zootaxa 4126 (3):301–337 Erichson WF (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Fenyes A (1909) New Aleocharinae (Staphylinidae, Col.) of the US. Entomol News Proc Entomol Sect Acad Nat Sci Philadelphia 20:418–425

335 Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera insectorum, vol XXVII, Fascicule 173A. M. Nijhoff, Den Haag, 453 pp + 7 pls Heer O (1839) (1838–1841) Fasciculus secundus [pp 145–360]. In: Fauna Coleopterorum Helvetica. Pars I. Turici [Zurich], Orellii, Fuesslini et Sociorum: xii + 652 pp Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belge Entomol 40:3–126 Klimaszewski J, Pohl G, Pelletier G (2003) Revision of the Nearctic Silusa (Coleoptera, Staphylinidae, Aleocharinae). Can Entomol 135(2):159–186 Klimaszewski J, Pelletier G, Majka C (2004) A revision of Canadian Leptusa Kraatz (Col., Staphylinidae, Aleocharinae): new species, new distribution records, key and taxonomic considerations. Belg J Entomol 6:3–42 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48 Klimaszewski J, Godin B, Pelletier G, Savard K (2008) Six new species and records of aleocharine beetles from the Yukon and Alaska (Coleoptera: Staphylinidae: Aleocharinae). Can Entomol 140 (3):265–291 Klimaszewski J, Webster RP, Savard K (2009) Review of the rove beetle species of the subtribe Gyrophaenina Kraatz (Coleoptera, Staphylinidae) from New Brunswick, Canada: new species, provincial records and bionomic information. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:81–170 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Bourdon C (2012) Further contributions to the aleocharine fauna of the Yukon Territory, Canada (Coleoptera, Staphylinidae). In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:207–237 Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF,

336 Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Kraatz G (1856) Staphylinii. Naturgeschichte der Insecten Deutschlands. Erste Abtheilung Coleoptera. Zweiter Band. Erste, zweite Lieferung. Bogen 1–24. Nicolai, Berlin, viii + 376 pp Majka CG, Klimaszewski J (2008) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:85–114 Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39 Mäklin FW (1852) [New species and notes]. In: Mannerheim CG: Zweiter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Impériale Nat Moscou 25 (2):283–372 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp [separate, re-issued in Mémoires Présentées à l’Académie Impériale des Sciences de St.-Pétersbourg 1:415–501, in 1831] McLean JA, Klimaszewski J, Li A, Savard K (2009) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species. Part 1. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:5–17

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Tribe Homalotini Heer, 1839

Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Pace R (1989) Monografia del genere Leptusa Kraatz (Coleoptera, Staphylinidae). Mem Museo Civ Storia Nat Verona (2) (A: Biologica) 8:1–307 Paykull G (1800) Fauna Suecica. Insecta. Tomus III. J.F. Edman, Upsala, 459 pp Seevers CH (1951) A revision of the North American and European staphylinid beetles of the subtribe Gyrophaenae (Aleocharinae, Bolitocharini). Fieldiana Zool 32(10):657–762 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana Zool 71:vi + 289 pp Stephens JF (1832) (1832–1835) Illustrations of British Entomology... Mandibulata, V. Baldwin & Cradock, London. pp 1–240, pls 24–26 Strand A (1938) Gyrophaena orientalis n. sp. (Col., Staph.). Notulae Entomol 18(2):39–40 Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom. Berlingska Boktryckeriet, Lund [3] + 290 pp Webster RP, Klimaszewski J, Sweeney J, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:83–118

Tribe Placusini Mulsant and Rey, 1871

Tribe Placusini Mulsant and Rey, 1871 (Figs. 19.149–19.154) (Ashe 2001, Klimaszewski et al. 2001, 2018) Diagnosis. Body broad to moderately broad, moderately convex or flattened; pronotum with anterolateral angles obtuse, pubescence short, dense, on disc evenly distributed, directed approximately straight posteriad; labrum rounded medially with small a-sensillum; mandible with dorsobasal “velvety patch” modified to form a transverse row of large teeth, and without rows of denticles in ventral molar region; ligula short, broad and apically rounded; labial palpi short, each with two palpomeres; mesocoxae narrowly separated or contiguous; tarsal formula 4-4-5. Adults occur in subcortical habitats.

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to a maximum of half the mesocoxal length, narrowly triangular and pointed; mesocoxae contiguous medially; hind tarsi with basal segment elongate, at least as long as two following tarsomeres combined; male tergite VIII with a variable number of small to large spines or teeth, always with one incurved spine near each lateral margin; male sternite VIII obtusely angulate apically; median lobe of aedeagus in lateral view with a narrowly elongate, curved process of crista apicalis; flagellum long, with multiple coils basally (Tachyporoides group), or short and inconspicuous (Canadensis group); spermatheca simple, capsule small and roughly elongate-spherical, stem sinuate, sometimes with irregular posterior coils. Key to Species of Placusa

Genus Placusa Erichson, 1837 (Figs. 19.149–19.154) (Klimaszewski et al. 2001, 2018) Diagnosis. Body broad and strongly flattened; pronotum transverse, 1.5–1.7 times as broad as long, base arcuate, pubescence on disc usually short, dense, directed approximately straight posteriad, sometimes obliquely laterad, particularly along posterior part of median line and basally; terminal antennomere flattened, broad, moderately elongate; procoxal cavities open behind; process of mesoventrite short, projecting

1. Pronotum distinctly narrower than elytra at base (Figs. 19.149a and 19.151a) ......................2 – Pronotum about as wide as elytra at base or only slightly narrower (Figs. 19.150a, 19.152a– 19.154a) .............................................................3 2. Body uniformly black (Fig. 19.151a); male tergite VIII with two long lateral and three shorter median teeth (Fig. 19.151c); median lobe of aedeagus and spermatheca as illustrated (Fig. 19.151b, g–i) ..............Placusa vaga Casey – Body brown or brown with paler elytra and appendages (Fig. 19.149a); male tergite VIII with

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_19

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two lateral teeth and one short median tooth (Fig. 19.149c); median lobe of aedeagus and spermatheca as illustrated (Fig. 19.149b, f) ......... .......................................Placusa petulans Casey 3. Pronotum broadest closer to middle, flat and dull, granulation fine and very close, elytra with sides subparallel (Fig. 19.150a); male and female tergite VIII with highly elongate lateral teeth and lacking median teeth (Fig. 19.150c, e); median lobe of aedeagus and spermatheca as illustrated (Fig. 19.150b, g, h) .......Placusa tacomae Casey – Pronotum broadest behind middle, convex and glossy, granulation fine and distant, elytra distinctly widening apicad (Figs. 19.152a, 19.153a and 19.154a); male tergite VIII with short to moderately elongate sublateral teeth, and with smaller denticulations between (Figs. 19.152c, 19.153c and 19.154c); female tergite VIII entire and without projections (Figs. 19.152e, 19.153e and 19.154e); spermatheca as illustrated (Figs. 19.152g–i, 19.153g, h and 19.154g–i) ............................................................................4 4. Elytra slightly wider at apex than base of abdomen (Fig. 19.154a); male tergite VIII with numerous, elongate teeth medially (Fig. 19.154c); spermatheca as illustrated (Fig. 19.154g–i) ............................Placusa tachyporoides (Waltl) – Elytra distinctly wider at apex than base of abdomen (Figs. 19.152a and 19.153a); male tergite VIII with three very short median teeth (Figs. 19.152c and 19.153c); spermatheca not as above (Figs. 19.152g–i and 19.153g, h) ............5 5. Body length 2.0–2.7 mm; terminal antennomere strongly tapering apically (Fig. 19.153a); scutellum and adjoining area of elytra usually black, strongly contrasting with rest of elytra (Fig. 19.153a); female sternite VIII obtusely angulate apically (Fig. 19.153f) ............ ................Placusa pseudosuecica Klimaszewski – Body length 1.3–2.0 mm; terminal antennomere round apically (Fig. 19.152a); scutellum brown (Fig. 19.152a); aedeagus and spermatheca as illustrated (Fig. 19.152b, g–i); female sternite VIII almost evenly rounded

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Tribe Placusini Mulsant and Rey, 1871

apically (Fig. 19.152f).......................................... ............................... Placusa incompleta Sjöberg

Canadensis et al. 2001)

Species

Group (Klimaszewski

Diagnosis. Apical margin of male tergite VIII with sublateral spines long, curved (Figs. 19.149c, 19.150c and 19.151c); median lobe of aedeagus small, flagellum (if apparent) no longer than median lobe, bulbus with basal process wide, apico-ventral portion of bulbus broadly rounded. 114. Placusa petulans Casey (Fig. 19.149a–f) Placusa petulans Casey, 1911. Klimaszewski et al. 2001, Klimaszewski and Winchester 2002. Diagnosis. Body length 1.7–2.7 mm; colour more or less uniformly light rust-brown to brown or dark brown with rust-brown legs, elytra, and apex of abdomen (Fig. 19.149a); pubescence fine and short; forebody finely and closely granulate, punctation dense; head with genae shorter than eyes in dorsal view, arcuate and strongly narrowed basally in dorsal view, head 0.7 times as wide as maximum pronotal width; antennomeres VII–X at least twice as wide as long; pronotum distinctly narrower than elytra, transverse, maximum width 0.5–0.6 mm, length 0.3–0.4 mm, pronotal index ¼ 1.5; pubescence on disc directed approximately straight posteriad on both sides of median line; elytra transverse, maximum width 0.5–0.6 mm, length 0.3–0.4 mm, elytral index ¼ 1.7, pubescence on disc directed obliquely posteriad apically, approximately straight posteriad elsewhere. MALE. Tergite VIII with apical margin sinuately truncate between two long, curved, sharp sublateral teeth, with additional small tooth in middle (Fig. 19.149c); sternite VIII truncate apically; median lobe of aedeagus in lateral view with bulbus moderately large, tubus arcuate, apex truncate, flagellum short, coils in bulbus not apparent, process of crista apicalis wide, broadly arcuate

19

Tribe Placusini Mulsant and Rey, 1871

dorsally, slightly sinuate ventrally, and sharply pointed apically (Fig. 19.149b). FEMALE. Tergite VIII with apical margin arcuate (Fig. 19.149d); sternite VIII very obtusely produced apically (Fig. 19.149e); spermatheca with capsule small and roughly spherical, stem with middle and posterior portion strongly sinuate and S-shaped (Fig. 19.149f). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA, WA. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002). Habitat: Sitka spruce forest in Carmanah Valley, Vancouver Island, most adults were captured in transition zone, forest interior, and a few in clearcut. Collecting period: VI–X. Collecting method: pitfall traps. 115. Placusa tacomae Casey (Fig. 19.150a–h) Placusa tacomae Casey, 1893. Criddle 1922, Hatch 1957, Webster et al. 2009, Klimaszewski et al. 2001, 2011, 2016, 2018, Majka et al. 2011. Diagnosis. Body length 2.1–3.0 mm; colour dark brown or dark brown with legs, bases of antennae, elytra, and often pronotum lighter, rust-brown (Fig. 19.150a); pubescence fine and short; forebody finely and closely granulate, punctation dense; head with genae about as long as eyes in dorsal view, arcuate and strongly narrowed basally in dorsal view, head 0.8 times as wide as maximum pronotal width; antennomeres VII–X usually about twice as wide as long; pronotum transverse, broadest near middle, maximum width 0.5–0.6 mm, length 0.3–0.4 mm, pronotal index ¼ 1.5; pubescence on disc directed approximately straight posteriad on both sides of median line; elytra transverse, maximum width 0.5–0.7 mm, length 0.3–0.4 mm, elytral index ¼ 1.7, pubescence on disc directed obliquely posteriad apically, approximately straight posteriad elsewhere. MALE. Tergite VIII with apical margin sinuately truncate between two long, curved, sharp sublateral teeth

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(Fig. 19.150c); sternite VIII with apical margin triangularly produced, apex rounded (Fig. 19.150d); median lobe of aedeagus in lateral view with bulbus moderately large, tubus arcuate, apex truncate, flagellum short, coils in bulbus not apparent, process of crista apicalis wide, broadly arcuate dorsally, slightly sinuate ventrally, and sharply pointed apically (Fig. 19.150b). FEMALE. Tergite VIII unique in having apical margin with two long blunt lateral spine-like teeth and a broad subrectangular projection medially (Fig. 19.150e); sternite VIII very obtusely produced apically (Fig. 19.150f); spermatheca with capsule small and roughly spherical, stem with middle and posterior portion sinuate and S-shaped (Fig. 19.150g, h). Placusa tacomae is similar externally and in many genital features to the Palearctic P. complanata Erichson, recorded from Europe, Caucasus, and Siberia (Horion 1967). Externally the two species are not readily distinguishable, except that tergite VIII of the females are shaped differently. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, NF, NS, NT, ON, QC, SK, YT. USA: AZ, CO, MA, ME, MN, NY, WA, WI. New locality data: British Columbia, Hwy 16 W of Fishpan Lake N of Barrett, 54 30.660 N, 126 45.370 W, 834 m, 27.VII.2012, log pile, in Pinus contorta bark, A. Davies (CNC) 35 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: in various conifer and hardwood forests with conifers; white pine, red pine, jack pine, white spruce, sugar maple stands, rich Appalachian hardwood stand with some conifers, and mature red spruce and red maple forests (Klimaszewski et al. 2018). Adults under bark of conifers; under bark of fire-killed red pine (NF), under bark of red spruce infested with D. rufipennis Kirby and P. rufipennis (Kirby) (NB), and in bark of log pile of Pinus contorta; and in galleries of Dendroctonus ponderosae (Hopkins) (Alberta) and Ips pini (Say) (BC),

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presumably from conifers (Klimaszewski et al. 2018). In BC, one specimen was collected in log pile, in Pinus contorta Dougl. ex Loud. bark (CNC). Collecting period: V–IX. Collecting method: alpha-pinene baited Lindgren funnel traps, hand searching and collecting from under bark. 116. Placusa vaga Casey (Fig. 19.151a–i) Placusa vaga Casey, 1911. Gusarov 2003, Majka and Klimaszewski 2008, Webster et al. 2009, Klimaszewski et al. 2001, 2008, 2011, 2018, Brunke et al. 2012. Diagnosis. Body length 1.8–2.6 mm; colour uniformly dark brown to black, tarsi and palps reddish-brown (Fig. 19.151a); pubescence fine and short; forebody finely and closely granulate, punctation dense; head with genae about as long as eyes in dorsal view, subparallel behind eyes, then arcuately narrowed basad, 0.7–0.8 times as wide as maximum pronotal width; antennomeres VII–X usually less than twice as wide as long; pronotum moderately flattened, distinctly narrower than elytra at base, transverse, maximum width 0.4 mm, length 0.2–0.3 mm, pubescence on disc directed slightly obliquely posteriad on both sides of median line and approximately laterad near base; elytra transverse, maximum width 0.5–0.6 mm, length 0.4 mm, elytral index ¼ 1.3; pubescence on disc directed obliquely posteriad apically, approximately straight posteriad elsewhere. MALE. Tergite VIII with two long, sharp, incurved lateral teeth and three shorter median teeth (rarely two), blunt and subequal in length (Fig. 19.151c); sternite VIII obtusely triangularly produced apically, apex rounded (Fig. 19.151d); median lobe of aedeagus in lateral view with bulbus moderately large, tubus curved obtusely ventrad basally, ventral margin nearly straight to blunt apex, flagellum short, coils in bulbus not apparent, process of crista apicalis moderately wide, broadly arcuate dorsally, sharply pointed apically (Fig. 19.151b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 19.151e); sternite VIII slightly produced apically, apex rounded (Fig. 19.151f);

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Tribe Placusini Mulsant and Rey, 1871

spermatheca with capsule small, roughly elongate-spherical, stem with median portion narrowly U-shaped, posterior portion curled (Fig. 19.151g–i). Similar to the European P. atrata but differs in having the body colour nearly black (light brown to dark brown or nearly black with two brownish spots on the elytra in P. atrata), and the spermatheca with the stem narrowly U-shaped distally and more curved posteriorly. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, NS, NT, ON, QC, SK, YT. USA: CA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: mixed forests, old growth Abies balsamea, Picea glauca, Populus tremuloides with Picea glauca stand, oak savannah, and Populus forest. In scolytid galleries, under bark of Populus species (common), at a sap flow on a recently cut Populus sp., on sappy Populus wood, in decaying (moldy) corncobs and cornhusks, and in drift material on a river margin. Collecting period: V–IX. Collecting method: Multi-Pher 7 traps, pitfall traps; Lindgren funnel traps baited with alphapinene and 95% ethanol, and with 70% ethanol as preservative; malaise traps, hand searching subcortical habitats, sifting litter and decaying corncobs and cornhusks. Tachyporoides Species Group (Klimaszewski et al. 2001) Diagnosis. Apical margin of male tergite VIII with sublateral teeth usually very short; median lobe of aedeagus large, flagellum long, coiled in bulbus, at least twice as long as median lobe when everted, aedeagal process thin (except in P. despecta), and apico-ventral portion of bulbus keel-shaped. 117. Placusa incompleta Sjöberg { (Fig. 19.152a–i) Placusa incompleta Sjöberg, 1934. Palm 1959, Horion 1967, Klimaszewski and

19

Tribe Placusini Mulsant and Rey, 1871

Winchester 2002, Webster et al. 2009, Klimaszewski et al. 2001, 2011, 2015, 2016, 2018, Brunke et al. 2012, Work et al. 2013. Diagnosis. Body length 1.5–2.0 mm; colour uniformly dark brown or dark brown with legs, basal antennal segments, pronotum, elytra, and usually base of abdomen light brown (Fig. 19.152a); pubescence fine and short; forebody finely and closely granulate, punctation dense; head with genae shorter than eyes in dorsal view, subparallel behind eyes, then arcuately narrowed basally (sometimes slightly dilated), head 0.6 times as wide as maximum pronotal width; antennomeres VII–X usually at least twice as wide as long; pronotum moderately flattened, transverse, maximum width 0.5 mm, length 0.3 mm, pronotal index ¼ 1.6; pubescence on disc directed slightly obliquely posteriad on both sides of median line; elytra transverse, maximum width 0.5–0.6 mm, length 0.3 mm, elytral index ¼ 1.8; pubescence on disc directed obliquely apically, approximately straight posteriad elsewhere. MALE. Apical margin of tergite VIII truncate with two small obliquely triangular lateral teeth and usually three minute medial teeth (Fig. 19.152c); sternite VIII obtusely produced apically, apex rounded (Fig. 19.152d); median lobe of aedeagus in lateral view with enlarged bulbus, tubus arcuate, apex blunt, flagellum long, coiled 3–8 times in bulbus, process of crista apicalis very narrow, bent basally, straight to sharply pointed apex (Fig. 19.152b). FEMALE. Tergite VIII truncate apically (Fig. 19.152e); sternite VIII rounded apically (Fig. 19.152f); spermatheca with capsule asymmetrical, stem coiled and irregularly looped posteriad (Fig. 19.152g–i). There is moderate variation in the form of posterior loops of spermatheca. DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, NB, NF, NS, ON, QC, SK. USA: WA. New locality data: British Columbia, BColl., Hwy 16 W of Fishpan Lake, N of Barrett, 54 30.660 N, 126 45.370 W, 834 m, 26 and 27.

341

VII.2012, log pile, in Pinus contorta bark, A. Davies (CNC) 7 males, 7 females. COLLECTION AND HABITAT DATA. Habitat: in Abies balsamea stands, Acer saccharum stands, yellow birch forest, spruce moss forest, mature red spruce and red maple stand, old-growth white spruce and balsam fir forest (Klimaszewski et al. 2018), and in BC in coastal montane forests of central Vancouver Island with mature Douglas fir (Pseudotsuga menziesii (Mirb.)) and western hemlock (Tsuga heterophylla (Raf.) Sarg.), also in log pile, in Pinus contorta Dougl. ex Loud. bark. In Europe, Betula, Populus, Quercus, and conifer forests. Adults from under bark of tamarack infested by eastern bark beetle, under bark of red spruce infested with Dendroctonus rufipennis (Kirby) and Polygraphus rufipennis (Kirby). One from gilled mushroom. Collecting period: VI–IX. Collecting method: Lindgren funnel traps with 70–75% ethanol, or baited with alpha-pinene and 95% ethanol, Lindgren funnel trap with frontalin and ethanol as attractants for Coleoptera, Luminoc pit-light trap, pitfall traps, carrion trap, hand searching subcortical habitats, and sifting mushroom. Many specimens were trapped near the forest floor. 118. Placusa pseudosuecica Klimaszewski (Fig. 19.153a–h) Placusa pseudosuecica Klimaszewski, in Klimaszewski et al. 2001. Klimaszewski et al. 2015, 2016, 2018. Diagnosis. Body length 2.0–2.7 mm; colour uniformly light to dark brown or dark brown with legs, basal antennal segments, elytra, base and tip of abdomen light brown to rust-brown (Fig. 19.153a); pubescence moderately long; forebody finely and closely granulate, punctation moderately dense; head with genae about as long as eyes in dorsal view, broadly arcuate and gradually narrowed basally (not dilated), head 0.6 times as wide as maximum pronotal width; antennomeres VII–X usually less than twice as

342

wide as long (average 1.8 times as wide as long); pronotum broadest near middle, moderately flattened, transverse, maximum width 0.5–0.6 mm, length 0.3–0.4 mm, pronotal index ¼ 1.5; pubescence on disc directed slightly obliquely posteriad on both sides of median line; elytra transverse, wider apically than at base, maximum width 0.7–0.8 mm, length 0.3–0.4 mm, elytral index ¼ 2.1; pubescence on disc directed obliquely apically, approximately straight posteriad elsewhere. MALE. Apical margin of tergite VIII truncate, with two short, acutely triangular lateral teeth, and 3 min median teeth (Fig. 19.153c); sternite VIII obtusely angulately produced apically, apex rounded (Fig. 19.153d); median lobe of aedeagus in lateral view with enlarged bulbus, tubus arcuate, apex blunt, flagellum long, coiled 1.5–5 times in bulbus, process of crista apicalis narrow, broadly arcuately tapering to sharply pointed apex (Fig. 19.153b). FEMALE. Tergite VIII arcuate apically (Fig. 19.153e); sternite VIII obtusely triangularly produced apically, apex rounded (Fig. 19.153f); spermatheca with capsule large, spherical, stem (duct) narrowly U-shaped, slightly sinuate posteriorly (Fig. 19.153g, h).

19

Tribe Placusini Mulsant and Rey, 1871

Comments. Placusa pseudosuecica is externally and genitalically similar to the European P. suecica Johnson and Lundberg and P. cribrata Johnson and Lundberg. It differs externally from both species in having antennomeres VI–X less transverse, usually distinctly less than twice as broad as long (distinctly more than twice as broad as long in P. suecica and P. cribrata), and the pronotum is narrower in P. cribrata (width 0.5 mm, length 0.3 mm, pronotal index ¼ 1.6). The median lobe of the aedeagus and male tergite VIII are similar in the three species. The spermatheca provides the most reliable features and consistent differences for specific identification. In P. pseudosuecica, the spermatheca has the longest stem (duct), which is more broadly arched at the middle and is sinuate at the base in both Quebec and British Colombia specimens, whereas it is either short and slightly arched posteriorly in P. cribrata, or longer but straight posteriorly in P. suecica. 119. Placusa tachyporoides (Waltl) { (Fig. 19.154a–i)

DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, ON, QC, SK. USA: not recorded.

Aleochara tachyporoides Waltl, 1838. Horion 1967, Moore and Legner 1975, Klimaszewski et al. 2001, 2007, 2015, 2016, 2018, Majka and Klimaszewski 2008.

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: balsam fir stands, black spruce stands, white spruce stands, a spruce/fir stand, trembling aspen dominated stands, and a sugar maple stand; in BC—western hemlock, Sitka spruce (Picea sitchensis (Bongard)), and Pacific silver fir (Abies amabilis (Douglas ex Louden)) forest. In general, the species seems to prefer mature conifer forests in eastern as well as western Canada. Nothing has been recorded about the specific habitat associations, but it is probably found in subcortical habitats like its congeners. Collecting period: VI–X. Collecting method: Malaise traps, Lindgren funnel traps, and ethanol-baited Lindgren funnel traps.

Diagnosis. Body length 1.8–3.0 mm; colour uniformly dark brown or dark brown with legs, basal antennal segments and elytra light brown to yellowish/reddish brown (Fig. 19.154a); pubescence fine and short; forebody finely and closely granulate, punctation moderately dense; head with genae shorter than eyes in dorsal view, broadly arcuate and gradually narrowed basally (not dilated); head 0.6 times as wide as maximum pronotal width; antennomeres VII–X usually less than twice as wide as long; pronotum moderately flattened, transverse, maximum width 0.5 mm, length 0.3 mm, pronotal index ¼ 1.6; pubescence on disc directed slightly obliquely posteriad on both sides of median line; elytra transverse, maximum width 0.5–0.7 mm, length 0.3–0.4 mm, elytral index ¼ 1.7; pubescence on disc directed

19

Tribe Placusini Mulsant and Rey, 1871

obliquely posteriad apically, approximately straight posteriad elsewhere. MALE. Apical margin of tergite VIII truncate, with two larger lateral incurved teeth and several (3–8, usually 5, in combinations: 1+1+2, 2+1+2, 1+1+1, 2+1+3, 2 +2+3, 3+1+3, 3+2+3) medial teeth varying in length from short and sharply pointed to long, rounded apically and approaching the length of the lateral teeth (Fig. 19.154c); sternite VIII obliquely triangularly produced apically, apex rounded (Fig. 19.154d); median lobe of aedeagus with enlarged bulbus, tubus arcuate, ventral margin slightly sinuate, apex blunt in lateral view, flagellum long, coiled 1.5–3 times in bulbus, process of crista apicalis narrow, slightly sinuate and sharply pointed apically (Fig. 19.154b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 19.154e); sternite VIII broadly rounded apically (Fig. 19.154f); spermatheca with capsule asymmetrical, stem U-shaped, doubled back to irregularly coiled base (Fig. 19.154g–i). Placusa tachyporoides is highly variable in body size, colour, antennal structure, shape of the spermatheca, and apical dentation of male tergite VIII. This species may be confused with P. incompleta but in P. tachyporoides the male tergite VIII has longer and more numerous teeth, and the U-shaped stem of the spermatheca is broader and longer.

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DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, NB, NS, ON, QC, SK. USA: CA, MA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: Sugar maple stands, balsam fir, and black spruce stands, mixed trembling aspen, white spruce, and white pine stands, and a yellow birch forest in QC. In NB, collected from Lindgren funnel traps in old-growth northern hardwood forest, a red oak forest, an old jack pine forest, and in a rich Appalachian hardwood forest. In Europe, in Acer, Alnus, Betula, Carpinus, Fagus, Populus, Quercus, Ulmus, Sorbus, and coniferous forests. Two individuals from NB sifted from pile of fairly fresh corncobs and cornhusks, otherwise nothing has been recorded about its microhabitat associations. Collecting period: V–IX. Collecting method: Lindgren funnel traps baited with alpha-pinene and 95% ethanol or with 95% ethanol only, Luminoc pit-light traps, some in unbaited Multi-Pher pitfall traps, and flight interception traps, and pitfall traps. There are some indications that this species, like P. incompleta, is attracted to ethanol-baited traps and to light.

344

Fig. 19.149 (a–f) Images of Placusa petulans Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) female tergite VIII; (e) female sternite

19

Tribe Placusini Mulsant and Rey, 1871

VIII; (f) spermatheca. (b, c, f) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

19

Tribe Placusini Mulsant and Rey, 1871

Fig. 19.150 (a–h) Images of Placusa tacomae Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g, h)

345

spermatheca. (b, g, h) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

346

Fig. 19.151 (a–i) Images of Placusa vaga Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

19

Tribe Placusini Mulsant and Rey, 1871

(e) female tergite VIII; (f) female sternite VIII; (g–i) spermatheca. (b, g–i) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

19

Tribe Placusini Mulsant and Rey, 1871

Fig. 19.152 (a–i) Images of Placusa incompleta Sjöberg: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite

347

VIII; (g–i) spermatheca. (b, g–i) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

348

Fig. 19.153 (a–h) Images of Placusa pseudosuecica Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; f, female sternite

19

Tribe Placusini Mulsant and Rey, 1871

VIII; (g, h) spermatheca. (b, g, h) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

19

Tribe Placusini Mulsant and Rey, 1871

Fig. 19.154 (a–i) Images of Placusa tachyporoides (Waltl): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e) female tergite

349

VIII; (f) female sternite VIII; (g–i) spermatheca. (b, c, g–i) after Klimaszewski et al. (2001). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

350

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II ZooKeys [Special Issue] 186:119–206 Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7 [1892–1894]:281–606 [see reference in Chapter 7 for explanation of date] Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing, Lancaster, 259 pp Criddle N (1922) The entomological record, 1921. Annu Rep Entomol Soc Ontario 52 [1921]:57–70 Erichson WF (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Gusarov VI (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353:1–134 Hatch MH (1957) The beetles of the Pacific Northwest. Part II: Staphyliniformia. Univ Wash Publ Biol 16:IX + 384 pp Horion A (1967) Faunistik der Mitteleuropȁischen Kȁfer. Bd. XI: Staphylinidae. 3. Teil: Habrocerinae bis Aleocharinae (ohne Subtribus Athetae). Űberlingen -Bodensee, P.C.V. Schmidt, xxiv + 419 pp Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belge Entomol 40:3–126 Klimaszewski J, Pelletier G, Germain C, Hébert C, Humble LM, Winchester NN (2001) Diversity of Placusa (Coleoptera: Staphylinidae, Aleocharinae) in Canada, with descriptions of two new species. Can Entomol 133(1):1–47 Klimaszewski J, Langor D, Savard K, Pelletier G, Chandler DS, Sweeney J (2007) Rove beetles (Coleoptera: Staphylinidae) in yellow birch-dominated stands of southeastern Quebec, Canada: diversity, abundance, and description of a new species. Can Entomol 139 (6):793–833 Klimaszewski J, Langor DW, Work TT, Hammond JHE, Savard K (2008) Smaller and more numerous harvesting gaps emulate natural forest disturbances: a biodiversity test case using rove beetles (Coleoptera, Staphylinidae). Divers Distrib 14(6):969–982 Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera,

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Tribe Placusini Mulsant and Rey, 1871

Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Majka CG, Klimaszewski J (2008) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:85–114 Majka CG, Chandler DS, Donahue CP (2011) Checklist of the beetles of Maine, USA. Empty Mirrors Press, Halifax, Nova Scotia, 328 pp Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Mulsant E, Rey C (1871) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens [Huitième Branche, Bolitocharaires]. Deyrolle Fils, Paris. [4] + 321 pp + [5, explic.] + [4, table, erratum], pls 1–5 Palm T (1959) Die Holz- und Rinden-käfer der süd- und mittelschwedischen Laubbäume. Opusc Entomol Suppl 16:5–374 + 93 figs Sjöberg O (1934) Zwei neue Bolitocharinen aus Schweden. Entomol Tidskr 55:281–285 Waltl J (1838) Verzeichniss der um Passau vorkommenden seltnern Käfer [cols. 263–267]; Um Passau vorkommende neue Arten von Käfern [cols. 267–273]. In: Beiträge zur nähern naturhistorischen Kenntniss des Unterdonaukreises in Bayern. Isis von Oken 1838(4):250–273 Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:171–248 Work TT, Klimaszewski J, Thiffault E, Bourdon C, Paré D, Bousquet Y, Venier L, Titus B (2013) Initial responses of rove and ground beetles (Coleoptera, Staphylinidae, Carabidae) to removal of logging residues following clearcut harvesting in the boreal forest of Quebec, Canada. ZooKeys 258:31–52

Tribe Athetini Casey, 1910

Tribe Athetini Casey, 1910 (Figs. 20.155–20.257) (Seevers 1978; Ashe 2001; Klimaszewski and Winchester 2002; Elven et al. 2010, 2012; Klimaszewski et al. 2018b) Comments. The tribe Athetini is by far the largest evolutionary lineage of aleocharines. It is one of the most difficult and poorly known tribes of beetles in North America and elsewhere. This situation is caused by various factors: enormous species diversity, overwhelming number of species descriptions without illustrations, the small size of the majority of species, difficult-toobserve diagnostic features, convergent character states, extensive literature, and the lack of studies of genitalic structures by most previous authors due to a lack of awareness of the value of genitalic characters. Most Nearctic species were described by Casey (1885, 1893, 1906, 1910, 1911), and are grouped in a large number of genera (subgenera), with often wrongly assigned species, genera (subgenera) poorly defined with overlapping characteristics. Ranking at the generic or subgeneric level is a serious problem, particularly in the large genus Atheta (sensu lato). At the present stage of knowledge of this genus, it is neither practical nor possible to elevate subgenera of Atheta to a generic status and place all the species in generic taxa because these taxa, are little understood and inadequately defined. The elevation of subgenera to generic rank taxa should be executed when our knowledge of

20

subgeneric taxa will be adequate. In this book, we treat the genus Atheta in a broad sense (sensu lato), and like most European authors (e.g., Palm 1970; Benick and Lohse 1974), we use subgeneric classification with the exceptions of those genera that are clearly delimited e.g., Acrotona, Amischa, Boreophilia, Earota, Philhygra, etc. Elven et al. (2012) conducted a molecular phylogenetic analysis that included many Nearctic genera (Amischa, Atheta, Boreostiba, Boreophilia, Dalotia, Hydrosmecta, Liogluta, Lypoglossa, Meronera, Mocyta, Myrmedonota, Stethusa, Strigota, and Tarphiota) and confirmed their position within Athetini. The current classification of Nearctic athetines is inadequate and many subgenus-level taxa currently recognized cannot be clearly delimited and diagnosed. Subtribe Athetina Casey, 1910 (Ashe 2001; Klimaszewski et al. 2018b) Comments. This subtribe is very poorly defined so a meaningful diagnosis is not possible. Currently the subtribe includes many different groups of athetines and likely does not represent a monophyletic group. We have adopted a broad definition of this subtribe to include diverse groups of the genus Atheta and those genera/subgenera that do not belong to the tribe Lomechusini. Only one genus of Athetini in British Columbia— Paragoniusa—is not assigned to this subtribe,

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_20

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but its subtribe affiliation is unclear. The genera and species within species groups are listed in alphabetical order. In recent molecular studies (Elven et al. 2010, 2012), a diverse group of athetine genera, including two genera occurring in British Columbia (Aloconota and Earota) were included in a newly proposed tribe Geostibini, which was considered a sister group of Lomechusini. Morphologically, the tribe Geostibini was defined by a simple autapomorphy of a reduced sensillum “a” of the epipharynx. We believe that more morphological evidence should be presented before this new classification can be accepted and therefore in this book these genera are listed alphabetically under a broadly defined tribe Athetini. The emphasis in this book is on species identification and grouping diverse genera in a tribe that cannot be easily morphologically defined would cause serious problems for the users. The genera Aloconota and Earota are more easily identifiable when left in Athetini. Key to Genera of Athetini Casey 1. Head either broadly oval or widened posteriad so the greatest width is behind eyes, constricted at base, eye small less than length of temple (Fig. 20.158a); mesocoxal cavities not margined; ligula short, broad and entire ............................... ..................................... Amischa C.G. Thomson – Head and other characters not as above ........2 2. Body form distinct (Figs. 20.240a, 20.250a, and 20.251a–20.255a); integument of forebody strongly granulose, microsculpture coarse (Figs. 20.240a, 20.250a, and 20.251a–20.255a); species restricted in distribution to sandy sea beaches ...............................................................3 – Body form not as above; integument of forebody not strongly granulose as above, microsculpture variable but weak in most; species living in terrestrial habitats and may be only accidentally found on sandy sea beaches (exception Adota Fig. 20.155, but integument not strongly granulose)...........................................................5

20

Tribe Athetini Casey, 1910

3. Elytra distinctly shorter than pronotum (Figs. 20.240a, and 20.252a–20.254a); hind wings absent.......................................................4 – Elytra longer than pronotum (Figs. 20.250a and 20.251a); hind wings present ........................ .................................................. Tarphiota Casey 4. Elytra with shoulders rounded (Fig. 20.240a), surface dull (Fig. 20.240a); elytra in most specimens much paler that remainder of body and appearing cream-white (Fig. 20.240a) ..............................................Pontomalota Casey – Elytra with shoulders angular (Figs. 20.252a, 20.253a, and 20.254a), elytra dark of the same colour as remainder of body or slightly paler (Figs. 20.252a and 20.253a) ....... Thinusa Casey 5. Pronotum with hypomera not visible in lateral view (edge of pronotum entire in lateral view and without visible structure below); body as illustrated (Figs. 20.226a–20.228a, 20.248a, and 20.249a) .............................................................6 – Pronotal hypomera clearly or partially (Lypoglossa) visible in lateral view; body not as above ..................................................................7 6. Antennae thick, antennomeres V–X more or less transverse (Figs. 20.248a and 20.249a); body narrowly elongate, densely punctate, particularly on abdomen, dorsal surface with fine white pilose pubescence (Figs. 20.248a and 20.249a; pronotum approximately as broad as maximum width of elytra (Figs. 20.248a and 20.249a); tergite VIII in both sexes with the basal line (antecostal suture) joining the base of tergite (Figs. 20.248d, f, and 20.249d, f), but not the sides of the disc as in other aleocharines; apical margin of female sternite VIII with row of strong microsetae on its dorsal side ........Strigota Casey – Antennae thin, antennomeres V–X subquadrate or slightly transverse (Figs. 20.226a–20.228a); body broadly to narrowly elongate, moderately densely punctate, pubescence different (Figs. 20.226a–20.228a); pronotum often broader than maximum width of elytra; tergite VIII in both sexes with the basal line joining sides of the disc (Figs. 20.226d, f, 20.227d, and 20.228c, e); apical

20

Tribe Athetini Casey, 1910

353

margin of female sternite VIII with less strongly developed apical microsetae.................................. ..................................... Mocyta Mulsant and Rey

– Basal antennomere not distinctly swollen (Figs. 20.189a–20.194a); abdomen subparallel............ Atheta (Microdota) Mulsant and Rey

7. Body slender, subparallel, length 1.4–3.0 mm (on average 2.0 mm) (Figs. 20.189a–20.194a, 20.207a, 20.215a, 20.230a, and 20.232a); head with genae elongate in dorsal view, and at least as long as diameter of eye; pronotum as wide as base of elytra or only slightly narrower .............................................................8 – Body less slender and broader, length on average 3.0 mm; other characters different or in different combinations.......................................... 12

12. Body form distinctive, head and pronotum much narrower than elytra which is at least 1.2 times as wide pronotum (Figs. 20.255a and 20.256a); abdomen oval in outline (Figs. 20.255a and 20.256a); forebody with pubescence long and shaggy, erect on pronotum (Figs. 20.255a and 20.256a) .....Trichiusa Casey – Body form and pubescence not as above....13

8. Eyes small, one tenth to half as long as genae in dorsal view or completely reduced (Figs. 20.215a, 20.230a, and 20.232a); elytra at suture as long as or shorter than pronotum along midline (Figs. 20.215a, 20.230a, and 20.232a) ............................................................................9 – Eyes larger, longer that half length of genae in dorsal view (Figs. 20.189a–20.194a, and 20.207a); elytra at suture longer than pronotum along midline ...................................................11 9. Pronotum highly glossy and distinctly margined laterally, pronotal pubescence and punctation sparse (Fig. 20.232a); genitalia as illustrated (Fig. 20.232b, c, h)................. Pelioptera Kraatz – Pronotum moderately glossy and not distinctly margined laterally, pronotal pubescence moderately dense (Figs. 20.215a and 20.230a); genitalia not as above .....................................................10 10. Elytra subparallel (Fig. 20.215a); ligula divided into two contiguous lobes; posterior part of spermathecal stem swollen (Fig. 20.215h) ..................................... Geostiba C.G. Thomson – Elytra broadening posteriad (Fig. 20.230); ligula divided into two widely separated lobes; posterior part of spermathecal stem not swollen (Fig. 20.230g)........Ousipalia Casey 11. Basal antennomere distinctly swollen (Fig. 20.207a); abdomen distinctly arcuate laterally........................................ Clusiota Casey

13. Pronotum exceptionally broad and shieldlike, 1.5–2.3 times as wide as long (Figs. 20.216a, 20.224a, and 20.257a)............. 14 – Pronotum not as above, at most 1.4 times as wide as long .....................................................16 14. Pronotum 1.5 times as wide as long (Figs. 20.224a and 20.225a) ................................ ...............................................Lypoglossa Fenyes – Pronotum at least 2 times as wide as long (Figs. 20.216a and 20.257a) ............................ 15 15. Pubescence of forebody very short and dense (Fig. 20.257a); pronotum about 2 times as wide as long with basal angles of disc slightly angulate to rounded and basal margin distinctly sinuate (Fig. 20.257a)....................................................... .....Paragoniusa Maruyama and Klimaszewski [genus not recorded yet in BC but most likely present there] – Pubescence of forebody moderately long and moderately dense (Fig. 20.216a); pronotum about 1.6 times as wide as long with basal angles of disc strongly angulate and basal margin arcuate (Fig. 20.216a)..............................Goniusa Casey 16. Pronotum with pubescence on midline of disc directed anteriad and posteriad basally and inwards towards midline in central area of disc (Fig. 20.247a); body highly glossy; male tergite VII with a weak to moderately strong median carina....................................... Seeversiella Ashe – Pronotum with different pattern of pubescence; male tergite III with lateroapical angles

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not prolonged into spines and without median carina................................................................17 17. Mandibles both appearing bifid (left mandible with tip bifid and right mandible bears one apical and one subapical tooth creating illusion of bifid mandible); body subparallel; pronotum slightly transverse (at most 1.3 times as wide as long) to nearly subquadrate, integument glossy, microsculpture weak (Figs. 20.243a–20.246a) ...........................................Schistoglossa Kraatz – Mandibles acutely pointed and not bifid; other characters not as above ....................................18 18. Body narrowly subparallel (Figs. 20.156a, 20.157a, 20.217a–20.223a, and 20.233a– 20.239a); head about as long as pronotum or slightly shorter, genae at least as long as eyes in dorsal view (Figs. 20.156a, 20.157a, 20.217a– 20.223a, and 20.233a–20.239a); elytra at suture at least as long as pronotum............................. 19 – Characters or combination of characters not as above ................................................................21 19. Empodial bristles much longer than tarsal claws; pronotum subquadrate (Figs. 20.156a and 20.157a), often with distinctive pattern of pubescence (wavy, V-shaped) in males; male tergite VIII with apical margin bearing two lateral teeth and two median smaller teeth or protuberances (Figs. 20.156c and 20.157d); spermatheca S-shaped (Figs. 20.156g and 20.157h) ................................... Aloconota C.G. Thomson – Characters or combination of characters not as above ................................................................20 20. Pronotum with pubescence on disc directed posteriad along midline, either posteriad or posterolaterad elsewhere (Figs. 20.217a– 20.223a); apical margin of male tergite VIII often with broad rectangular projection, often crenulate, with two lateral teeth, punctation often asperate near apex of disc (Figs. 20.217c, 20.218c, and 20.219c); spermatheca large (Figs. 20.219g, 20.220g, 20.221g, 20.222h, and 20.223g) ....................... Liogluta C.G. Thomson – Pronotum with pubescence on disc directed anteriad along midline, laterad elsewhere

20

Tribe Athetini Casey, 1910

(Figs. 20.233a–20.239a); male tergite VIII truncate apically, without additional structures, punctation not asperate (Figs. 20.233c, 20.234d, 20.235d, 20.236c, 20.237d, 20.238c, and 20.239c); male sternite VIII very elongate, often projecting posteriad under pygidium (Figs. 20.233d, 20.234e, 20.235e, 20.236d, 20.237e, 20.238d, and 20.239d); spermatheca vestigial and difficult to find, additional membraneous structures present in female pygidium (Figs. 20.233g, 20.234h, 20.235h, 20.236g, 20.237h, i, 20.238g, and 20.239g) ....................... ............................... Philhygra Mulsant and Rey 21. Infraorbital carina absent; body flattened in most (Figs. 20.209a–20.212a); forebody with distinct microsculpture; mainly subcortical species ........................ Dinaraea C.G. Thomson – Infraorbital carina partial or complete but always present; body convex to moderately flattened; forebody with weak microsculpture in most; not subcortical species............................ 22 22. Pubescence on pronotum and elytra dense, appearing silky (Figs. 20.157a and 20.214a) (Except Dochmonota, Fig 20.213a); apical antennomere almost as long as previous three combined (Except Dochmonota, Fig 20.213a); ligula with wide base, split into two lobes in apical third; mesocoxae broadly separated by distance about equal to length of isthmus ............ 23 – Combination of characters not as above .....25 23. Body small, length 2.2–2.5 mm, narrowly ovoid, with dense punctation (Fig. 20.213a); antennomeres VI–X subquadrate to slightly transverse (Fig. 20.213a); male sternite VIII highly elongate, apical part triangularly produced, apex obtuse (Fig. 20.213e); median lobe of aedeagus with large sac-shaped crista apicalis on apical margin of prominent triangular projection of bulbus (Fig. 20.213b); spermathecal capsule hemispherical, with narrow, very deep apical invagination (Fig. 20.213h).................................. ...............................Dochmonota C.G. Thomson – Body larger, over 2.5 mm long, other combination of characters not as above.....................24

20

Tribe Athetini Casey, 1910

24. Antennomeres VII–X strongly transverse (Fig. 20.214a); pubescence on pronotum and elytra short, dense, appearing silky, directed approximately straight posteriad (Fig. 20.214a); male tergite VIII produced apically and not serrate (Fig. 20.214d); median lobe of aedeagus and spermathecal as illustrated (Fig. 20.214b, c, h); occurs in terrestrial habitats ................................. ............................................Earota Mulsant and – Antennomeres VII–X subquadrate to slightly transverse (Fig. 20.155a); pubescence on pronotum and elytra moderately long and dense, slightly silky, directed mostly laterad and forming wavy pattern on elytra (Fig. 20.155a); male tergite VIII truncate apically and with four rounded teeth (Fig. 20.155d); spermathecal capsule tubular and flattened (Fig. 20.155h); occurs in debris on sea beaches.................................Adota Casey 25. Procoxal cavities separated from lateral margin of pronotum by long and wide hypomeron; apical margin of male tergite VIII entire, without teeth or crenulation (Figs. 20.204e, 20.205d, and 20.206d); body broad in most (Figs. 20.204a, 20.205a, and 20.206a)......................................26 – Procoxal cavities reaching lateral margin of pronotum or separated from lateral margin of pronotum by a narrow part of hypomeron; apical margin of male tergite VIII usually with teeth or some degree of crenulation in most; body moderately broad or narrow ....................................... 27 26. Ligula divided to base into two divergent lobes; antennomeres VII–X quadrate (Fig. 20.204a) or moderately transverse to elongate ................................Boreophilia Benick – Ligula divided into two lobes only in apical half; antennomeres VII–X moderately to distinctly transverse (Figs. 20.205a, 20.206a) .......... ................................................. Boreostiba Lohse 27. Elytron virtually lacking emargination at the posterolateral angle (Fig. 20.231a); interior edge of lacinia without spines; ligula reduced and not apparent; median lobe of aedeagus very narrow and lacking clearly visible athetine bridge (Fig. 20.231b–e)..... Paraleptonia Klimaszewski

355

– Elytron with emargination at the posterolateral angle; interior edge of lacinia with spines; ligula present and split apically; median lobe of aedeagus with clearly visible athetine bridge ..28 28. Entire body covered by strong isodiametric microsculpture (Figs. 20.241a and 20.242a); frontal margin of labrum concave; last tarsomere of all tarsi lacking empodial setae; tarsal claws of different length; median lobe of aedeagus and spermathecal as illustrated (Figs. 20.241b, c, h and 20.242b, c, h) ............................................. ........................ Psammostiba Yosii and Sawada – Characters not as above or in different combination......................................................29 29. Median lobe of aedeagus with highly enlarged bulbus and very short, broad triangular tubus in lateral view (Fig. 20.208b); spermatheca with large capsule divided by a median seam, apical half narrower, hemispherical, stem short and strongly sinuate (Fig. 20.208g); male tergite VIII with two lateral teeth and distinctive broadly V-shaped apical emargination with several irregular in size small teeth (Fig. 20.208c); habitus as illustrated (Fig. 20.208a).............. Dalotia Casey – Median lobe of aedeagus with moderatelysized bulbus and moderately narrow to broad subparallel or triangular tubus in lateral view (Figs. 20.159b, 20.160b, 20.161b, c, and 20.162b, c); spermatheca variable in shape but with club-shaped capsule and moderate to long sinuate stem looped posteriorly in most species (Figs. 20.159h, 20.160h, 20.161h, and 20.162i); male tergite VIII often with two lateral teeth and differently shaped apical margin of disc (Figs. 20.160d, 20.164d, 20.165d, and 20.168d); habitus variable, as illustrated (Figs. 20.160a– 20.201a) .......................... Atheta C.G. Thomson

Genus Adota Casey, 1910 (Fig. 20.155a–h) (Casey 1910; Fenyes 1920; Seevers 1978; Lohse and Smetana 1985; Ashe 2001; Gusarov 2003d)

356

Diagnosis. Body length 2.2–3.2 mm, body parallel-sided and flat, colour brown to black, elytra dark brown with some reddish tinge, antennae black to brown, legs brown to yellowish brown, entire body with strong and dense isodiametric microsculpture; head as long as wide; eyes large, temple length to eye length ratio 0.8–1.2; infraorbital carina very short, ends in posterior portion of temples; antennomere IV–VI subquadrate, VII–X subquadrate to slightly transverse, terminal one without coeloconic sensilla, about as long as articles IX–X combined; labrum transverse, with concave anterior margin; mandibles broad, right mandible with a small medial tooth, dorsal molar area with velvety patch consisting of very small denticles; maxilla with galea projecting slightly beyond apex of lacinia; apical lobe of galea covered with numerous fine and short setae; internal margin of galea with long subapical setae; apical 1/3 of lacinia with row of closely spaced spines, middle portion produced medially and covered with numerous setae; ligula short, with broad base and split apex; medial area of prementum with 2 pores and with 8–23 pseudopores, lateral areas each with two asetose pores, single setose pore and 7–12 pseudopores; labial palpus with setae a, b, g and d present (Fig. 10, in Gusarov 2003d); mentum with concave anterior margin; pronotum slightly transverse, with microsetae directed anteriorly in midline, in lateral portions of disc microsetae directed laterally, macrosetae short; hypomera fully visible in lateral view; mesosternal process narrow, extending about 3/5 length of mesocoxal cavities, metasternal process short, mesoventrite and mesosternal process not carinate medially; relative lengths of mesoventral process: isthmus: metaventral process in ratio of about 3:1:1; mesocoxal cavities margined posteriorly; mesocoxae narrowly separated; tarsal formula 4-5-5, metatarsomere I as long as II; all tarsomeres with long setae; one empodial seta, as long as claws; claws strong of equal length; elytral posterior margin straight; wings fully developed; abdominal tergites III–VI with moderate basal impressions, tergum VII as long as

20

Tribe Athetini Casey, 1910

tergum VI, punctation on tergites VI–VII only slightly sparser than on III–V, tergum VII with wide white palisade fringe. Adota can be distinguished from Atheta by the concave anterior margin of the labrum; by basal impression on the tergum VI, and by lacking distinct medial lamellae (sclerites) of internal sac. Adota differs from Psammostiba in having apical margin of male tergite VIII with 2–4 teeth (serrate in Psammostiba), and tarsi with single empodial seta and equally long claws. 120. Adota maritima (Fig. 20.155a–h)

(Mannerheim)

Homalota maritima Mannerheim, 1843. Casey 1910, Fenyes 1920, Moore and Legner 1975, Seevers 1978, Lohse and Smetana 1985, Gusarov 2003d (complete list of synonyms). Diagnosis (adopted from Gusarov 2003d). Body length 2.5–3.2 mm, colour dark brown or black with light brown tarsi (Fig. 20.155a); head surface dull, with strong and dense isodiametric microsculpture, with fine punctation, distance between punctures equals 1–2 times their diameter; frons with sparser punctation, in both sexes with longitudinal impression; temple length to eye length ratio 1.0–1.2; antennomere II about 1.1 times as long as antennomere III, IV–V slightly elongate (length to width ratio 1.2–1.3), VI–VII as long as wide, VIII–X slightly transverse (length to width ratio 0.8); pronotum subquadrate or slightly transverse, 1.1 times as wide as head, width 0.47–0.61 mm, length 0.41–0.51 mm, width to length ratio 1.2, with strong and dense isodiametric microsculpture; punctation finer than on head, poorly visible; elytra much wider (0.63–0.89 mm) and longer (0.57–0.79 mm; measured from humeral angle) than pronotum (elytral length to pronotal length ratio 1.5), 1.1 times wider than long, surface matte, with strong and dense isodiametric microsculpture; punctation poorly visible; abdominal tergites dull with strong and dense

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Tribe Athetini Casey, 1910

isodiametric microsculpture and fine punctation, distance between punctures equals 2–3 times their diameter on tergites III–V and 3–5 times on tergites VI–VII, apical margin of tergite VII with white palisade fringe. MALE. Tergite VIII with posterior margin emarginate medially and with four rounded teeth (Fig. 20.155d); sternite VIII parabolic posteriorly (Fig. 20.155e); tubus of median lobe of aedeagus slightly sinuate ventrally, with narrowly rounded apex in lateral view (Fig. 20.155b); in dorsal view as illustrated (Fig. 20.155c). FEMALE. Tergite VIII arcuate apically (Fig. 20.155f); sternite VIII broadly rounded apically (Fig. 20.155g); spermatheca with tubular capsule bearing broad and short apical invagination, stem narrow, sinuate and narrowly looped posteriorly (Fig. 20.155h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA. New locality and habitat data: British Columbia, Queen Charlotte Is., NW corner Graham Is., Lepas Bay, 17.VIII.1983, sifting seaweed and forest floor litter, J.M. Campbell (CNC) 1 male; 50 sex undetermined; BC, Queen Charlotte Is., 12.8 km S Tlell R. Bridge, 17.VII.1983, 03–15, ex rotting seaweed, J.M. Campbell (CNC) 3 males, 4 females, 3 sex undetermined; BC, Queen Charlotte Is., Tlell, 17.VII.1983, 83-16, ex rotting seaweed, J.M. Campbell (CNC) 2 females; BC, Queen Charlotte Is., Ramsay Is., N shore, 1.5 km E of west end, 11.VIII.1983, 83-74, sifting seaweed, J.M. Campbell (CNC) 5 sex undetermined; BC, Queen Charlotte Is., 14 km E Masset, 13.VII.1983, 83-5, treading Carex marsh and sifting Alnus litter, J.M. Campbell (CNC) 1 sex undetermined; BC, Queen Charlotte Is., Louise Is., Skedans, 6. VIII.1983, 83-59, sifting beach wrack and on beach at night, J.M. Campbell (CNC) 1 male, 1 sex undetermined; BC, Queen Charlotte Is., Mamin R., 1 km SE Juskatla, 500 , 30.VII.1983, 83-46, under rock at river edge, J.M. Campbell (CNC) 1 sex undetermined; BC, Queen Charlotte Is., Skidegate, 17.VII.1983, 83-17, ex rotting seaweed, J.M. Campbell (CNC) 1 sex undetermined;

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Vancouver Is., Long Beach, 12 mi S Tofino, 23. V.1968, A. Smetana (CNC) 1 male, 1 female. COLLECTION AND HABITAT DATA. Habitat: mainly litoral habitats, found in decomposing seaweed on sandy seashores, under seaweed on rocks with Diptera larvae (Gusarov 2003d); under rock at river edge, treading Carex marsh and sifting Alnus litter, and under rock at river edge. Collecting period: V– VIII, X. Collecting method: sifting decomposing seaweed and algae; treading Carex marsh and sifting Alnus litter.

Genus Aloconota C.G. Thomson, 1858 (Figs. 20.156a–g and 20.157a–h) (Klimaszewski and Peck 1986; Gusarov 2003a; Klimaszewski et al. 2011, 2018b) Diagnosis. Body narrowly subparallel; pronotum and elytra often with complex pubescence pattern in males (wavy, V-shaped); ligula divided into two lobes at base; prementum with two medial, widely separated setae, lateral pseudopores absent; twin pores of labial palpomere I small; mesocoxal cavities narrowly separated; process of mesoventrite pointed at apex; hind tarsus with tarsomere I longer than II to about as long as II and III combined, empodial seta distinctly longer than tarsal claw; male tergite VIII with apical margin obliquely produced laterally, with two small, rounded medial teeth or tubercles as well as two sublateral teeth or projections; median lobe of aedeagus narrowly elongate and without distinct sclerites, median bridge wide but incomplete medially, flagellum thin and longer than median lobe; spermatheca very short, swollen toward base. Key to Species of Aloconota 1. Antennomeres VII–X subquadrate to slightly transverse (Fig. 20.157a); elytra at suture slightly longer than pronotum along midline (Fig. 20.157a); spermathecal capsule facing upward (Fig. 20.157h); tergite VIII with two

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minute lateral teeth, and two inconspicuous broadly triangular rounded teeth medially (Fig. 20.157d)....................................................... ...................Aloconota carmanah Klimaszewski – Antennomeres VII–X slightly elongate (more in females than in males) (Fig. 20.156a); elytra at suture as long as pronotum along midline (Fig. 20.156a); spermathecal capsule facing laterad (Fig. 20.156g); tergite VIII with two minute lateral teeth, and two distinct narrowly triangular rounded teeth medially (Fig. 20.156c)................. ........................Aloconota cambrica (Wollaston)

121. Aloconota cambrica (Wollaston) { (Fig. 20.156a–g) Homalota cambrica Wollaston, 1855. Moore and Legner 1975, Klimaszewski and Winchester 2002, Smetana 2004. Diagnosis. Body length 2.6–2.8 mm, fattened, narrowly subparallel, colour brown to dark brown, legs, and posterior part of elytra slightly paler; forebody densely punctate and pubescent, punctation slightly asperate, microsculpture meshy, particularly well visible on pronotum, sculpticells slightly convex (Fig. 20.156a); head elongate and strongly depressed medially, nearly as broad as pronotum, postocular region elongate, twice as long as maximum diameter of eye, tempora with vestigial carinae, not visible laterally when head slightly retracted into thorax or visible only at very base of tempora; antennae slender, as long as pronotum and elytra combined, basal three antennomeres strongly elongate, IV–X slightly elongate, and terminal one strongly elongate and about as long as two preceding antennomeres combined; pronotum narrowly trapezoidal in shape, flattened, slightly elongate, hypomeron visible across basal two-thirds of pronotal length in lateral view, pubescence radiating from midline of disk anteriad, laterad along broadly arched lines and obliquely posteriad; elytra flat, slightly elongate, broader than pronotum, posterior margins broadly arcuate and without angular emarginations,

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pubescence directed obliquely posteriad; abdomen subparallel, four first visible tergites impressed at base; basal metatarsomere only slightly longer than the following one. MALE. Tergite VIII with two minute lateral teeth, and two distinct narrowly triangular rounded teeth medially (Fig. 20.156c); sternite VIII rounded apically (Fig. 20.156d); tubus of median lobe of aedeagus long, slightly about straight in lateral view (Fig. 20.156b). FEMALE. Tergite VIII truncate apically (Fig. 20.156f); sternite VIII broadly rounded apically (Fig. 20.157f); spermatheca S-shaped, capsule compressed facing laterad (Fig. 20.156g). DISTRIBUTION. Origin: Palearctic, adventive in Canada. CANADA: BC. USA: CA. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002). Habitat: in BC adults were captured in Sitka spruce forest of Carmanah Valley, Vancouver Island, in forest interior, tranzition zone, and clearcut. Majority of specimens were collected in clearcut zone. Collecting period: VI. Collecting method: Malaise traps. 122. Aloconota (Fig. 20.157a–h)

carmanah

Klimaszewski

Aloconota carmanah Klimaszewski, Klimaszewski and Winchester, 2002.

in

Diagnosis. Body length 2.5–2.9 mm, flattened, narrowly subparallel, colour light brown to dark brown, usually appendages and elytra paler (Fig. 20.157a); forebody densely punctate and pubescent with meshy microsculpture particularly well visible on pronotum, sculpticells slightly convex; head elongate and slightly depressed medially, nearly as broad as pronotum, postocular region elongate, about 1.5 longer than maximum diameter of eye, tempora with vestigial carinae, not visible laterally when head slightly retracted into thorax or visible only at very base of tempora; antennae slender, as long as pronotum and elytra combined, basal three antennomeres

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strongly elongate, IV–VI slightly elongate, VII–X slightly transverse, progressing apically, and terminal one strongly elongate and almost as long as two preceding antennomeres combined; pronotum flattened, slightly elongate, narrowly trapezoidal in shape, hypomeron visible for about two-thirds of the entire pronotal length in lateral view, pubescence radiating from midline of disk anteriad medioapically and laterad along broadly arched lines laterally; elytra flat, slightly elongate, broader than pronotum, posterior margin of each elytron broadly arcuate and without angular emarginations, pubescence directed obliquely posteriad; abdomen subparallel, four first visible tergites impressed at base; basal metatarsomere only slightly longer than the following one. MALE. Tergite VIII with two minute lateral teeth, and two inconspicuous broadly triangular rounded teeth medially (Fig. 20.157d); sternite VIII rounded apically (Fig. 20.157e); tubus of median lobe of aedeagus short, slightly sinuate in lateral view (Fig. 20.157b), in dorsal view as illustrated (Fig. 20.157c). FEMALE. Tergite VIII truncate apically (Fig. 20.157f); sternite VIII rounded apically (Fig. 20.157g); spermatheca S-shaped, capsule compressed facing dorsad (Fig. 20.157h). DISTRIBUTION. Origin: Palearctic, adventive in Canada. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002). Habitat: adults were captured in Sitka spruce forest of Carmanah Valley, Vancouver Island, in tranzition zone. Collecting period: VI, VII. Collecting method: Malaise traps.

Genus Amischa C.G. Thomson, 1858 (Fig. 20.158a–e) (Muona 1990; Ashe 2001; Klimaszewski et al. 2018b) Diagnosis. Body ovoid or subparallel, rather loosely articulated, length 2.5–4.8 mm in most; integument dark brown to black, often with paler

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elytra, usually with meshed microsculpture; head either broadly oval or widened posteriad so the greatest width is behind eyes, clearly constricted basally; eyes small, shorter than length of temples; pronotum with pubescence on disc forming distinct pattern, usually directed lateroposteriad from midline, but with specific patterns in different subgenera; hypomera completely visible in lateral view; mesocoxae narrowly separated; mesocoxal cavities not margined; process of mesoventrite slender; tarsal formula 4-5-5; athetine bridge located in dorsoanterior part of median lobe of aedeagus; spermatheca with capsule club-shaped, perpendicularly bent at middle, stem long, tightly coiled posteriorly. Comments. This genus contains several species in Canada. In this book, we recognise only one adventive species, A. analis (Gravenhorst), in British Columbia, but there is at least one more species that we cannot put a name on at the present time because this genus requires a major revision in North America. In Amischa, the median lobe of the aedeagus offers few diagnostic characters for species recognition and the shape of the spermatheca and tergite VIII of both sexes offer much better characters for species identification. 123. Amischa analis (Gravenhorst) { (Fig. 20.158a–e) Aleochara analis Gravenhorst, 1802. Bernhauer 1907, Moore and Legner 1975, Muona 1984, 1990, Klimaszewski et al. 2005, 2007a, 2011, 2013a, 2016d, 2018b, Majka and Klimaszewski 2008a. Diagnosis. Body narrowly subparallel, moderately convex, length 2.2–2.4 mm; colour light to dark brown with antennae, palpi, legs and often apical part of abdomen paler (Fig. 20.158a); forebody moderately glossy, pubescence fine and sparse; head abruptly constricted at base, almost as wide as pronotum; antennomere IV subquadrate, V–X slightly to distinctly

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transverse; pronotum slightly transverse and as wide as elytra; elytra at suture slightly longer than pronotum; abdomen subparallel for most of its length. MALE: not found in Canada. FEMALE. Apical margin of tergite VIII obliquely produced laterally, with broad, deep, rounded medial emargination (Fig. 20.158b); sternite VIII arcuate apically (Fig. 20.158c); spermatheca with capsule tubular, angularly bent at middle, with deep apical invagination, stem long, thin, tightly coiled posteriorly (Figs. 20.158d, e). DISTRIBUTION. Origin: Palearctic, now cosmopolitan, adventive in Canada and USA. CANADA: BC [new provincial record], LB, NB, NF, NS, ON, PE, QC, SK. USA: CA, CO, IN, MA, ME, MI, NH, NY, PA, VT, WA. We report this species from BC for the first time here based on the following: British Columbia, Lander, Imperial road, 49.07 N, 119.33W , 15.V.1961, coll. A.T. Wilkinson (UBC, LFC) 6 females; Osoyoos, WR, Black Sage Rd., 7. VI.1995/7.VII.1995, 6.V.1994/8.VI.1994, Purshia association, pitfall trap, coll. G.G.E. Scudder (UBC) 2 females; Osoyoos, IR1, Brights Winery, 6.V.1994/2.VI.1994, Purshia association, pitfall trap, coll. G.G.E. Scudder (UBC) 1 female; Osoyoos, near Haynes Ecol. Res., 1.V.1995/26.V.1995, 24. V.1999/30.VI.1999, 6.VI.2000/6.VII.2000, Artemisia–Purshia association, pitfall trap, coll. G.G.E. Scudder (UBC) 3 females; Osoyoos, Haynes Ecol. Res., 4.IV.1995/1.V.1995, 11. X.1998/11.IV.1999, 11.V.1999/24.V.1999, 9. V.2000/6.VI.2000, recovery after fire, pitfall trap, coll. G.G.E. Scudder (UBC) 4 females. COLLECTION AND HABITAT DATA. Habitat: open and forested habitats; agricultural fields, dry habitats with Artemisia–Purshia association and recovery after fire habitats (BC), pasture/meadow (commonly collected by sweeping grasses), urban fields, in grass litter in garden and lawn, coastal sand dunes amidst vegetation, river margins in flood debris and drift material, old-growth eastern white cedar swamp in moist

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sphagnum, black spruce bog in sphagnum, and red spruce forest in pitfall traps. Collecting period: IV–X. Collecting method: sifting various kinds of litter and moss, pitfall traps, sweeping grasses.

Genus Atheta C.G. Thomson, 1858 [sensu lato] (Figs. 20.159–20.201, 20.256, and 20.257) (Strand and Vik 1964; Benick and Lohse 1974; Seevers 1978; Lohse et al. 1990; Ashe 2001; Klimaszewski and Winchester 2002; Gusarov 2003a; Elven et al. 2010; Klimaszewski et al. 2018b) Diagnosis. Body ovoid or subparallel, rather loosely articulated with very flexible abdomen, length 2.5–4.8 mm in most species; colour dark brown to black, integument usually with meshed microsculpture; suborbital carina partial or complete; ligula split; pronotum with pubescence on disc forming distinct pattern, usually directed lateroposteriad from midline, but with specific patterns in different subgenera; hypomera completely visible in lateral view; mesocoxae narrowly separated; process of mesoventrite slender; tarsal formula 4-5-5; athetine bridge of median lobe of aedeagus partial (divided dorsally), or complete and located in dorso-basal part of median lobe of aedeagus; spermatheca with club-shaped capsule and long stem that is narrowly looped posteriorly in most species (there are many spermathecal forms present in Atheta). Comments. This genus is treated here as Atheta sensu lato, similar as in Klimaszewski et al. (2018b), and embraces a wide range of species with diverse morphology forming the Atheta complex. It includes several subgenera that are not sufficiently delimited to justify elevating them to generic status, and such an act would bring only more confusion at the present stage of knowledge on this group. It is, however, a temporary solution and most of the present subgenera, when properly defined, should be elevated to a

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generic status. The subgenera of Atheta are listed below in alphabetical order because the relationships between them are not clear. For species identification, dissection and examination of genitalic structures is necessary. Key to Subgenera of Atheta (Species of Atheta currently considered to be incertae sedis are not included in the key to subgenera of Atheta. Should the key not work well for the specimens at hand, the user should compare genital structures of their specimens to those species listed as incertae sedis). 1. Body strongly flattened (Fig. 20.185a); pubescence on elytra forming wavy pattern near sutural area of disc (Fig. 20.185a); male tergite VIII with two minute lateral teeth and minute medial emargination (Fig. 20.185d); spermatheca more or less evenly tubular (Fig. 20.185h) .............................. ..........Hydrosmectomorpha Klimaszewski et al. – Body more or less convex; pubescence on elytra forming different patterns; male tergite VIII and spermathecal differently shaped ..........2 2. Body usually subparallel, highly glossy (Figs. 20.160a and 20.161a); length 3.5–4.4 mm; forebody with microsculpture fine or not evident; elytra often with granulose texture; antennomeres VI–X slightly transverse, quadrate or elongate (Figs. 20.160a and 20.161a); spermatheca with stem irregularly twisted and bearing small tuberculate projections (Figs. 20.160h and 20.161i) ................. Atheta s. str. C.G. Thomson – Combination of characters not as above .......3 3. Apical part of abdomen with long, arcuate and robust macrosetae, particularly on ventral side (Fig. 20.188a); genitalia as illustrated (Fig. 20.188b, c, h)............................................... ............................. Metadimetrota Klimaszewski – Apical part of abdomen with setae not as above; genitalia shaped differently ....................4 4. Pronotum rounded, highly convex and glossy, all pubescence on disc directed straight or obliquely posteriad (Fig. 20.159a); pronotal hypomera narrowly visible in lateral view; male

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tergite VIII truncate apically, without teeth on apical margin [BC species] (Fig. 20.159d); median lobe of aedeagus in dorsal view with small bulbus and long, narrow tubus (Fig. 20.159c), ventral margin of tubus long, arcuate in lateral view, internal sac as illustrated (Fig. 20.159b)................ Alaobia C.G. Thomson – Combination of characters not as above .......4 5. Body narrowly subparallel, length 2.5–2.7 mm (Fig. 20.162a); pronotum rectangular in shape; abdomen broadening apically; infraorbital carinae incomplete; antennomeres VII–X distinctly transverse and incrassate (Fig. 20.162a); male tergite VIII with entire apical margin subrectangularly produced, rounded laterally, shallowly sinuate along margin, without teeth (Fig. 20.162e); tubus of median lobe of aedeagus in lateral view with ventral margin irregularly, shallowly sinuate, apex narrowly triangular (Figs. 20.162b, c)..................... ........................................ Bessobia C. G. Thomson – Combination of characters not as above .......6 6. Body length 3.0–3.5 mm; apical and lateral abdominal macrosetae very strong; antennomeres VII–X elongate (Fig. 20.163a); male tergite VIII shallowly emarginate apically (Fig. 20.163d); median lobe of aedeagus in dorsal view gradually tapering from near base of bulbus to apex of tubus (Fig. 20.163c), in lateral view with crista apicalis trapezoidal, apex of tubus broad, rounded (Fig. 20.163b), spermatheca with stem elbowed at base of capsule and near middle, with single curl at base (Fig. 20.163h) ................................... .................................Chaetida Mulsant and Rey – Combination of characters not as above .......7 7. Body length 2.2–4.0 mm (usually 2.8–3.0 mm); at least pronotum with coarse asperate punctation (Figs. 20.166a–20.184a); lateral margins of pronotum, elytra, and middle and hind tibiae with distinct bristles; elytra of the same colour as pronotum or paler but not strongly contrasting with rest of the body; antennomeres IV–X subquadrate, slightly transverse, or rarely slightly elongate; apical margin of male tergite VIII obliquely produced laterally, to broadly truncate apex which is usually bounded by teeth or

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angulate projections (Figs. 20.166d, 20.167d, 20.169d, 20.170d, and 20.171d); median lobe of aedeagus and spermatheca diverse in form (Figs. 20.166b, c, h, 20.167b, c, h, 20.168b, c, h, and 20.169b, c, h) ............................................ ...............................Dimetrota Mulsant and Rey – Combination of characters not as above .......8 8. Large species, length 3.0–4.0 mm; body broad, subparallel, broadest at elytra; antennomeres VII–X subquadrate to moderately transverse; abdomen broad, arcuate laterally (Figs. 20.186a, and 20.187a); tubus of the median lobe of aedeagus with two lateral narrow and diverging projections in dorsal view (Figs. 20.186c, and 20.187c); spermatheca with small spherical capsule and stem thin, very long, sinuate, and narrowly looped posteriorly (Fig. 20.186h).............................. Lamiota Casey – Body size variable; smaller in most, length 2.8–3.7 mm; other character not as above or in different combination .........................................9 9. Elytra yellowish to yellowish-red, strongly contrasting with dark brown pronotum, frequently darker baso-medially (Fig. 20.196a); antennomeres V–X distinctly transverse (Fig. 20.196a); apical margin of male tergite VIII truncate, crenulate, with sublateral teeth (Fig. 20.196d); genitalia as illustrated (Figs. 20.196b, c, h) ...................Pseudota Casey – Combination of characters not as above .....10 10. Head, pronotum and abdomen dark brown, elytra yellowish-brown (Fig. 20.198a); pronotum without distinct asperate punctation; antennomeres V–X moderately transverse (Fig. 20.198a); apical margin of male tergite VIII abruptly produced in middle 2/5, apex sinuately truncate, angulate laterally (Fig. 20.198e); median lobe of aedeagus as illustrated (Fig. 20.198b–d); spermatheca with basal part of capsule almost as broad as small spherical apex, stem elbowed at base of capsule and near middle, with single tight curl at base (Fig. 20.198i, j) ........Tetropla Mulsant and Rey – Head, pronotum and abdomen uniformly dark brown to black, or pronotum orange and elytra

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Tribe Athetini Casey, 1910

yellowish (A. pennsylvanica); pronotum often with distinct asperate punctation; other characters in different combination...................................11 11. Forebody finely to moderately punctate (Figs. 20.189a–20.194a); male tergite VIII truncate or arcuate apically and without teeth (Figs. 20.189a–20.194a); spermatheca with entire capsule broad, club-shaped, broadest at rounded apex, stem angulately bent at base of capsule (Figs. 20.189g, 20.190h, 20.191h, 20.192h, and 20.193h), median lobe of aedeagus as illustrated (Figs. 20.189b, 20.190b, c, 20.191b, c, 20.192b, c, 20.193b, c, and 20.194b, c).............................. .............................. Microdota Mulsant and Rey – Forebody coarsely and densely punctate (Figs. 20.164a and 20.165a); male tergite VIII densely setose and with apical teeth (Figs. 20.164d and 20.165d); spermatheca with capsule not entirely broad, club shaped (Figs. 20.164h and 20.165h), median lobe of aedeagus as illustrated (Figs. 20.164b, c and 20.165b, c) ........... Datomicra Mulsant and Rey

(Subgenus Alaobia C.G. Thomson, 1858) (Fig. 20.159a–h) (Gusarov 2003a) Diagnosis. Body narrowly ovoid, length 2.5–3.0 mm; integument highly glossy, unevenly coloured; antennomere IV subquadrate, VII–X distinctly transverse; cervical carina not diverged; ligula fusiform; pronotum rounded, with all pubescence on disc directed straight or obliquely posteriad; pronotal hypomera narrowly visible in lateral view; male tergite VIII truncate apically; median lobe of aedeagus in dorsal view with small bulbus and very elongate narrow tubus; in lateral view ventral margin of tubus long, arcuate, internal sac without distinct sclerites; spermatheca with moderately large club-shaped capsule with narrow and moderately long apical invagination, stem angulately bent at base of capsule, narrowly looped posteriorly and irregularly twisted at base. Comments. This subgenus is not monophyletic (Elven et al. 2012). Generally, all included

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Tribe Athetini Casey, 1910

species have the same superficial shape of the spermatheca. The two species included here are rather dissimilar, but in the absence of a more detailed revision we maintain the current classification. 124. Atheta (Alaobia) ventricosa Bernhauer (Figs. 20.159a–h) Atheta (Atheta) ventricosa Bernhauer, 1907. Gusarov 2003a, Klimaszewski et al. 2005, 2007b, 2008b, 2011, 2015a, 2018b, Majka and Klimaszewski 2008b, 2010, Stefani et al. 2016. Diagnosis. Body narrowly ovoid, moderately convex, length 2.5–3.0 mm; colour brown to dark brown, with antennomeres I–III, palps, legs, often pronotum and part of elytra, and basal and apical part of abdomen paler, yellowish-brown (Fig. 20.159a); integument highly glossy; pubescence fine and sparse, denser on elytra; head slightly narrower than pronotum; antennae moderately thick, antennomere IV subquadrate, VII–X distinctly transverse, each about 1.2–1.4 times as wide as long; pronotum round, highly convex, moderately transverse, widest at middle, distinctly narrower than elytra, all pubescence on disc directed latero-posteriad; elytra at suture about as long as pronotum; abdomen mostly subparallel. MALE. Tergite VIII with apical margin obliquely produced laterally, broadly shallowly emarginate medially (Fig. 20.159d); sternite VIII parabolically rounded from base to apex (Fig. 20.159e); median lobe of aedeagus in dorsal view with bulbus small, roughly rhomboid, tubus long, subparallel and gradually tapering apicad (Fig. 20.159c), in lateral view tubus long, ventral margin curved obliquely ventrad basally, almost straight to middle, then thin and slightly arcuate to narrowly rounded apex (Fig. 20.159b). FEMALE. Tergite VIII with apical margin obliquely produced laterally, truncate in about apical 1/3 (Fig. 20.159f); sternite VIII broadly parabolically rounded from base to apex (Fig. 20.159g); spermatheca with capsule large, club-shaped with apex broadly cone-shaped, stem arcuate, looped and irregularly twisted posteriorly (Fig. 20.159h).

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DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK, YT. USA: AK, CA, DC, IN, MA, NC, NH, NJ, NY, PA, VT, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, found in various forest types usually in decaying fungi (often common) and leaf litter; mature mixed forest in polypore fungi, mixed forest with cedar in litter at margin of vernal pond, old spruce forest in Tricholoma sp. in various stages of decay, spruce and balsam fir forests in decaying lobster mushrooms, mature red spruce and eastern white cedar forests in decaying mushrooms and leaf litter, old-growth eastern white cedar swamp in moss and litter at base of tree (6 April, probably overwintering site), balsam fir stand in moldy conifer duff, and in Carex marsh in litter at base of tree. In NF, in unbaited and carrion-baited pitfall traps and flight intercept traps in various coniferous, mixed wood, and riparian forests. Elsewhere, in conifer forests and organic litter in mixed forests. Collecting period: IV–X. Collecting method: sifting fungi and various kinds of forest litter, flight intercept traps, unbaited and carrion-baited pitfall traps. (Subgenus Atheta C.G. Thomson, 1858) (Figs. 20.160a–h, 20.161a–i) (Benick and Lohse 1974; Muona 1979; Klimaszewski and Winchester 2002; Klimaszewski et al. 2018b) Diagnosis. Body length 3.5–4.4 mm, narrowly subparallel, elongate, usually highly glossy; pronotum with pubescence on disc directed laterad from midline; pronotal hypomera fully visible in lateral view; antennomere IV slightly elongate to subquadrate, IV–X slightly transverse, quadrate or slightly elongate; spermatheca with stem thin, irregularly twisted and bearing small tuberculate projections. Comments. The shape of spermatheca constitutes an apomorphy of this group. This subgenus contains mostly northern species.

364

Key to Species of Subgenus Atheta 1. Antennomeres VII–X elongate (Fig. 20.160a); apical margin of male tergite VIII with fine teeth (crenulate) (Fig. 20.160d); male sternite VIII notched apically (Fig. 20.160e) ........................... ...... Atheta (Atheta) graminicola (Gravenhorst) – Antennomeres VII–X transverse (Fig. 20.161a); apical margin of male tergite VIII entire, without teeth (crenulate) (Fig. 20.161e); male sternite VIII pointed apically (Fig. 20.161f) ............................... ..................... Atheta (Atheta) ringi Klimaszewski

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Tribe Athetini Casey, 1910

(Fig. 20.160f); sternite VIII broadly rounded apically (Fig. 20.160g); spermatheca with capsule U-shaped tubular, with apical part broader pitcher-shaped, stem irregularly twisted and coiled with small granulate projections (Fig. 20.160h). DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, LB, MB, NB, NF, NT, ON, QC, SK, YT. USA: AK, OR.

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: mostly associated with various wetland habitats in NB; 125. Atheta (Atheta) graminicola (Gravenhorst) Carex marsh in emergent vegetation (treading, (Fig. 20.160a–h) very common), splashing exposed bedrock next to waterfalls, in leaf litter and moss under alders Aleochara graminicola Gravenhorst, 1806. next to small brook in Carex marsh, among Lohse and Smetana 1985, Lohse et al. 1990, cobblestones, gravel, and sand on gravel bar Gusarov 2003a, Webster et al. 2009, near outflow of brook into a river, in hardwood Klimaszewski et al. 2011, 2015a, 2018b. forest in moist leaves on vernal pond margin, upper margin of sea beach in drift material Diagnosis. Body narrowly ovoid, moderately (mostly grass litter), in leaf litter at base of tree convex, length 3.5–4.4 mm; colour dark brown, in red oak and red maple forest, in leaf litter in often with antennae, palps, legs, pronotum/elytra crotch of silver maple in silver maple swamp in (or elytra only) and apical part of abdomen light early April (probably overwintering site); two reddish-brown (Fig. 20.160a); integument glossy; individuals collected with aerial net in late afterpubescence fine and sparse, denser on elytra; head noon (15:00–18:00 h) when snow still present in slightly narrower than pronotum; antennae shaded areas of woods. In QC, found in leaf litter robust, with all antennomeres elongate, each next to stream and in Salix litter at two localities about 1.5 times as long as wide; pronotum in YT. In NL, collected using a flight intercept rounded, moderately transverse, widest at middle, trap in a mixed forest. Collecting period: IV– narrower than elytra; elytra at suture at least as VIII. Collecting method: sifting litter, treading long as pronotum; abdomen arcuately narrowed marsh vegetation, aerial net, flight intercept traps, to apex posteriorly. MALE. Tergite VIII with pitfall traps. apical margin obliquely produced laterally, broadly truncate and coarsely crenulate apically 126. Atheta (Atheta) ringi Klimaszewski between two rounded angulate teeth (Fig. 20.161a–i) (Fig. 20.160d); sternite VIII parabolically Atheta ringi Klimaszewski, in Klimaszewski rounded from base to apex except narrowly emarginate at apex (Fig. 20.160e); median lobe of and Winchester, 2002. aedeagus in dorsal view with bulbus broad and short, abruptly constricted at base of triangular Diagnosis. Body length 3.2–4.0 mm, modertubus (Fig. 20.160c); in lateral view tubus deeply ately convex, narrowly subparallel, dark brown, constricted at base, ventral margin highly sinuate sometimes head and basal part of abdomen nearly to curved, moderately narrowly rounded apex black, elytra, legs and basal antennal segment (Fig. 20.160b). FEMALE. Tergite VIII with api- rust-brown (Fig. 20.161a); forebody moderately cal margin broadly truncate, sinuate laterally sparsely punctate and pubescent, pubescence

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Tribe Athetini Casey, 1910

long, punctation slightly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head slightly elongate, round, slightly depressed medially, distinctly narrower than pronotum, postocular region as long as or slightly shorter than the maximum diameter of eye, tempora carinate, carinae strong basally and diffused apically; antenna slender and long, slightly widening apically, approximately as long as pronotum and elytra combined, antennomeres I–III strongly elongate, IV slightly elongate, V quadrate, VI–X slightly transverse (up to about one third broader than long), terminal one elongate and nearly as long as two preceding antennomeres combined, basal antennomere IV glossy and remaining matte; pronotum impressed mesobasally, slightly transverse, approximately rectangular in shape, hypomeron visible for almost entire pronotal length in lateral view, pubescence directed anteriad at one third of apical midline, obliquely laterad across arcuate lines medially and obliquely posteriad elsewhere; elytra moderately convex, slightly transverse, distinctly broader than pronotum, posterior margins broadly arcuate and with small but distinct shallow lateral emarginations, pubescence directed obliquely posteriad, forming wavy pattern basally; abdomen subparallel, three first visible tergites strongly impressed at base; basal metatarsomere about one third longer than the following one. MALE. Tergite eight transverse and broadly rounded apically (Fig. 20.161e); sternite eight slightly transverse, rounded apically and with tip truncate or shallowly emarginated (Fig. 20.161f); median lobe of aedeagus with broadly oval large bulbus and short triangular tubus in dorsal view (Fig. 20.161d), in lateral view tubus straight ventrally and feebly pointing at apex (Fig. 20.161b, c). FEMALE. Tergite VIII transverse and truncate apically (Fig. 20.161g); sternite VIII transverse and rounded apically (Fig. 20.161h); spermatheca with pitcher-shaped and sinuate capsule and irregularly twisted posterior duct (Fig. 20.161i).

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DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski and Winchester 2002). Habitat: adults were captured in forest litter in Sitka spruce forest of Carmanah Valley, Vancouver Island, in forest interior, transition zone, and clearcut, with most captured in transition zone. Collecting period: VI–VIII. Collecting method: pitfall traps. (Subgenus Bessobia C.G. Thomson, 1858) (Figs. 20.162a–i) (Lohse et al. 1990; Smetana 2004; Klimaszewski et al. 2018b) Diagnosis. Body narrowly subparallel, length 2.5–2.7 mm; integument slightly glossy, unevenly coloured; antennomere IV slightly transverse, VII–X distinctly transverse, about twice as wide as long; pronotum rectangular with all pubescence on disc directed obliquely laterad from midline; pronotal hypomera fully visible in lateral view; male tergite VIII with entire apical margin subrectangularly produced, rounded laterally, broadly sinuate along margin, without teeth; median lobe of aedeagus in dorsal view with large oval bulbus and very short, rightangled triangular tubus, in lateral view ventral margin of tubus irregularly shallowly sinuate; spermatheca capsule small, club-shaped, with small apical invagination, sinuate stem narrowly looped posteriorly. 127. Atheta (Bessobia) cryptica (Lohse) (Fig. 20.162a–i) Bessobia cryptica Lohse, in Lohse et al. 1990. Klimaszewski et al. 2008b, 2011, 2015a, 2018b. Diagnosis. Body elongate, narrowly subparallel, moderately convex, length 2.5–2.7 mm; colour brown, often with reddish tinge, basal three antennomeres, palps, legs, pronotum, and elytra light brown to reddish-brown, moderately glossy

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(Fig. 20.161a); pubescence fine and dense, denser on elytra; head slightly narrower than pronotum, impressed between eyes; antennae moderately thick and incrassate, antennomere IV slightly transverse, V–X distinctly transverse, each at least 1.5 times as wide as long; pronotum rectangular in shape, transverse, widest at middle, slightly narrower than elytra; elytra at suture about as long as pronotum along midline; abdomen mostly subparallel and widening apicad to about 3/4 of its length, then moderately narrowing apically. MALE. Tergite VIII with entire apical margin subrectangularly produced, lateral angles rounded and posterior margin shallowly sinuate (Fig. 20.161e); sternite VIII rounded apically (Fig. 20.161f); median lobe of aedeagus with large oval bulbus and very short triangular tubus in dorsal view (Fig. 20.161d), in lateral view ventral margin of tubus bent obtusely ventrad at base, irregularly, shallowly sinuate to narrowly triangular apex (Fig. 20.161b, c). FEMALE. Tergite VIII arcuate apically (Fig. 20.161g); sternite VIII broadly rounded apically (Fig. 20.161h); spermatheca with small club-shaped capsule, with small apical invagination, and sinuate stem narrowly looped posteriorly (Fig. 20.161i). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NF, QC, YT. USA: not recorded. New locality data: British Columbia, 160 km W of Watson Lake, 27.VI.2015, 59.9056131.4924, 872 m, Firekill, decidous, sifting, coll. B. Godin and D. Horwood (BGC, LFC) 2 males, 3 females; BC, Summit Lake, 58.6616124.5215, 1238 m, 10.VI.2013, Willow/aspen litter, sifting, coll. B. Godin and D. Horwood (BGC, RBCM) 2 males, 2 females. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: collected using pitfall traps in fir forests (NF) and sifted from willow (Salix sp.) leaf litter (YT). In BC, specimens were taken from litter in firekilled decidous forest. Collecting period: V–VII. Collecting method: sifting forest litter, pitfall traps.

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Tribe Athetini Casey, 1910

(Subgenus Chaetida Mulsant and Rey, 1873b) (Fig. 20.163a–h) (Smetana 2004; Klimaszewski et al. 2018b) Diagnosis. Distinct for its Oxypoda-like body form, abdomen tapering apically and with strong lateral macrosetae; macrosetae also present on lateral margins of pronotum and hind femora; antennomeres V–X distinctly elongate; male and female tergite VIII truncate apically and with broad, shallow median emargination; median lobe of aedeagus simple, tubus slightly arcuate ventrally, apex broadly rounded; spermatheca with long and mostly straight stem with single curl at base. 128. Atheta (Chaetida) longicornis (Gravenhorst) { (Fig. 20.163a–h) Aleochara chaetida Gravenhorst, 1802. Muona 1984, Smetana 2004, Klimaszewski et al. 2007a, 2011, 2015a, 2018b, Webster et al. 2009, Majka and Klimaszewski 2008b, Michaud et al. 2010. Diagnosis. Body narrowly ovoid (Oxypodalike), length 3.0–3.6 mm, colour brown to black, with elytra and often apex of abdomen yellowishbrown or reddish-brown, integument moderately glossy, forebody with coarse asperate punctation (Fig. 20.163a); pubescence fine and sparse, denser on elytra; head distinctly narrower than pronotum, slightly transverse; antennae robust, antennomeres IV–V subquadrate and VI–X elongate; pronotum rectangular, transverse, slightly narrower than elytra; elytra at suture as long as pronotum; abdomen strongly tapering apically with strong macrosetae. MALE. Apical margin of tergite VIII sinuate laterally, broadly shallowly emarginate apically (Fig. 20.163d); sternite VIII elongate, semicircularly rounded apically (Fig. 20.163e); median lobe of aedeagus in dorsal view narrowly ovoid, gradually tapering to apex (Fig. 20.163c); in lateral view tubus short, slightly curved ventrad basally, ventral margin slightly arcuate to broadly rounded apex (Fig. 20.163b). FEMALE. Tergite VIII similar to that of male but

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Tribe Athetini Casey, 1910

apical margin rounded laterally (Fig. 20.163f); sternite VIII rounded apically with minute median emargination (Fig. 20.163g); spermatheca with capsule club-shaped, stem long, elbowed at base of capsule and near middle, with single curl at base (Fig. 20.163h). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: BC, NB, NF, NS, QC, SK. USA: CA, MN, OR. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: open habitats where it occurs in cow and horse dung, carrion, compost, rotting mushrooms, and other rotting organic substrates. Some flying individuals collected with aerial net in late afternoon (15:00–18:00 h) early in spring in mixed forest opening. Collecting period: IV–X. Collecting method: sifting organic litter, pitfall traps, collected with butterfly net during late afternoon. (Subgenus Datomicra Mulsant and Rey, 1873b) (Figs. 20.164a–h and 20.165a–h) (Smetana 2004; Klimaszewski et al. 2011, 2018b; Lee and Ahn 2017) Diagnosis. Body length 0.9–2.5 mm; integument of pronotum distinctly transverse, usually densely and coarsely punctate, matte with finely reticulate microsculpture, setae along midline directed anteriad; antennomeres VII–X distinctly transverse. This subgenus generally is similar to Dimetrota but the body size is smaller. Comments. Species of this subgenus were recently reviewed for South Korea, including some species occurring in Canada (Lee and Ahn 2017). Key to Species of Subgenus Datomicra 1. Pronotal punctures very dense, distance between punctures equal to diameter of a puncture, integument slightly glossy (Fig. 20.164a); four teeth of male tergite VIII short, crenulationlike (Fig. 20.164d); apex of median lobe of

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aedeagus narrow in lateral view (Fig. 20.164b); spermatheca with stem thin and long (Fig. 20.164h)....................................................... ................ Atheta (Datomicra) celata (Erichson) – Pronotal punctures moderately dense, distance between punctures greater than diameter of a puncture, integument strongly glossy (Fig. 20.165a); apical margin of male tergite VIII with two large lateral and two smaller median teeth (Fig. 20.165d); apex of median lobe of aedeagus broad in lateral view (Fig. 20.165b); spermatheca with stem sinuate thick and short (Fig. 20.165h).............................. ... Atheta (Datomicra) dadopora C.G. Thomson

129. Atheta (Datomicra) celata (Erichson)  (Fig. 20.164a–h) Homalota celata Erichson, 1837. Bernhauer 1907, Casey 1910 [as Datomicra wrangler], 1911 [as Pseudota nanulina], Benick and Lohse 1974, Majka et al. 2006, Majka and Klimaszewski 2008a, Klimaszewski et al. 2011, 2015a, 2018b, Lee and Ahn 2017. Datomicra wrangleri Casey, 1910. New synonymy. Lectotype (male): USA, Alaska, (Fort Wrangel in orig. description), Wick. [ham]; Casey determ. wrangleri-9; Casey bequest 1925; Lectotypus Datomicra wrangleri Casey des. V.I. Gusarov 2000 (USNM). Present Lectotype designation. Paralectotype: USA, Alaska, Wick. [ham]; Casey determ. wrangleri-2; Casey bequest 1925 (USNM) 1 female. Examined. Pseudota nanulina Casey, 1911. New synonymy. Lectotype (male): Canada, Metlakatla, B. Col., Keen; Type USNM 39152; Casey bequest 1925; Lectotypus Pseudota nanulina Casey, V.I. Gusarov des. 2000 (USNM). Present Lectotype designation. Diagnosis. Body small, subparallel, length 1.8–2.2 mm; colour uniformly dark brown, with legs and posterior section of elytra slightly paler

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rusty-brown, integument slightly glossy, forebody with coarse and dense asperate punctation and meshed microsculpture (Fig. 20.164a); head slightly narrower than pronotum, slightly transverse; antennae slender, antennomere IV subquadrate and V–X distinctly transverse; pronotum distinctly transverse, about as wide as elytra; elytra at suture about as long as pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically, and with two minute lateral and two minute median teeth on apical margin (Fig. 20.164d); sternite VIII rounded apically (Fig. 20.164e); median lobe of aedeagus with enlarged bulbus and short arcuate ventrally tubus in lateral view (Fig. 20.164b), in dorsal view tubus broadly triangular (Fig. 20.164c). FEMALE: tergite VIII truncate apically and slightly concave medially (Fig. 20.164f); sternite VIII arcuate apically (Fig. 20.164g); spermatheca with tubular L-shaped capsule bearing very long apical invagination, stem straight and half-looped posteriorly (Fig. 20.164h). DISTRIBUTION. Origin: Previously reported as adventive in North America, but it is probably Holarctic, recorded from Europe, across northern Russia, and North and South Korea. CANADA: BC, NB, NF, NS, SK. USA: AK, MA, WV. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: various forest types in decaying organic materials, also associated with mammal and bird nests; old jack pine forest in well rotted mushroom, red spruce and yellow birch forest in rotting mushrooms, mixed forest in decaying chanterelle mushrooms, near mixed forest in decaying corncobs (moldy) and cornhusk. One individual captured in carrionbaited pitfall trap, others on sandy beach of a lake, and in wet spruce litter. Specimens also found in nests of boreal and saw-whet owls, barred owls, and nests of mammals. Collecting period: V–X. Collecting method: sifting organic litter and mushrooms, carrion-baited pitfall traps. 130. Atheta (Datomicra) dadopora Thomson  (Fig. 20.165a–h)

C.G.

Tribe Athetini Casey, 1910

Atheta dadopora Thomson, 1867. Casey 1910 [as Datomicra decolourata, D. inopia, D. schematica and D. stilla], Gusarov 2003a, Klimaszewski et al. 2005, 2007b, 2008a, 2011, 2015a, 2018b, Majka and Klimaszewski 2008a, b, c, Majka and Sikes 2009, Lee and Ahn 2017. Diagnosis. Body length 1.8–2.2 mm, small and narrow, subparallel; colour uniformly dark brown to black, with tibiae and tarsi rusty-brown, integument slightly glossy, forebody with coarse and dense asperate punctation and meshed microsculpture (Fig. 20.165a); head slightly narrower than pronotum, slightly transverse; antennae slender, antennomere IV subquadrate, and V–X slightly transverse; pronotum distinctly transverse, about as wide as elytra; elytra at suture about as long as pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically, with two lateral sharply pointed and two median rounded teeth (Fig. 20.165d); sternite VIII slightly produced apically (Fig. 20.165e); median lobe of aedeagus with large oval bulbus and short and broad tubus rounded apically in dorsal view (Fig. 20.165c), tubus straight and short with broad triangular apex in lateral view (Fig. 20.165b). FEMALE. Tergite VIII slightly emarginated apically (Fig. 20.165f); sternite VIII rounded apically with minute median emargination (Fig. 20.165g); spermatheca with capsule bearing apical projection, stem short and sinuate (Fig. 20.165h). Males differ from those of other similar species in the shape of the two rounded median teeth of tergite VIII. DISTRIBUTION. Origin: Holarctic, recorded from Europe, China, India, Japan and South Korea. CANADA: AB, BC, LB, NB, NF, NS, ON, PE, QC, SK, YT. USA: AK, NY, PA, RI. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in various mixed, coniferous, and hardwood forest types, usually in decaying materials such as decaying fungi and carrion; from Lindgren funnel trap in

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Tribe Athetini Casey, 1910

red spruce forest with red maple and balsam fir, unbaited pitfall traps in red spruce forest, spruce forest in decaying fleshy fungi, mixed forests on Pleurotus sp. on Populus log, red oak and red maple forest in gilled mushrooms, alder swamp on remains of dead deer, hardwood forests in gilled fungi, and rich Appalachian hardwood forest in fleshy fungi in various stages of decay. Also in compost, poplar/spruce litter, carrion, bison and caribou dung, and under the bark of decaying spruce logs. Collecting period: V–IX. Collecting method: sifting various kinds of organic materials, flight intercept traps, Lindgren funnel traps, unbaited and carrion-baited pitfall traps. (Subgenus Dimetrota Mulsant and Rey, 1873a) (Figs. 20.166–20.184) (Benick and Lohse 1974; Seevers 1978; Lohse et al. 1990; Klimaszewski et al. 2011, 2018b) Diagnosis. Forebody with distinct meshed or irregularly hexagonal microsculpture, punctation asperate; head slightly narrower than pronotum; eyes of moderate size, subequal in length to genae in dorsal view, infraorbital carina present; glossae divided into two lobes in apical half; pronotum transverse with hypomeron partially or fully visible in lateral view, pronotum with pubescence on disc directed mostly anteriad at midline, laterad or obliquely posteriad elsewhere, lateral bristles thick and long; elytra at suture subequal in length to length of pronotum along midline, hind margins without sinuation, pubescence on disc directed obliquely posteriad, sometimes forming wavy patterns; legs moderately long, basal article of hind tarsus about subequal in length to second one; abdomen sparsely punctate and glossy. Comments. Canadian species are arranged into species groups, and species within groups are arranged in alphabetical order. Only a key to species groups is provided for this subgenus because it contains clusters of morphologically similar species within species groups and positive identification requires examination of genitalic characters. The reader should first identify the taxon to species groups and then examine and compare genitalic characters to those for all

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known species in the group for specific identification. In case species do not match any of listed species in this publication user should also consult species included in eastern Canada (Klimaszewski et al. 2018b). Key to Species Groups of Subgenus Dimetrota (Modified from Klimaszewski et al. 2018b) [The morphology of the antennae provides good characters to classify species into species groups, however there are some exceptions to this rule. These exceptions are indicated in the key as exceptions, i.e., having characters different from those given in the particular couplet]. 1. Pubescence of forebody appearing somewhat silky (Fig. 20.166a); median lobe of aedeagus in dorsal view with tubus short, broadly triangular (Fig. 20.166c), in lateral view tubus short and straight ventrally (Fig. 20.166b); internal sac with thin and inconspicuous structures in bulbus (Figs. 20.166b, c); male tergite VIII with apical margin shallowly emarginate, with very small lateral teeth (Fig. 20.166d); spermatheca with tubular capsule, stem highly sinuate and narrowly looped posteriorly (Fig. 20.166h) ........................ ..............................................Alesi species group – Pubescence of forebody not silky, other characters not as above ......................................2 2. Antennomeres VIIII–X subquadrate to transverse (Campbelli: Fig. 20.167a; Columbica: Figs. 20.168a, 20.169a; Crassicornis: Figs. 20.170a–20.173a; Fanatica: Figs. 20.175a, 20.176a; Hampshirensis: Fig. 20.177a; Picipennis: Fig. 20.178a; Prudhoensis: Figs. 20.179a–20.184a)......................................3 – Antennomeres VIII–X slightly elongate (Fig. 20.174a); median lobe of aedeagus in dorsal view with bulbus moderate-sized, tubus long, subparallel, triangular apically (Fig. 20.174c), in lateral view tubus straight and produced ventrally at apex (Fig. 20.174b); male tergite VIII with apical margin approximately straight or slightly angularly emarginate with small lateral teeth (Fig. 20.174d); spermatheca with capsule long, tubular, with tubular appendix and small apical

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invagination, stem sinuate (Fig. 20.174h)............ ........................................Districta species group 3. Body small, length 2.0–2.6 mm, punctation of forebody dense and aspirate, distance between punctures shorter that two diameter of a puncture (Fig. 20.177a); microsculpture of forebody weak; male tergite VIII with two long and thin spineliked lateral teeth (Fig. 20.177d); internal sac of median lobe of aedeagus with two narrowly elongate sclerites in bulbus (Fig. 20.177b, c); spermatheca with capsule tubular, strongly curved medially and narrowed apically, apex pointed ventrad, apical invagination absent, stem sinuate and looped posteriorly (Fig. 20.177h) ................. ............................. Hampshirensis species group – Body medium-sized to larger, length over 2.6 mm in most; punctation of forebody less dense; other characters not as above..................4 4. Body strongly glossy; punctation of forebody dense and asperate; median lobe of aedeagus in dorsal view with bulbus oval in outline and streamlined with base of tubus, tubus short and triangular (Figs. 20.175a and 20.176a); in lateral view tubus broad, slightly to strongly sinuate ventrally and apex broadly rounded not produced ventrad (Figs. 20.175b and 20.176b); male tergite VIII with apical margin truncate, with small lateral teeth, apical margin slightly crenulate (Figs. 20.175d and 20.176d); male sternite VIII with long macrosetae (Figs. 20.175e and 20.176e); spermatheca with capsule spherical or broadly tubular rounded apically, stem sinuate and looped posteriorly (Figs. 20.175h and 20.176h) ........................ Fanatica species group – Body moderately glossy; genital characters not as above..............................................................5 5. Median lobe of aedeagus with large oval bulbus and short and triangular tubus in dorsal view (Figs. 20.167c, 20.168c, 20.169c, 20.170c, 20.171c, 20.172c and 20.173c), in lateral view apical part of tubus strongly produced ventrad (Figs. 20.167b, 20.168b, 20.169b, 20.170b, 20.171b, 20.172b and 20.173b) .........................6 – Median lobe of aedeagus not as above .........8

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Tribe Athetini Casey, 1910

6. Median lobe of aedeagus in lateral view with apical part of tubus moderately long and apical part strongly produced ventrally (Figs. 20.167b, 20.168b and 20.169b); male tergite VIII with apical margin straight or shallowly emarginate, teeth variable in size and shape (Figs. 20.167d, 20.168d and 20.169d) ........................................ 7 – Median lobe of aedeagus in lateral view with tubus almost straight in basal half and moderately produced ventrally at apex, apical part of tubus short and bent (Figs. 20.170b, 20.171b, 20.172b and 20.173b); internal sac with two very elongate sclerites in bulbus and additional complex structures in tubus (Figs. 20.170b, c, 20.171b, c, 20.172b, c and 20.173b, c) .................................. .............Crassicornis (=Modesta) species group 7. Antennomeres VII–X strongly transverse (Fig. 20.167a); integument of forebody with coarse punctation and distinct microsculpture; apical part of median lobe of aedeagus narrow in lateral view (Fig. 20.167b); spermatheca with capsule moderately broad, club-shaped, tubular, with apical invagination narrow, deep, located off-centre; stem moderately wide and long, posterior loop narrow (Fig. 20.167h) ........................ ..................................... Campbelli species group – Antennomeres VII–X slightly transverse (Figs. 20.168a and 20.169a); integument of forebody with fine punctation and weak microsculpture; apical part of median lobe of aedeagus moderately broad in dorsal view (Figs. 20.168b and 20.169b); spermatheca with capsule narrow, club-shaped, tubular, with apical invagination small; stem very thin and extremely long, posterior loop narrow (Figs. 20.168h and 20.169h) ..................... Columbica species group 8. Tubus of median lobe of aedeagus in dorsal view gradually or triangularly narrowing from apex of bulbus (Figs. 20.179c, 20.189c, 20.181c, 20.182c, 20.183c and 20.184c), in lateral view with ventral margin distinctly curved ventrad only in apical half (Figs. 20.179b, 20.189b, 20.181b, 20.182b, 20.183b and 20.184b); male tergite VIII with apical margin straight to obtusely emarginate, lateral teeth not extending beyond margin (Figs. 20.179, 20.189,

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Tribe Athetini Casey, 1910

20.181, 20.182, 20.183 and 20.184); spermatheca with stem with only a single long, sinuate loop posteriorly (Figs. 20.179d, 20.189d, 20.181d, 20.182d, 20.183d and 20.184d) ........................... ................................. Prudhoensis species group – Tubus of median lobe of aedeagus in dorsal view with sides parallel to slightly constricted from apex of bulbus (Fig. 20.178c), in lateral view tubus with entire ventral margin arcuate, straight, or abruptly bent ventrad near apex (Fig. 20.178b); internal sac with two mediumsized, elongate sclerites in bulbus and additional structures in tubus (Fig. 20.178b, c); spermatheca with stem long, thin, with multiple dense, irregular coils posteriorly (Fig. 20.178h) ...................... .....................................Picipennis species group

REVIEW OF SPECIES GROUPS AND SPECIES OF SUBGENUS DIMETROTA

Alesi species group (Brunke et al. 2012; Klimaszewski et al. 2018b) Diagnosis. Antennae robust with antennomeres V–X moderately transverse; integument of forebody highly glossy with dense microsculpture of irregularly hexagonal sculpticells; elytra with punctation finely asperate, pubescence appearing somewhat silky; median lobe of aedeagus in dorsal view with enlarged bulbus and short and triangular tubus, in lateral view tubus short and straight ventrally; internal sac with thin and inconspicuous structures in bulbus; male tergite VIII with shallowly concave apical margin and very small lateral teeth; spermatheca with tubular capsule and stem highly sinuate and narrowly looped posteriorly. Comments. This group is tentatively assigned to the subgenus Dimetrota, because the two species included here have weakly asperate punctation on the forebody in comparison to strongly asperate in other Dimetrota species, the shape of spermatheca and male tergite VIII are also unique in Alesi group. Atheta alesi was originally placed

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in the subgenus Microdota but its size and the size of its sibling species exceeds the size range of much smaller species in the subgenus Microdota. The shape of median lobe of aedeagus and spermatheca resemblance those of some Palearctic Atheta (Bessobia) species. The taxonomic status of this group is unsettled. 131. Atheta (Dimetrota) alesi Klimaszewski and Brunke (Fig. 20.166a–h) Atheta (Microdota) alesi Klimaszewski and Brunke, in Brunke et al. 2012. Atheta (Dimetrota) alesi: Klimaszewski et al. 2015c, 2018b, Webster et al. 2016. Diagnosis. Body narrowly subparallel, slightly flattened, length 2.4–2.6 mm; colour dark brown, with legs/tarsi, basal parts of antennae and elytra yellowish-brown; integument highly glossy (Fig. 20.166a); forebody with punctation fine and sparse, with coarse irregularly hexagonal microsculpture; pubescence short, appearing silky; pronotum with pubescence on disc directed laterad and obliquely posteriad from midline, that on elytra directed obliquely posteriad; head rectangular, slightly narrower than pronotum; antennomeres V–X transverse; pronotum transverse, about 1/6 narrower than base of elytra; elytra transverse, at suture about as long as pronotum or slightly longer; abdomen arcuate laterally. MALE. Tergite VIII with apical margin with broadly reversed V-shaped emargination, margin with some crenulation, and with small teeth-like angular lateral edges (Fig. 20.166d); sternite VIII rounded apically (Fig. 20.166e); tubus of median lobe of aedeagus straight ventrally and triangular at apex in lateral view (Fig. 20.166b); in dorsal view tubus short, triangular apically, and bulbus broadly oval and large (Fig. 20.166c). FEMALE. Tergite VIII arcuate apically (Fig. 20.166f); sternite VIII rounded apically (Fig. 20.166g); spermatheca with tubular, sinuate capsule, and sinuate broad stem narrowly looped posteriorly (Fig. 20.166h).

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DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], ON. USA: not recorded. We report this species from BC for the first time here based on the following: British Columbia, Creston, near Kootenay Pass, 1500–1800 m, 18.VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 male, 2 females; near Cherryville at Monashee Summit, 1100–1200 m, 8.VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 female. COLLECTION AND HABITAT DATA. Habitat: the ON specimens were found in open fields, farmland where this species occurs in Marmota monax (L.) burrows; also collected from raised pan trap in agricultural hedgerow. Collecting period: IV–V. Collecting method: sifting litter from burrows, canopy trap. Comments. The BC specimens agree in external morphology and structures of genital characters with the Ontario population, and the only difference we could find was less contrasting colour of elytra in comparison to the rest of the body in BC specimens. No habitat data was specified for the BC specimens. Campbelli Species Group (Klimaszewski et al. 2018b) Diagnosis. Antennae slender, antennomeres V– X moderately to distinctly transverse; integument of forebody moderately glossy, with microsculpture moderate to coarse, consisting of irregularly hexagonal sculpticells; median lobe of aedeagus in dorsal view with bulbus enlarged, tubus short and triangular, in lateral view tubus straight and strongly produced ventrally at apex; internal sac with two coarse, falciform sclerites in bulbus; male tergite VIII with apical margin shallowly concave (sinuate medially), with very small lateral teeth, or truncate with large apical teeth and apical margin crenulate or emarginate; spermatheca with capsule club-shaped, tubular with apical invagination narrow and moderately deep, stem sinuate.

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Tribe Athetini Casey, 1910

132. Atheta (Dimetrota) remulsa Casey (Fig. 20.167a–h) Atheta (Atheta) remulsa Casey, 1910. Casey 1910 [as the synonyms A. asperiola and A. frugalis], Klimaszewski et al. 2005, 2007b, 2011, 2016d, 2018b, Dollin et al. 2008, Majka and Klimaszewski 2008b. Diagnosis. Body narrowly subparallel, slightly flattened, length 2.4–2.8 mm; colour dark brown to black, with legs/tarsi, basal parts of antennae and elytra yellowish-brown or reddish-brown; integument moderately glossy (Fig. 20.167a); forebody with punctation dense, coarse and finely asperate, with coarse meshed microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad, forming arcuate lines from midline, that on elytra directed obliquely posteriad forming wavy pattern posteriorly; head slightly narrower than pronotum; antennae moderately robust, antennomeres V–X distinctly transverse; pronotum transverse, about 1/5 narrower than width of base of elytra; elytra transverse, at suture as long as pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically, with two lateral teeth and medially emarginated broad projection (Fig. 20.167d); sternite VIII rounded apically (Fig. 20.167e); tubus of median lobe of aedeagus straight basally and strongly produced ventrally in apical half in lateral view (Fig. 20.167b); in dorsal view bulbus broadly oval and tubus broadly triangular (Fig. 20.167c). FEMALE. Tergite VIII slightly emarginate medially at apex (Fig. 20.167f); sternite VIII rounded apically (Fig. 20.167g); spermatheca with tubular, clubbed-shaped capsule with asymmetrically located apical invagination, and with long sinuate stem narrowly looped posteriorly (Fig. 20.167h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK, YT. USA: AK, NY. New locality data: USA, Alaska, Taylor Hwy, mi 82, 8.VII.1968, A. Smetana (CNC) 1 female.

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Tribe Athetini Casey, 1910

COLLECTION AND HABITAT DATA. Habitat: in various forest types, usually in mushrooms; yellow birch/balsam fir forest, red spruce forest, old-growth hemlock forest, mixed forests, deciduous forests, and a red oak forest; most individuals in gilled mushrooms and fleshy fungi in various stages of decay. Also from leaf litter and moss in Carex marsh and forest litter. Collecting period: V–X. Collecting method: sifting litter and fungi, pitfall traps, pitfall Luminoc traps, flight intercept traps. Columbica Species Group, New Diagnosis. Antennae slender, antennomeres V– X moderately transverse; integument of forebody moderately glossy, with microsculpture moderate to coarse, consisting of irregularly hexagonal sculpticells; median lobe of aedeagus in dorsal view with bulbus large, oval in shape, tubus short and triangular, in lateral view tubus straight basally and apically strongly produced ventrad, apical part triangular; internal sac with two long, falciform sclerites in bulbus; male tergite VIII with apical margin variable in shape; spermatheca with capsule club-shaped, stem thin, very long, sinuate and narrowly looped posteriad. 133. Atheta (Dimetrota) stercoris Fenyes [¼A. columbica Bernhauer] (Fig. 20.168a–h) Atheta stercoris Fenyes, 1920. As Dimetrota: Moore and Legner 1975. Diagnosis. Body moderately convex, broad, subparallel, length 3.7 mm, dark brown to black, elytra with some reddish tinge, apical part of abdomen, and legs paler, antennae uniformly dark brown (Fig. 20.168a); forebody moderately densely punctate and pubescent, pubescence long, punctation slightly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head subquadrate, round posteriorly, distinctly narrower than pronotum, postocular region shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; antennae

373

slender, shorter than pronotum and elytra combined, antennomere IV subquadrate to slightly transverse, VII–X slightly transverse, basal 3 antennomeres glossy and remaining matte; pronotum impressed mesobasally, transverse, approximately rectangular in shape, basal margin sinuate, hypomeron visible for two thirds of pronotal length in lateral view, pubescence directed from midline of disc anteriad at apical third, on sides nearly horizontally laterad medially, and obliquely laterad posteriorly; elytra slightly convex, distinctly transverse, broader than pronotum, at suture slightly longer than pronotum along midline, posterior margins feebly sinuate and with shallow, scarcely visible, lateral emarginations, pubescence directed straight or slightly obliquely posteriad; abdomen subparallel, strongly glossy, four first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII with two distinct arcuate lateral teeth and margin between teeth strongly sinuate (Fig. 20.168d); sternite VIII parabolic apically (Fig. 20.168e); median lobe of aedeagus broadly oval, apex narrowly triangular in dorsal view (Fig. 20.168c); in lateral view tubus with ventral margin about straight basally and arcuate apically, apex narrowly triangular and produced ventrad (Fig. 20.168b). FEMALE. Tergite VIII transverse and broadly truncate and slightly emarginated medially (Fig. 20.168f): sternite VIII transverse, broadly rounded apically (Fig. 20.168g); spermatheca with capsule small, hemispherical apically and with elbowed neck posteriorly, stem thin, long, broadly U-shaped medially, and narrowly looped posteriorly (Fig. 20.168h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. New locality data: British Columbia, Queen Charlotte Is., 4.7 km N Rennell, Sound Rd., Ghost Cr., 19.VII.1983, 83-24, 700 ft., ex gravel along edge of stream, J.M. Campbell (CNC) 1 male, 1 female; BC, Queen Charlotte Islands, Moresby Is., Tekakia Lk., 21.VII.1983, 83-28, 30000 , J.M. Campbell (CNC) 1 male, 1 female;

374

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Queen Charlotte Is., Moresby Is., Mt. Moresby, 26.VII.1983, sifting bear dung, J.M. Campbell (CNC) 2 males, 4 females; Queen Charlotte Is., Tlell Riv., end Richardson Rd., 17.VII.1983, 83-14, ex rotting Boletus, coll. J.M. Campbell (CNC) 1 male, 2 females.

Etymology. Named for our friend, Neville Winchester, professor of the University of Victoria, BC, Canada, who dedicated his life to study of insects and who pioneered canopy arthropod studies in Canada. He also collected the type series of this species.

COLLECTION AND HABITAT DATA. Habitat: found in transition zone of Sitka spruce forest (Carmanah Valley), in gravel along edge of stream; and in bear dung (QCI). Collecting period: VI–IX. Collecting method: pitfall traps, sifting organic litter, sifting bear dung, ex rotting Boletus, hand picking from gravel along edge of stream.

Diagnosis. Body narrowly elongate, subparallel, length 2.9–3.2 mm; colour dark brown to black, with bases of antennae, central part of elytra, and legs or tarsi only reddish-brown (Fig. 20.169a); integument moderately glossy, forebody with fine, dense punctation and meshed microsculpture; head round, almost as wide as pronotum and elytra, with eyes shorter than genae in dorsal view; antennae moderately robust, antennomere IV subquadrate, V–X slightly to moderately transverse; pronotum evenly arcuate laterally, widest at middle of its lenghth, all angles broadly rounded; elytra at suture as long as pronotum along midline; abdomen subparallel. MALE. Tergite VIII with apical margin truncate, with two minute lateral teeth and margin between teeth crenulate (Fig. 20.169d); sternite VIII parabolic apically, base of disc sinuate (Fig. 20.169e); median lobe of aedeagus in lateral view with tubus straight basally and strongly produced ventrally in alpical half (Fig. 20.169b), in dorsal view bulbus large and oval in shape, tubus narrowly elongate and narrowly triangular apically (Fig. 20.169c). FEMALE. Tergite VIII with apical margin arcuate, apex truncate (Fig. 20.169f); sternite VIII rounded apically, antecostal suture strongly sinuate (Fig. 20.169g); spermatheca with capsule narrowly club-shaped and stem very thin, highly sinuate and narrowly looped posteriorly (Fig. 20.169h).

134. Atheta (Dimetrota) winchesteri Klimaszewski, sp. n. (Fig. 20.169a–h) Holotype (male): Canada, British Columbia, Mt. Cain, 50.13 N, 126.21 W, 20.VII-5. VIII.1997, 16/6 PIT 10, coll. N. Winchester (CNC). Paratypes: Canada, British Columbia, Mt. Cain, 50.13 N, 126.21 W, 9.VI-23.VI.1997, 17.4 PIT 2, coll. N. Winchester (LFC) 2 females; Canada, British Columbia, Mt. Cain, 50.13 N, 126.21 W, 23.VI-7.VII.1997, 16.6 PIT 4, coll. N. Winchester (CNC, LFC, RBCM) 17 males, 37 females; British Columbia, Mt. Cain, 50.13 N, 126.21 W, 27.VI-13. VII.1996, coll. N. Winchester (LFC) 1 male, 1 sex undetermined; British Columbia, Mt. Cain, 50.13 N, 126.21 W, 7.VII-20. VII.1997, 16.6 PIT4, coll. N. Winchester (CNC, LFC, RBCM) 12 males, 34 females; Canada, British Columbia, Mt. Cain, 50.13 N, 126.21 W, 20.VII-5.VIII.1997, 16/6 PIT 10, coll. N. Winchester (CNC, LFC, RBCM) 13 males, 27 females; British Columbia, Mt. Cain, 50.13 N, 126.21 W, 5.VIII-18. VIII.1997, 16.6 PIT 3, N. Winchester (CNC, LFC, RBCM) 3 males, 3 females; British Columbia, Mt. Cain, 50.13 N, 126.21 W, 18.VIII-1. IX.1997, 16.6 PIT, coll. N. Winchester (LFC) 2 females; British Columbia, Mt. Cain, 50.13 N, 126.21 W, 1.IX-19.IX.1997, 16.6 PIT 8, N. Winchester (CNC, LFC, RBCM) 2 males, 2 females.

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Macrohabitat: boreal forest. Collecting period: VI–IX. Collecting method: pitfall traps.

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Tribe Athetini Casey, 1910

Crassicornis (=Modesta) species group, new interpretation (Gusarov 2003a; Klimaszewski et al. 2018b) Diagnosis. Antennae slender, antennomeres V– X moderately to distinctly transverse; integument of forebody highly glossy, with coarse microsculpture consisting of irregularly shaped hexagonal sculpticells; pubescence on disc of elytra directed approximately straight posteriad; median lobe of aedeagus in dorsal view with bulbus large, elongate, oval, tubus moderately long and triangular apically, in lateral view tubus arcuate or straight in basal half and more or less produced ventrally at apex, bulbus of internal sac with two highly elongate sclerites and additional complex structures; male tergite VIII with apical margin straight or convex, with lateral teeth small or large and rounded; spermatheca with capsule moderately large, club-shaped, with small apical invagination, stem long, thin, sinuate, and coiled and twisted posteriorly. 135. Atheta (Dimetrota) godini Klimaszewski, sp. n. (Fig. 20.170a–h) Holotype (male): Canada, British Columbia, Valemont, Yellowjacket ck., 24-29.VI. 2014, pitfall, Kinbasket # 1181, 52.703 N, 119.049 W, 771 m, R-Mature, coll. Charlene Wood (RBCM). Paratypes: BC, Valemont, Yellowjacket ck., 52.703 N, 119.049 W, 24-29. VI. 2014, pitfall, Kinbasket # 1179, 1181, 771 m, R-Mature, coll. Charlene Wood (RBCM) 1 male, 1 female; Valemont, Yellowjacket ck., 52.703 N, 119.049 W, 3-8.VI.2014, pitfall, Kinbasket # 1053, 1054, 1059, 1064, 1074, R-Mature, coll. Charlene Wood (RBCM) 12 males, 10 females; BC, 13 km N Topley, 54.616 N, 126.3 W, VIIIIX.2012, pitfall # 389, 1132 m, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 male; BC, Log Lk., 65 km N Prince George, 54.2 N, 122.616 W, 780 m, pitfall # 208, OM3C1SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 2 males.

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Etymology. This species is dedicated to Benoit Godin, Whitehorse, Yukon, a very good friend of the first author and collaborator in many aleocharine projects in Canada. Diagnosis. Body length 3.9–4.2 mm, head, most of antennae, pronotum and abdomen dark brown to almost black, and elytra except for scutellar and lateral regions, 2–3 basal antennomeres, and legs yellow brown (Fig. 20.170a); microsculpture of forebody strong, sculpticells hexagonal, punctation moderately dense; forebody slightly flattened; moderately glossy; head moderately narrower than pronotum, postocular area shorter than diameter of eye seen from above; eyes large; antennomere IV subquadrate, VII–X subquadrate; pronotum strongly transverse, broadest at middle, truncate basally, pubescence on midline directed mostly anteriad and on side laterad and obliquely laterad; elytra broad, subparallel, at suture slightly longer than pronotum along midline; abdomen subparallel. MALE. Tergite VIII truncate apically without teeth but angular laterally (Fig. 20.170d); sternite VIII parabolic apically (Fig. 20.170e); tubus of median lobe of aedeagus short, venter slightly sinuate, apex triangular and slightly produced ventrally in lateral view (Fig. 20.170b), in dorsal view tubus broadly oval with two narrow lateral projections of internal sac and semicircular apical sclerites (Fig. 20.170c); internal sac as illustrated (Fig. 20.170b, c). FEMALE. Tergite VIII truncate apically, slightly emarginate medially (Fig. 20.170f); sternite VIII rounded apically (Fig. 20.170g); spermatheca not found; pygidium as illustrated (Fig. 20.170h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: sub-boreal spruce forest. Collecting period: VI–IX. Collecting method: pitfall traps. 136. Atheta (Dimetrota) Bernhauer (Fig. 20.171a–h)

metlakatlana

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Atheta (Atheta) metlakatlana Bernhauer, 1909. Casey 1910 [as Athetalia], Hatch 1957 [as Stethusa]. As Atheta: Moore and Legner 1975, Klimaszewski and Winchester 2002. Diagnosis. Body moderately convex, narrowly oval, length 3.5–4.0 mm, dark brown, elytra light brown to yellowish brown and sometimes with rust brown tinge, legs yellowish brown, antenna uniformly dark brown with basal segment light brown to yellowish brown (Fig. 20.171a); forebody densely punctate and pubescent, pubescence long, punctation strongly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head as broad as long or slightly elongate, round, little narrower than pronotum, postocular region shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; antennae slender and long, slightly incrassate, as long as the pronotum and elytra combined, antennomeres VII–X slightly transverse, terminal one elongate and as long as two preceding antennomeres combined, basal 4 antennomeres glossy and remaining matte; pronotum impressed mesobasally, strongly transverse, approximately rectangular in shape, pubescence directed anteriad at one third of apical midline, and obliquely laterad across arcuate lines elsewhere; elytra moderately convex, slightly elongate or about as long as wide, broader than pronotum, posterior margins broadly arcuate and with shallow lateral emarginations, pubescence directed obliquely posteriad; abdomen subparallel, strongly glossy, three first visible tergites deeply impressed at base; basal mesotarsomere as long as the following one. MALE. Tergite VIII transverse, with two short, broad, and rounded lateral teeth, margin between teeth obtuse concave or straight and crenulate (Fig. 20.171d); sternite VIII about as long as wide, and parabolic apically (Fig. 20.171e); in dorsal view median lobe of aedeagus with enlarged, broadly suboval bulbus and tubus short, triangular (Fig. 20.171c), in lateral view tubus short, straight mesomedially, apex slightly produced ventrally (Fig. 20.171b). FEMALE. Tergite VIII transverse and truncate apically

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Tribe Athetini Casey, 1910

(Fig. 20.171f); sternite VIII transverse truncate apically and with fringe of long and dense apical setae (Fig. 20.171g); spermatheca with capsule hemispherical and round apically, stem narrowly elongate and broadly looped posteriorly with two small coils at apex (Fig. 20.171h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: OR. New locality data: British Columbia, Queen Charlotte Is., Port Clement, 20.VII.1983, 83-25, sifting mushrooms, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Lyall Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, coll. J.M. Campbell (CNC) 2 males; Queen Charlotte Is., Skowkona Cr., 7.9 km NW Queen Charlotte City, 29.VII-4. VIII.1983, 83-55A, ex rotting pile of mushrooms, J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Moresby Is., Haswell Bay, 1.5 km SW Hoskins Point, 13.VIII.1983, 83-79, sifting mushrooms, J.M. Campbell (CNC) 1 male. COLLECTION AND HABITAT DATA. Habitat: transition zone and forest interior in Sitka spruce forest, Carmanah Valley; rotting mushrooms, marten dung on thick moss. Collecting period: VI–IX. Collecting method: sifting forest litter, marten dung and mushrooms, pitfall traps. 137. Atheta (Dimetrota) pseudometlakatlana Klimaszewski and Godin (Fig. 20.172a–h) Atheta (Dimetrota) pseudometlakatlana Klimaszewski and Godin, in Klimaszewski et al. 2008b [new subgeneric combination]. Diagnosis. Externally, this species is hardly distinguishable from its sister species A. metlakatlana (see description and Fig. 20.171a–h). There are minute external differences between the two species, and A. pseudometlakatlana may be distinguished from A. metlakatlana by the following: slightly more transverse antennomeres VII–X

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Tribe Athetini Casey, 1910

(Fig. 20.172a), and slightly shorter and more transverse elytra (Fig. 20.172a). The two species may be easily differentiated by the shape of tubus of median lobe of aedeagus and the internal sac structures. In A. metlakatlana tubus of median lobe is almost straight and horizontal with short subapical part and apex slightly produced ventrad in lateral view (Fig. 20.171b), and in A. pseudometlakatlana tubus is strongly produced ventrad with long subapical part and apex strongly produced ventrad in lateral view (Fig. 20.172b). The terminal segments of male and female and spermatheca are very similar in the two species. Females may be identified with certainty only by associations with males and slight differences in the shape of the capsule of the spermatheca (Figs. 20.171h and 20.172h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], YT. USA: not recorded. We report this species from BC for the first time here based on the following: British Columbia, Boya Lake, 59.3639-129.1035, 698 m, 28. VI.2015, mushroom, sifting, coll. B. Godin and D. Horwood (BGC) 1 female; Log Cabin, 59.7589-134.9669, 902 m, 14.VI.205, mushroom, soil sifting, coll. B. Godin and D. Horwood (BGC, LFC) 9 males, 3 females; Log Lake, 65 km N of Prince George, 54.2122.616, 780 m, VIII-IX.2012, pitfall # 208, OM3C1SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 male, 3 females; Valemont, N of Packsaddle Creek, 52.78119.164, 767 m, 24-29.VI.2014, Kinbasket # 1212, R-Mature, coll. Charlene Wood (RBCM) 3 females. COLLECTION AND HABITAT DATA. Habitat: sub-boreal spruce forest, forest litter, fungi. Collecting period: VI–IX. Collecting method: sifting forest litter and fungi, pitfall traps. 138. Atheta (Dimetrota) wheelerae Klimaszewski, sp. n. (Fig. 20.173a–h)

377

Holotype (male): Canada, British Columbia, Queen Charlotte Is., Yaku, NW corner, Graham Is., 17.VIII.1983, 83-92, sifting mushrooms and dung, J.M. Campbell (CNC). Paratypes: Canada, BC, Queen Charlotte Is., Moresby Is., Cumsheva, 14.VIII.1983, 83-82, sifting squirrel midden besides totem pole, J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Moresby Is., Haswell Bay, 1.5 km SW Hoskins Point, 13.VIII.1983, sifting mushrooms, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Moresby Is., Mt. Moresby, 26.VII.1983, 83-39, 2100 feet, sifting bear dung, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Skowkona Cr., 7.9 km NW Queen Charlotte City, 29.VII-4.VIII.1983, ex rotting pile of mushrooms, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Burnaby Is., N shore, Section Cove, 11 and 12.VIII.1983, 83-76, sifting moss under dung and mushrooms, and sifting moss and mushrooms, J.M. Campbell (CNC) 1 male, 2 females; Queen Charlotte Is., 4.7 km N Rennell Sound Rd., Ghost Cr., 19. VII.1983, 83-24, 700 ft., ex gravel along edge of stream, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Lyall Is., Gate Cr., 10. VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, coll. J.M. Campbell (CNC) 1 male, 1 female; Queen Charlotte Is., Port Clement, 20.VII.1983, 83-25, sifting mushrooms, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Tlell Riv., end Richardson Rd., 17.VIII.1983, ex rotting Boletus, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Slatechuck Mt., 14.VII.1983, 83-7, sifting Sphagnum at edge of seep., 2200 ft., coll. J.M. Campbell (CNC) 1 male. Etymology. This species is dedicated to Dr. Erica Wheeler, Head of Collection Care and Conservation at the Royal British Columbia Museum, Victoria, for her role in supporting BC biodiversity projects. Diagnosis. Body length 3.9–4.3 mm, head, most of antennae, pronotum and abdomen dark brown

378

to almost black, and elytra except for some dark brown irregular patches, bases of antennae and legs yellow brown (Fig. 20.173a); microsculpture of forebody strong, sculpticells hexagonal, punctation moderately dense; head moderately narrower than pronotum, postocular area shorter than diameter of eye seen from above; eyes large; antennomere IV subquadrate, VII–X moderately transverse; pronotum transverse, evenly arcuate laterally, truncate basally, pubescence on midline directed mostly anteriad and on side laterad and obliquely laterad; elytra broad, subparallel, at suture slightly longer than pronotum along midline; abdomen subparallel. MALE. Tergite VIII truncate apically without teeth but angular laterally (Fig. 20.173d); sternite VIII parabolic apically (Fig. 20.173e); tubus of median lobe of aedeagus short, slightly arcuate basally and straight apically, apex rounded in lateral view (Fig. 20.173b), in dorsal view bulbus large, oval, tubus with two narrow lateral projections (Fig. 20.173c); internal sac with two eleongate apical sclerites as illustrated (Fig. 20.173b, c). FEMALE. Tergite VIII truncate apically, slightly sinuate medially (Fig. 20.173f); sternite VIII truncate apically and slightly emarginated medially, with row of strong apical setae (Fig. 20.173g); spermatheca with club-shaped capsule with minute apical invagination, stem long, arcuate, looped posteriad and twisted apically (Fig. 20.173h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: squirrel midden, rotting mushrooms, marten dung and other dung on thick moss, in gravel along edge of stream. Collecting period: VII, VIII. Collecting method: sifting moss, dung and mushrooms. Districta Species Group (Klimaszewski et al. 2018b) Diagnosis. Antennae thick and robust, antennomeres V–X elongate or subquadrate;

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Tribe Athetini Casey, 1910

integument of forebody moderately glossy, with fine microsculpture consisting of irregularly shaped hexagonal sculpticells; median lobe of aedeagus in dorsal view with medium-sized bulbus and long and apically triangular tubus, in lateral view tubus straight or slightly produced ventrally at apex; internal sac with two moderately elongate sclerites in bulbus; male tergite VIII with approximately straight apical margin and small lateral teeth; spermatheca with long tubular capsule bearing tubular extention and small apical invagination, and with sinuate stem. 139. Atheta (Dimetrota) districta Casey (Fig. 20.174a–h) Atheta (Atheta) districta Casey, 1911. Klimaszewski et al. 2005, 2007b, 2011, 2016d, 2018b, Majka and Klimaszewski 2008b. Diagnosis. Body narrowly elongate, slightly flattened, length 2.8–3.3 mm; colour dark brown to black, with legs/tarsi, basal parts of antennae and elytra yellowish-brown (Fig. 20.174a); integument moderately glossy; forebody with punctation dense, finely asperate on elytra; with fine isodiametric microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad from midline, that on elytra directed obliquely or straight posteriad; head distinctly narrower than pronotum; antennae thick, antennomeres V–X slightly elongate; pronotum transverse, about 1/7 narrower than width of base of elytra; elytra transverse, at suture as long as pronotum; abdomen arcuate laterally. MALE. Tergite VIII truncate apically, angular laterally and straight or slightly concave medially (Fig. 20.174d); sternite VIII parabolic apically (Fig. 20.174e); in lateral view tubus of median lobe of aedeagus long, slightly arcuate basally and straight medially and with moderately thin and rounded apical part (Fig. 20.174b); in dorsal view bulbus moderately large and suboval in shape, tubus long and narrow with triangular apex (Fig. 20.174c). FEMALE. Tergite VIII arcuate apically (Fig. 20.174f); sternite VIII rounded apically (Fig. 20.174g); spermatheca with tubular

20

Tribe Athetini Casey, 1910

capsule bearing arched apical tubular projection with small apical invagination, stem thin, looped and swollen posteriorly (Fig. 20.174h). The median lobe of the aedeagus and particularly the spermatheca of this species are similar to its sibling species A. pseudodistricta, from which it differs in having antennomeres V–X elongate and the median lobe of the aedeagus shaped differently. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, in various forest types usually in dung or decaying mushrooms; eastern white cedar swamp in moose dung, mature red spruce and eastern white cedar forest in deer dung, mixed forest in bear dung, in decaying (moldy) corncobs and cornhusks near mixed forest, regenerating mixed forest in decaying mushrooms, near lakeshore in decaying fungi, old jack pine forest in rotten Boletus mushrooms, red maple swamp in leaf litter near stream, in unbaited pitfall traps in red spruce forest, and a few collected at u.v. light. Elsewhere, captured in flight intercept traps and carrion-baited pitfall traps in coniferous and mixed forests (NF) and on coastal barrens (LB); also from spruce litter, and dry and decaying mushrooms. Collecting period: VI–IX. Collecting method: sifting litter, dung, and fungi, pitfall traps, flight intercept traps. Fanatica Species Group (newly defined here) Diagnosis. Body length 2.5–3.3 mm, subparallel, antennae slender, antennomeres V–X subquadrate to moderately transverse; integument of forebody glossy, with microsculpture distinct; punctation of forebody dense and asperate; median lobe of aedeagus in dorsal view with bulbus oval in outline and streamlined with base of tubus, tubus short and triangular; in lateral view tubus broad, slightly to strongly sinuate ventrally and apex broadly rounded not produced

379

ventrad; internal sac with two complex sclerites in bulbus; male tergite VIII with apical margin truncate, with small lateral teeth, apical margin slightly crenulate; male sternite VIII with long macrosetae; spermatheca with capsule spherical or broadly tubular rounded apically, apical invagination present, stem sinuate and looped posteriorly. 140. Atheta (Dimetrota) fanatica Casey (Fig. 20.175a–h) Atheta (Atheta) fanatica Casey, 1910. Casey 1911, Majka et al. 2006 [as Atheta irrita], Klimaszewski et al. 2011, 2012, 2015a, 2018b, Bousquet et al. 2013, Webster et al. 2009 [as Atheta irrita]. Atheta omissa Casey, 1910. Synonymized by Gusarov 2003b. Atheta vicaria Casey, 1910. Synonymized by Gusarov 2003b. Diagnosis. Body narrowly subparallel, slightly flattened, length 2.0–2.3 mm; colour dark brown to almost black, with legs/tarsi, and maxillary palpi yellowish-brown or reddish-brown (Fig. 20.175a); integument moderately glossy; forebody with punctation dense and asperate, with fine microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad from midline, forming transversely arcuate pattern, that on elytra directed obliquely posteriad forming wavy pattern posteriorly; head slightly narrower than pronotum; antennae slender, antennomeres V–X slightly to distinctly transverse; pronotum transverse, about 1/7 narrower than width of base of elytra; elytra transverse, at suture slightly longer than pronotum; abdomen arcuate laterally. MALE. Apical margin of tergite VIII truncate, minutely crenulate, slightly obtusely angulate laterally (Fig. 20.175d); sternite VIII parabolic apically (Fig. 20.175e); tubus of median lobe of aedeagus highly angular medioventrally and with broadly rounded apex in lateral view (Fig. 20.175b); in dorsal view bulbus large and oval, tubus, short, broadly triangular (Fig 20.175c). FEMALE. Tergite VIII truncate

380

apically (Fig. 20.175f); sternite VIII rounded apically (Fig. 20.175g); spermatheca with sac-shaped, tubular capsule, and sinuate stem narrowly looped posteriorly (Fig. 20.175h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NS, QC, SK, YT. USA: NV. COLLECTION AND HABITAT DATA. Habitat: Atheta fanatica appears be associated with owl nests and other decaying organic material on standing trees. In NB, this species was a common inhabitant of nests of barred owls that nest in tree holes in large trees and artificial nest boxes in hardwood, silver maple, and mixed forests. The material in nests usually consisted of rich decaying organic material with bones, fur, owl pellets and portions of dead prey items and the contents often had a strong urine smell. This species was also found in nest contents of the great horned owl (NB), boreal owl, and northern saw-whet owl (NS). It also has been found in Pleurotus sp. (oyster mushrooms) in the early stages of decay on dead standing trembling aspen trees in oak forests, Populus stands, hardwood forests, and mixed forests. One specimen from Lindgren funnel trap in red pine forest (NB) and one from NF captured in a carrionbaited pitfall trap in a spruce/moss forest. Collecting period: V–VIII. Collecting method: sifting owl nest contents, mushrooms, and litter, carrion-baited pitfall traps, Lindgren funnel trap. Comments. This species was reported from different parts of North America under different synonymic names (Klimaszewski et al. 2011). It is related to the western Palearctic Atheta vaga (Heer), but in A. fanatica the shape of the median lobe of the aedeagus is slightly different, with the tubus having a more pronounced median projection in lateral view, and the apical margin of male tergite VIII is not as distinctly emarginate. Both species have the tubus of median lobe very broad and evenly coalescent with the bulbus in dorsal view, and the shapes of the structures of the

20

Tribe Athetini Casey, 1910

internal sac of the median lobe are similar (Lohse et al. 1990: Fig. 20.173). 141. Atheta (Dimetrota) munsteri Bernhauer  (Fig. 20.176a–h) Atheta (Dimetrota) munsteri Bernhauer, 1902. Lohse et al. 1990 (as Dimetrota); Bousquet et al. 2013 (as Atheta (Dimetrota)); Smetana 2004 (as Boreophilia); Schülke and Smetana 2015 (as Boreophilia); Klimaszewski et al. 2019 (as Atheta (Dimetrota)). Lectotype (male): Dovre; Fauvel vidis; x; Je ne l’ai pas; male; Chicago NHMus.; M. Bernhauer Collection; Munsteri Bernh.; Type Norwegen; ischnocera tps?; Lectotypus male Atheta munsteri Bernhauer, V.I. Gusarov des. 1998; Dimetrota munsteri (Bernh.) V.I. Gusarov (FMNH). Present lectotype designation. Paralectotypes: Dovre; munsteri, Brh. Type, Norwegen; Paralectotype male Atheta munsteri Bernhauer, V.I. Gusarov des. 1998; Dimetrota munsteri (Bernh.) V.I. Gusarov (FMNH) 1 male; Dovre; Münster; münsteri, B., Norwegen; CNHM 1955; Eduard Knirch Palearctic coll.; Paralectotype female Atheta munsteri Bernhauer, V.I. Gusarov des. 1998; Dimetrota munsteri (Bernh.) V.I. Gusarov (FMNH) 1 female. Diagnosis. Body narrowly subparallel, slightly flattened, length 2.6–3.0 mm; colour uniformly black (Fig. 20.176a); integument moderately glossy; forebody with punctation sparse and asperate, with fine microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad from midline, and that on elytra directed obliquely posteriad forming slightly wavy pattern posteriorly; head slightly narrower than pronotum; antennae slender, antennomeres V–X subquadrate to slightly transverse; pronotum transverse, about 1/3 narrower than width of base of elytra; elytra transverse, at suture slightly longer than pronotum; abdomen arcuate laterally. MALE. Apical margin of tergite VIII truncate, minutely crenulate, slightly

20

Tribe Athetini Casey, 1910

obtusely angulate laterally (Fig. 20.176d); sternite VIII parabolic apically with strong macrosetae (Fig. 20.176e); tubus of median lobe of aedeagus highly sinuate medio-ventrally and with broadly rounded apex in lateral view (Fig. 20.176b); in dorsal view bulbus large and oval, tubus short, broadly triangular (Fig 20.176c). FEMALE. Tergite VIII truncate apically, apical margin feebly concave (Fig. 20.176f); sternite VIII rounded apically (Fig. 20.176g); spermatheca S-shaped, with spherical capsule, and strongly sinuate stem looped posteriorly (Fig. 20.176h). DISTRIBUTION. Origin: Holarctic. CANADA: BC [new provincial record], MB, NT, YT. USA: AK. We report this species from BC for the first time here based on the following: British Columbia, Log Cabin, 59.7589-134.9669, 902 m, 14. VI.2015, mushroom, soil sifting, coll. B. Godin and D. Horwood (LFC) 1 female. COLLECTION AND HABITAT DATA. Habitat: fungi/litter in unspecified forest. Collecting period: VI. Collecting method: sifting forest litter and fungi. Comments. This BC record should be considered as a tentative identification because it is based on a single female. A male from BC is essential to validate this record. We designated the lectotype from the syntypes because Gusarov labelled the male syntype as a lectotype but did not publish his designation. Klimaszewski et al. (2019) excluded this species from the genus Boreophilia Benick on the grounds of morphological and DNA studies. Hampshirensis Species Group (newly defined here) Diagnosis. Body small, length 2.0–2.6 mm, antennae slender, antennomeres V–X moderately transverse; integument of forebody slightly glossy, with microsculpture weak; punctation of forebody dense and asperate; median lobe of

381

aedeagus in dorsal view with bulbus oval in outline, tubus short and triangular; in lateral view tubus nearly straight ventrally and apex not produced ventrad; internal sac with two narrowly elongate sclerites in bulbus; male tergite VIII with apical margin truncate, with thin and long lateral teeth, apical margin slightly crenulate; spermatheca with capsule tubular, strongly curved medially and narrowed apically, apex pointed ventrad, apical invagination absent, stem sinuate and looped posteriorly. The taxonomic position of this group is unsettled and is tentatively assigned to the subgenus Dimetrota. 142. Atheta (Dimetrota) Bernhauer (Fig. 20.177a–h)

hampshirensis

Atheta (Dimetrota) hampshirensis Bernhauer, 1909. Lohse and Smetana 1985 [as Dimetrota moesta], Klimaszewski and Winchester 2002, Gusarov 2003a [list of synonymies including Dimetrota retrusa Casey from Metlakatla, BC], Klimaszewski et al. 2005, 2011, 2016b, 2018b, Majka and Klimaszewski 2008b, Majka and Sikes 2009. Diagnosis. Body narrowly oval, slightly flattened, length 2.2–2.6 mm; colour dark brown to black, with legs/tarsi, basal parts of antennae yellowish brown and elytra light brown with rusty tinge (Fig. 20.177a); integument moderately glossy; forebody with punctation dense and asperate, with fine microsculpture; pubescence on disc of pronotum directed laterad from midline, that on elytra directed obliquely posteriad; head slightly narrower than pronotum; antennae slender, antennomeres V–X slightly transverse; pronotum transverse, about 1/6 narrower than width of base of elytra; elytra transverse, at suture as long as pronotum; abdomen arcuate laterally and strongly tapering apically. MALE. Tergite VIII irregularly truncate apically with two long, thin, sharp lateral spinelike dents (Fig. 20.177d); sternite VIII rounded apically (Fig. 20.177e); tubus of median lobe of aedeagus straight ventrally and with thin, rounded apex in lateral view (Fig. 20.177b); in dorsal view bulbus large and

382

oval, tubus narrowly triangular (Fig. 20.177c). FEMALE. Tergite VIII truncate apically (Fig. 20.177f); sternite VIII rounded apically (Fig. 20.177g); spermatheca with capsule tubular, hooked apically and stem sinuate and looped posteriorly (Fig. 20.177h). Externally, this species is superficially similar to Atheta (Datomicra) celata, A. (D.) dadopora, A. (D.) whitehorsensis and Strophogastra penicillata, but it differs in having the body more compact and robust, the teeth of male tergite VIII long, and in the shape of the median lobe of the aedeagus and spermatheca. The placement of A. hampshirensis in Dimetrota is not settled and its affiliation with Datomicra is unclear. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC. USA: AK, CA, NC, NH, NY, PA, RI. New locality data: British Columbia, Queen Charlotte Is., Tlell R., end Richardson Rd., 17. VII.1983, 83-14, 18.VII.1983, 83-18b, ex rotting Boletus, coll. J.M. Campbell (CNC) 6 females; Queen Charlotte Is., Slatechuck Mt., 14.VII.1983, 83-7, sifting Sphagnum at edge of seep., 2200 ft., coll. J.M. Campbell (CNC) 1 male, 1 female; Queen Charlotte Is., 4.7 km N Rennell Sound Rd., Ghost Cr., 19.VII.1983, 83-24, 700 ft., ex gravel along edge of stream, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Yakoun R., 26 km NW Queen Charlotte City, 21.VIII.1983, 83-101, sifting under mushrooms and moss, coll. J.M. Campbell (CNC) 1 female. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in eastern and central Canada it occurs in various coniferous, mixed, and hardwood forests, usually in decaying organic materials such as rotting fungi, dung, and carrion; in NB, specimens from black spruce forest on gilled mushrooms, jack pine forest in decaying mushrooms, from pitfall traps in red spruce forest, old-growth hemlock forest in decaying bracket fungi (polypore), various mixed forests in gilled mushrooms and decaying fleshy and gilled mushrooms in various stages of decay,

20

Tribe Athetini Casey, 1910

in decaying corncobs and cornhusks near mixed forest, alder swamp on remains of dead deer, rich Appalachian hardwood forest in fleshy fungi of various stages of decay, some from leaf litter and scat under great horned owl nest in hardwood forest; in NS, in compost, carrion, and mushrooms in coniferous and deciduous forests and open habitats. In NF, captured in carrion traps and flight intercept traps in mixed and coniferous forests. In BC, captured in rotting Boletus, in Sphagnum, and in gravel along edge of stream. Collecting period: V–IX. Collecting method: sifting fungi and litter, unbaited and carrionbaited pitfall traps, flight intercept traps. Picipennis Species Group (Klimaszewski et al. 2008a, 2018b) Diagnosis. Antennae slender, antennomeres V– X subquadrate to slightly transverse; integument of forebody highly glossy, with coarse microsculpture consisting of irregularly shaped hexagonal sculpticells; pubescence on disc of elytra directed approximately straight or obliquely posteriad; median lobe of aedeagus in dorsal view with bulbus moderate-sized, narrowly elongate, tubus long and apically triangular, in lateral view tubus arcuate, slightly sinuate or straight, with apical part slightly curved ventrad; internal sac with two medium-sized, elongate sclerites in bulbus; male tergite VIII with straight or convex and sinuate or crenulate apical margin, and with small or large lateral teeth; spermatheca very distinct, with capsule narrow, tubular, club-shaped, with small apical invagination, stem long, thin, twisted, with multiple coils. 143. Atheta (Dimetrota) picipennis (Mannerheim) (Fig. 20.178a–h) [¼A. cursor (Mäklin), ¼ A. carlottae (Casey)] Homalota picipennis Mannerheim, 1843. As Atheta: Moore and Legner 1975, Lohse and Smetana 1985, Klimaszewski and Winchester 2002, Bousquet et al. 2013.

20

Tribe Athetini Casey, 1910

Diagnosis. Body moderately convex, narrowly subparallel, length 2.9–3.5 mm, dark brown to nearly black, elytra usually paler, rust-brown with darker base near scutellum, legs and I–II basal antennomeres rust-brown or yellowish brown (Fig. 20.178a); forebody moderately sparsely punctate and pubescent, pubescence long, punctation strongly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head slightly elongate, round, narrower than pronotum, postocular region shorter than the maximum diameter of eye, tempora carinate, carinae strong throughout; antennae slender and moderately long, filiform, shorter than the pronotum and elytra combined, antennomeres VII–X subquadrate to slightly transverse, basal 4 antennomeres elongate, glossy and remaining matte; pronotum impressed mesobasally, strongly transverse, approximately rectangular in shape, hypomeron visible for two thirds of pronotal length in lateral view; pubescence directed anteriad at one third of apical midline, and obliquely laterad across arcuate lines elsewhere; elytra moderately convex, slightly elongate, and broader than pronotum, posterior margins broadly arcuate and with small and scarcely visible lateral emarginations, pubescence directed obliquely posteriad; abdomen arcuate laterally, four first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII with two rounded laterally apical teeth, margin between teeth shallowly emarginated and broadly V-shaped, and usually bearing minute irregular crenulation (Fig. 20.178d); sternite VIII rounded apically (Fig. 20.178e); median lobe of aedeagus with narrowly oval and narrowed anteriorly bulbus, and tubus long, subparallel and triangular apically in dorsal view (Fig. 20.178c); in lateral view tubus of median lobe arcuate basally and about straight apically in lateral view (Fig. 20.178b); internal sac with two pairs of large, elongate structures in bulbus (Figs. 20.178b, c). FEMALE. Tergite VIII transverse and truncate apically, slightly emarginated medially (Fig. 20.178f); sternite VIII transverse and broadly rounded

383

apically (Fig. 20.178g); spermatheca with narrowly elongate, tubular capsule slightly spherical apically and stem irregularly multiply twisted posteriorly (Fig. 20.178h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. New locality data: British Columbia, Queen Charlotte Is., Burnaby Is., N shore, Section Cove, 12.VIII.1983, 83-76, sifting moss under dung and mushrooms, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Moresby Is., Cumshewa, 14.VIII.1983, 83-82, sifting squirrel midden besides totem pole, coll. J.M. Campbell (CNC) 3 males, 3 females; Queen Charlotte Is., Moresby Is., Mt. Moresby, 26.VII.1983, 83-39, 21000 , sifting bear dung, J.M. Campbell (CNC) 23 males, 26 female, 37 sex undetermined; Queen Charlotte Is., 1 km W Moresby Camp, Pallant Cr., 3.VIII.1983, 83-50, sifting under dung in spruce forest, J.M. Campbell (CNC) 7 sex undetermined; Queen Charlotte Is., Moresby Is., 3 km NE Jedway, 12.VIII.1983, sifting moss along seepage and waterfall, J.M. Campbell (CNC) 6 sex undetermined; Queen Charlotte Is., Queen Charlotte Is., Skowkona Cr., 7.9 km NW Queen Charlotte City, 29.VII-4.VIII.1983, 83-55A, ex rotting pile of mushrooms, J.M. Campbell (CNC) 6 sex undetermined; Queen Charlotte Is., 7.9 km NW Queen Charlotte City, 16-19.VIII.1983, 83-19, dung and carrion traps, J.M. Campbell (CNC) 3 males, 3 females, 6 sex undetermined; Queen Charlotte Is., Louise Is., Skedans, 6.VIII.1983, 83-57, sifting bear dung, J.M. Campbell (CNC) 2 sex undetermined; Queen Charlotte Is., Tlell Riv., end Richardson Rd., 17.VII.1983, ex rotting Boletus, coll. J.M. Campbell (CNC) 2 males; Queen Charlotte Is., Yakoun Riv., 26 km NW Queen Charlotte City, 21.VIII.1983, sifting under mushrooms and moss, J.M. Campbell (CNC) 3 males, 3 females, 1 sex undetermined; Yakun Riv., NW corner Graham Is., 17.VIII.1983, 83-92, sifting mushrooms and dung, J.M. Campbell (CNC) 50 sex undetermined; Queen Charlotte Is., Ghost Creek Drainage, Ghost Main Road, 7.3 km NW Rennell Sound, 21.VIII.1983, 83-103, Berlese of

384

20

moss ex under deciduous shrubs around base of cedar, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., Lyall Is., Gate Cr., 10. VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, J.M. Campbell (CNC) 2males, 5 females, 2 sex undetermined; Queen Charlotte Is., 3.5 km SW Tow Hill, 22. VIII.1983, 83-110, under debris and kelp on beach, coll. J.M. Campbell (CNC) 3 males, 2 females. COLLECTION AND HABITAT DATA. Habitat: in BC, found in transition zone and forest interior of Sitka sprucer forest, under dung in spruce forest, in bear dung, moss along seepage and waterfall, rotting pile of mushrooms, under deciduous shrubs around base of cedar, and under debris and kelp on beach. Collecting period: VI–IX. Collecting method: sifting forest litter, kelp, moss, fungi, bear and martin dung. Comments. Moore and Legner (1975) recorded this species from AK, BC, NH, and southern CA. The records from NH and CA are doubtful, and should be regarded as erroneous unless proved otherwise. In BC, this species was recorded from Queen Charlotte Islands, Masset, and Carmanah Valley on Vancouver Island (Lohse and Smetana 1985, Klimaszewski and Winchester 2002). The male median lobe of aedeagus presented here is based on the type specimen from AK. We have found that there are two forms of median lobe of aedeagus, one elongate (Fig. 20.178b), and another slightly shorter and more rounded apically (Fig. 64 in Klimaszewski and Winchester 2002). We believe that the two forms represent intraspecific morphological variation. Prudhoensis Species et al. 2008a, 2018b)

Group (Klimaszewski

Diagnosis. Antennae slender, antennomeres V– X subquadrate to slightly transverse; forebody with integument moderately glossy, with moderately impressed microsculpture consisting of irregularly shaped hexagonal sculpticells, with

Tribe Athetini Casey, 1910

punctation moderately dense and asperate; pubescence on disc of elytra directed obliquely posteriad; median lobe of aedeagus in dorsal view with large, oval bulbus and moderately long and apically triangular tubus, in lateral view tubus arcuate or straight basally, with ventrally produced broad apical part; internal sac with two large, falciform sclerites in bulbus; male tergite VIII with apical margin straight to very obtusely emarginate, and with small or large lateral teeth; spermatheca S-shaped, with narrow, club-shaped capsule bearing small apical invagination, and with long, sinuate stem, narrowly looped posteriorly. 144. Atheta (Dimetrota) brumalis Casey (Fig. 20.179a–h) Atheta brumalis Casey, 1910. Hatch 1957. Moore and Legner 1975, Klimaszewski and Winchester 2002. Diagnosis. Body moderately convex, narrowly oval, length 3.0–3.5 mm, dark brown, legs yellowish rust-brown, elytra and tip of abdomen light brown and often with rust tinge, antennae usually with basal antennomere rust-brown (Fig. 20.179a); forebody densely punctate and pubescent, punctation slightly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head elongate and slightly depressed basally, slightly narrower than pronotum, postocular region elongate, shorter than the maximum diameter of eye, tempora carinate, carinae strong throughout; antennae slender and short, shorter than pronotum and elytra combined, antennomeres IV–VI subquadrate or slightly elongate, VII–X slightly transverse, basal 3 antennomeres glossy and remaining matte; pronotum slightly impressed mesobasally, transverse, approximately rectangular in shape, hypomeron visible across basal two thirds of pronotal length in lateral view, pubescence radiating from midline of disk obliquely laterad, almost horizontally laterad along broadly arcuate lines, and obliquely posteriad; elytra moderately

20

Tribe Athetini Casey, 1910

385

convex, transverse, much broader and longer than pronotum, posterior margins broadly arcuate and with small and shallow lateral emarginations, pubescence directed obliquely posteriad; abdomen subparallel, four first visible tergites impressed at base, first three strongly so; basal metatarsomere short and as long as the following one. MALE. Tergite VIII truncate apically, with two short, lateral teeth, margin between teeth either entire or slightly crenulate (Fig. 20.179d); sternite VIII rounded apically (Fig. 20.179e); tubus of median lobe of aedeagus broadly arcuate, subapical part narrowly subparallel, apex slightly projecting ventrally in lateral view (Fig. 20.179b); in dorsal view bulbus broadly oval and tubus triangular, constricted basally and rounded apically (Fig. 20.179c). FEMALE. Tergite VIII arcuate laterally and truncate medially at apical margin (Fig. 20.179f); sternite VIII broadly rounded apically (Fig. 20.179g); spermatheca with minute rounded capsule, as broad as or slightly broader than very thin stem, stem sinuate and looped posteriorly (Fig. 20.179h).

Point, 13.VIII.1983, 83-79, sifting mushrooms, J.M. Campbell (CNC) 2 males, 2 females; Queen Charlotte Is., Tlell R., end Richardson Rd., 17.VII.1983, ex rotting Boletus, 83-14, J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Lyall Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, J.M. Campbell (CNC) 2 males; Queen Charlotte Is., Port Clement, 20.VII.1983, 83-25, sifting mushrooms, J.M. Campbell (CNC) 1 male; Vancouver Is., Mt. Cain, 50.15 N, 126.25 W, 5. VIII-18.VIII.1997, 17.4 PIT 4, coll. N. Winchester (LFC) 1 male.

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA.

Comments. This species was originally described from a specimen from Metlakatla, BC.

New locality data: British Columbia, Queen Charlotte Is., Ghost Creek Drainage, Ghost Main Rd., 7.3 km NW Rennell, 21.VIII.1983, 83-103, Berlese of moss ex under deciduous shrubs around base of cedar, coll. J.M. Campbell (CNC) 3 males, 1 female; Queen Charlotte Is., Graham Is., Yaku, NW corner, 17. VIII.1983, 83-92, sifting mushrooms and dung, coll. J.M. Campbell (CNC) 5 males, 8 females; Queen Charlotte Is., Skowkona Cr., 7.9 km NW Queen Charlotte City, 29.VII-4.VIII.1983, 83-55A, ex rotting pile of mushrooms, coll. J.M. Campbell (CNC) 3 males, 2 females; Queen Charlotte Is., Burnaby Is., N shore, Section Cove, 12.VIII.1983, 83-76, moss under dung and mushrooms, coll. J.M. Campbell (CNC) 1 male, 1 female; same data as before except 11. VIII.1983, 83-75, sifting moss and mushrooms (CNC) 3 males, 2 females; Queen Charlotte Is., Moresby Is., Haswell Bay, 1.5 km SW Hoskins

145. Atheta (Dimetrota) Klimaszewski (Fig. 20.180a–h)

COLLECTION AND HABITAT DATA. Habitat: the BC, specimens were found in transition zone and the forest interior of Sitka spruce forest, and in rotting mushrooms, moss under deciduous shrubs around base of cedar, and from moss under dung. Collecting period: VI– X (most abundant in VIII-IX). Collecting method: pitfall traps, sifting mushrooms and moss and dung.

capsularis

Atheta capsularis Klimaszewski, in Klimaszewski et al. 2005. Klimaszewski et al. 2008a, 2011, 2018b, Work et al. 2013. Diagnosis. Body narrowly oval, moderately convex, length 2.8–3.2 mm; colour dark brown to black, with legs/tarsi, basal parts of antennae, elytra, and basal and apical parts of abdomen yellowish-brown, some specimens entirely yellowish brown (Fig. 20.180a); integument moderately glossy; forebody with punctation dense and asperate, with fine meshed microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad from midline, forming arcuate pattern, that on elytra directed obliquely posteriad, forming wavy pattern laterally; head slightly narrower than

386

pronotum; antennae slender, antennomeres V–X subquadrate; pronotum transverse, about 1/5 narrower than width of base of elytra; elytra transverse, at suture as long as pronotum; abdomen subparallel. MALE. Apical margin of tergite VIII angulately produced laterally, apex with broad, shallow, slightly sinuate medial emargination (Fig. 20.180d); sternite VIII parabolic apically (Fig. 20.180e); tubus of median lobe of aedeagus straight ventrally and with broad, rounded apical part produced slightly ventrally in lateral view (Fig. 20.180b); in dorsal view bulbus large, narrowly oval and broadest in basal third, tubus triangular, apex narrowly rounded (Fig. 20.180c). FEMALE. Tergite VIII truncate apically (Fig. 20.180f); sternite VIII rounded apically (Fig. 20.180g); spermatheca with club-shaped capsule with narrow apical invagination facing laterad, stem sinuate and with posterior loop (Fig. 20.180h). This species may be confused externally with Atheta terranovae, but it differs in being generally larger, in having the elytra yellowish and the body paler, in the shape and structures of the median lobe of the aedeagus, and in the unique shapes of the spermatheca and male tergite VIII. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], LB, NB, NF, ON, QC, YT. USA: not recorded. We report this species from BC for the first time here based on the following: British Columbia, 13 km N of Topley, VIII-IX.2012, pitfall # 389, 54.616-126.3, 1132 m, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 male, 1 female; Vancouver Is., Mt. Cain, 50.13 N, 126.22 W, 7.VII-20. VII.1997, 16.6 PIT 4, coll. N. Winchester (LFC) 1 male; Mt. Cain, 50.13 N, 126.22 W, 27.VII-10. VIII.1996, 16.2 P 1, coll. N. Winchester (LFC) 1 female; Mt. Cain, 50.13 N, 126.22 W, 5.VIII18.VIII.1997, pitfall traps 1, coll. N. Winchester (LFC, RBCM) 8 males, 3 females; Mt. Cain, 50.13 N, 126.22 W, 1.IX-19.IX.1997, pitfall traps, coll. N. Winchester (LFC, RBCM) 3 males, 1 female; Mt. Cain, 50.13 N, 126.22 W, 14.IX-28.IX.1996, Pan traps,

20

Tribe Athetini Casey, 1910

coll. N. Winchester (LFC, CNC) 2 males, 1 female; Mt. Cain, 50.13 N, 126.22 W, 28.IX12.X.1996, GMT, coll. N. Winchester (LFC, CNC) 2 females. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: red spruce, balsam fir, jack pine, mixed and riparian type forests. Most specimens were from pitfall traps and flight intercept traps; some from forest litter. This species feeds on several species of yeasts and fungal spores based on contents in the posterior gut. In BC, two specimens were captured in sub-boreal spruce forest. Collecting period: V– IX. Collecting method: sifting litter, pitfall traps, flight intercept traps, Pan traps. Comments. Externally and genitalically Atheta capsularis is similar to A. terranovae. The two species can be easily confused but A. capsularis differs in having distinctly transverse antennomeres VII-X, sinuate V-shaped emargination of male tergite VIII, broader apex of median lobe of aedeagus in lateral view, and in a distinct shape of spermatheca. 146. Atheta (Dimetrota) prudhoensis (Lohse) (Fig. 20.181a–h) Dimetrota prudhoensis Lohse, in Lohse et al. 1990. Gusarov 2003a, Klimaszewski et al. 2007b, 2008a, 2011, 2015c, 2018b, Webster et al. 2009, Majka and Klimaszewski 2008b. Diagnosis. Body narrow, subparallel, slightly flattened, length 3.3–3.5 mm; colour dark brown to black, with legs, antennae, elytra, and posterior part of abdomen yellowish brown or reddish brown (Fig. 20.181a); integument moderately glossy; forebody with punctation dense and asperate, with fine microsculpture; pubescence on disc of pronotum directed laterad and obliquely posteriad from midline, that on elytra directed obliquely and straight posteriad; head slightly narrower than pronotum; antennae slender, antennomeres V–X subquadrate to slightly transverse; pronotum transverse, about 1/5 narrower than width of base of elytra; elytra

20

Tribe Athetini Casey, 1910

transverse, at suture as long as pronotum; abdomen arcuate laterally. MALE. Tergite VIII with two small lateral teeth and broadly V-shaped apical emargination (Fig. 20.181d); sternite VIII parabolic apically (Fig. 20.181e); tubus of median lobe of aedeagus arcuate ventrally and with evenly moderately broad apical part, apex rounded and produced ventrally in lateral view (Fig. 20.181b); in dorsal view bulbus large and oval, tubus triangular apically (Fig. 20.181c). FEMALE. Tergite VIII truncate apically (Fig. 20.181f); sternite VIII rounded apically (Fig. 20.181g); spermatheca S-shaped, with tubular, clubbed-shaped capsule, and sinuate stem narrowly looped posteriorly (Fig. 20.181h). Atheta prudhoensis is closely related to A. burvelli, A. curvipennis, and possibly A. caribou, from which it differs in the shape of the male genitalia and in some external features. Females of this species may be confused with those of A. burwelli, A. curvipennis, and A. terranovae, but may be distinguished with certainty by association with males, which are more easily identifiable. DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NF, NS, ON, QC, SK, YT. USA: AK, NY, VT. Additional locality and collection data: British Columbia, Queen Charlotte Is., Moresby Is., 26. VIII.1983, 83-39, 21000 , sifting bear dung, coll. J.M. Campbell (CNC) 5 males, 1 female; Queen Charlotte Is., Burnaby Is., N shore, Section Cove, 11.VIII.1983, 83-75, sifting moss and mushrooms, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Ghost Main Rd., 7.3 km NW Rennell, 21.VIII.1983, 83-103, Berlese of moss ex under deciduous shrubs around base of cedar, coll. J.M. Campbell (CNC) 1 female; Vancouver Is., Mt. Cain, 50.13 N, 126.21 W, 18.VIII-1. IX.1997, 16.6 PIT 8, N. Winchester (LFC) 2 males. COLLECTION AND HABITAT DATA. Habitat: in many kinds of forests such as conifer dominated forests, spruce, white spruce, spruce

387

and fir, red pine, jack pine, birch, maple, oak, northern hardwood, hemlock, and mixed forests. Adults often common in decaying mushrooms and Russula sp. (gilled mushroom) but also were found in deer dung, carrion, compost, moss, and leaf litter. Collecting period: VI–X. Collecting method: sifting litter and mushrooms, pitfall traps, flight intercept traps. 147. Atheta (Dimetrota) rurigena Casey (Fig. 20.182a–e) Atheta (Dimetrota) rurigena Casey, 1911. Holotype (male): British Columbia, Yale, Wickham, rurigena Casey; Type USNM 39282; Casey bequest 1925 (USNM). Body missing except for abdomen and genitalia. Examined. Diagnosis (based on Casey’s original description and damaged holotype). Body subparallel, length 2.7 mm, punctation fine and sparse; head large, transverse, fourth-fifth as wide as pronotum, tempora prominent, parallel and abruptly oblique near base; eyes large; antennae, long, filiform, moderately thick, antennomeres IV–X slightly elongate; pronotum two-fifth wider than long; elytra short, strongly transverse, at suture scarcely a fifth longer than pronotum; abdomen broad with sides arcuate (Fig. 20.182a); basal metatarsomere as long as second metatarsomere (Fig. 20.182a). MALE. Tergite VIII sinuate apically, with shallow median emargination and without teeth (Fig. 20.182d); sternite VIII parabolic, rounded apically (Fig. 20.182e); tubus of median lobe of aedeagus straight basomedially, subapical part broad, apex rounded in lateral view (Fig. 20.182b), in dorsal view bulbus large, oval, and tubus moderately short, subparallel, broad and rounded at apex (Fig. 20.182c); internal sac with two long and broad sclerites in bulbus (Fig. 20.182b, c). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

388

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Tribe Athetini Casey, 1910

COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded.

DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NF, ON, QC, SK, YT. USA: AK, NY.

Comments. This species is known only from the male holotype with missing forebody. In the original description, Casey misidentified the sex of the holotype as a female because it was lacking lateral teeth present in most of the males in this group.

New locality data: British Columbia, Hwy 16 rest stop at Cluculz Creek, 53 54.52N, 123 35.220 W, 31.VII.2012, 776 m, in Salix and Alnus litter with moss, A. Davies (CNC) 1 female.

148. Atheta (Dimetrota) strigosula Casey (Fig. 20.183a–i) Atheta (Atheta) strigosula Casey, 1910. Klimaszewski et al. 2005, 2007b, 2008a, 2011, 2016d, 2018b, Majka and Klimaszewski 2008b; Brunke et al. 2012, Stefani et al. 2016. Diagnosis. Body broadly subparallel, length 3.6–4.0 mm, head and pronotum flattened medially; colour dark brown, with elytra, legs and bases of antennae light yellowish-brown or reddish-brown (Fig. 20.183a); integument moderately glossy, forebody with asperate punctation and meshed microsculpture, pubescence fine and sparse, on elytra denser; head slightly narrower than pronotum, subquadrate, disc impressed medially; antennae slender, with antennomeres IV–V slightly elongate, VI–X subquadrate to slightly transverse; pronotum distinctly transverse, widest at middle, slightly narrower than elytra (wider in male than in female); elytra at suture slightly longer than pronotum; abdomen widely arcuate laterally. MALE. Tergite VIII truncate apically, with two obtuse lateral teeth and crenulate posterior margin (Fig. 20.183d); sternite VIII parabolic apically (Fig. 20.183e); tubus of median lobe of aedeagus with broad and triangular apical part in lateral view (Fig. 20.183b), in dorsal view bulbus large, oval, and tubus moderately long and narrowly triangularly shaped (Fig. 20.183c). FEMALE. Tergite VIII truncate apically (Fig. 20.183f); sternite VIII rounded apically (Fig. 20.183g); spermatheca minute, capsule club-shaped, stem thin and sinuate (Fig. 20.183h, i).

COLLECTION AND HABITAT DATA. Habitat: occurs in various forest types such as red spruce, mature red spruce and eastern white cedar, yellow birch and balsam fir, mixed, red oak, rich Appalachian hardwood, northern hardwood, and other deciduous forests. Adults usually found in decaying mushrooms (often common) on forest floor but also in leaf litter and possibly carrion. In BC, one female was captured in Salix and Alnus litter with moss. Collecting period: VI–X (most common in IX in NB). Collecting method: sifting mushrooms and litter, carrionbaited and unbaited pitfall traps, flight intercept traps. 149. Atheta (Dimetrota) terranovae Klimaszewski and Langor (Fig. 20.184a–h) Atheta (Dimetrota) terranovae Klimaszewski and Langor, in Klimaszewski et al. 2011. Brunke et al. 2012, Webster et al. 2012, Klimaszewski et al. 2012, 2016d, 2018b, Work et al. 2013. Diagnosis. Body narrowly subparallel, moderately convex, length 2.5–3.0 mm; head, pronotum, and entire/or posterior of abdomen dark brown to nearly black, with appendages and elytra paler, yellowish brown mottled with dark brown (Fig. 20.184a); integument moderately glossy; forebody with punctation dense and asperate, with microsculpture moderately coarse; pubescence on disc of pronotum directed laterad from midline of disc, that on elytra directed obliquely posteriad; head distinctly narrower than pronotum; antennae slender, antennomeres V–X slightly transverse; pronotum transverse, about 1/8 narrower than width of base

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Tribe Athetini Casey, 1910

of elytra; elytra transverse, at suture at least as long as pronotum; abdomen subparallel. MALE. Tergite VIII with entire apical margin shallowly, obtusely angulately emarginate, lateral angles rectangular (Fig. 20.184d); sternite VIII parabolically rounded from base to apex (Fig. 20.184e); tubus of median lobe of aedeagus slightly produced ventrally at apex, with apical part narrowly elongate in lateral view (Fig. 20.184b); in dorsal view bulbus narrowly oval, tubus broad, triangular apically (Fig. 20.184c). FEMALE. Tergite VIII truncate apically (Fig. 20.184f); sternite VIII rounded apically (Fig. 20.184g); spermatheca S-shaped, with club-shaped capsule and stem sinuate, narrowly looped posteriorly (Fig. 20.184h). This species bears some external resemblance to Atheta capsularis and A. prudhoensis, but in A. terranovae the body is slightly smaller and narrower, the pronotum is less transverse, the punctation are denser, antennomeres VII–X are more transverse, the elytra are darker, and the shapes of the median lobe of the aedeagus and spermatheca are different. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], LB, NB, NF, ON, QC, SK, YT. USA: AK [new state record]. We report this species from BC and AK for the first time here based on the following: British Columbia, Log Lake, 65 km N of Prince George, 54.2-122.616, 780 m, VIII-IX.2012, pitfall # 208, OM3C1SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 male, 1 female; 13 km N Topley, 54.616-126.3, 1132 m, VIIIIX.2012, pitfall # 389, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 3 males, 7 females; Kootenay, Porcupine forest service road, 49.2646-117.1815, 820 m, 12.IX.2011, Berlese, moss, R.G.B. Bennett and D. &C. Copley (RBCM) 1 female; BC, Vancouver Is., Mt. Cain, 50.13 N, 126.22 W, 5.VIII-18. VIII.1997, 16.2 PIT9, N. Winchester (LFC) 1 male; Mt. Cain, 50.13 N, 126.21 W, 10.VIII23.VIII.1996, 16.6 PAN 2, N. Winchester (LFC) 2 males; Mt. Cain, 50.13 N, 126.21 W, 18.VIII-

389

1.IX.1997, 16.6 PIT 8, Winchester (LFC) 2 males, 2 female; Mt. Cain, 50.13 N, 126.22 W, 23.VIII-14.IX.1997, 16.2 PAN 2, Winchester (LFC) 1 male, 1 female; Mt. Cain, 50.13 N, 126.22 W, 1.IX-19.IX.1997, 16.2 PIT 2, 6 & 7, N. Winchester (LFC) 1 male, 2 female. USA, Alaska, Eureka Summit, 1000 m, Hwy 1, 5.VIII.1988, leg. R. Baranowski (ZMLU) 5 males, 2 females; near Malta-Muska Glacier, Hwy 1, 4.VIII.1988, leg. R. Baranowski (ZMLU) 1 male; 8-16 miles E of Willow, 7.VIII.1988, leg. R. Baranowski (ZMLU) 1 male. COLLECTION AND HABITAT DATA. Habitat: occurs in many coniferous, mixed and hardwood/deciduous forest types in Canada. Adults commonly found in decaying mushrooms (polypore and gilled) and sometimes in fresh mushrooms (Klimaszewski et al. 2018a). In BC, adults were found in sub-boreal spruce, in Sitka spruce forest and in moss. In NB, in an oak forest, rich Appalachian hardwood forest, northern hardwood forest, various mixed forests, red spruce forest, mature red spruce forest with red maple or yellow birch, black spruce forest, and eastern white cedar forests and swamps; specimens in fresh and decaying gilled mushrooms of various species, rotting lobster mushrooms, and a coral mushroom on spruce log. Also captured in unbaited and carrion-baited pitfall traps and flight intercept traps in various coniferous (sprucefeathermoss, spruce-lichen forests), mixed (spruce/aspen, riparian), and deciduous forests in NF and LB including some from decaying mushrooms; also from red maple, pine/willow, balsam fir/white birch forests, elsewhere. Collecting period: VI–X (most common in VIII and IX). Collecting method: sifting litter and mushrooms, carrion-baited and unbaited pitfall traps, flight intercept traps, Lindgren funnel traps. (Subgenus Hydrosmectomorpha Klimaszewski and Webster) (Fig. 20.185a–i) (Klimaszewski et al. 2018a, b, 2019) Diagnosis. Hydrosmectomorpha is distinguishable from other subgenera of Atheta by the

390

following combination of characters: body narrow, subparallel and flattened; elytra markedly broader than remainder of body; distinctive pattern of pubescence on pronotum and elytra (described above); antennomeres VIII–X distinctly to slightly elongate; ligula weakly subdivided apically; legs long; apical margin of male tergite VIII lacking prominent teeth; spermatheca small, with stem short and simple, straight or sinuate; all known species occur among cobblestones and gravel along margins of clear, fast flowing rivers, near the outflow of brooks into rivers, and along lake margins with wave action. The median lobe of the aedeagus in Hydrosmecta has a blade-like, triangular projection at the base of the tubus in lateral view to which parameres are attached, this structure is narrowly elongate along ventrally flat part of bulbus, in Hydrosmectomorpha. Male tergite VIII in Hydrosmecta is truncate apically and not modified, while in Hydrosmectomorpha it is truncate and slightly emarginate, deeply notched, or with minute teeth. The stem of the spermatheca in Hydrosmecta is coiled posteriorly, broadly looped, or irregularly twisted, while in Hydrosmectomorpha it is straight or slightly sinuate with small apical swelling. The external similarity of Hydrosmectomorpha species (e.g., flattened body, similar pronotal pubescence pattern) to those of Hydrosmecta represent convergent evolution and adaptation to live among cobblestones, gravel and sand along margins of clear, fast-flowing water. 150. Atheta (Hydrosmectomorpha) meduxnekeagensis Webster and Klimaszewski (Fig. 20.185a–i) Atheta (Hydrosmectomorpha) meduxnekeagensis Webster and Klimaszewski, in Klimaszewski et al. 2018a. Diagnosis. Body subparallel, flattened, length 2.6–3.0 mm (mean ¼ 2.8 mm (N ¼ 10), NB specimens); colour dark brown to almost black, with legs and elytra except for scutellar region paler, reddish- or yellowish-brown

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Tribe Athetini Casey, 1910

(Fig. 20.185a); integument slightly glossy, forebody with fine, moderately dense punctation and faint meshed microsculpture; head slightly narrower than pronotum, eyes large and slightly protruding; antennae moderately robust, antennomeres I–VII highly elongate, VIII–X slightly to distinctly elongate; pronotum widest at apical third (0.51 mm); elytra distinctly elongate and broader than pronotum (0.61 mm, at shoulders), flattened, at suture about as long as pronotum; abdomen subparallel, broadest before middle, gradually tapering apicad. MALE. Apical margin of tergite VIII slightly (NB specimens, Fig. 20.185d) to distinctly (BC specimens, Fig. 20.185e) sinuate, with only one small lateral denticle on each side (Fig. 20.185d, e); apical margin of sternite VIII moderately produced, rounded (Fig. 20.185f); in lateral view tubus of median lobe of aedeagus moderately wide, ventral margin arcuate in basal half, narrow and slightly curved ventrad apically (Fig. 20.185b BC, c NB). FEMALE. Apical margin of tergite VIII in NB specimens shallowly, broadly emarginate, angular near lateral margin (Fig. 20.185g), more produced, lobe-like laterally in BC specimens (Fig. 20.185h); sternite VIII broadly arcuate apically (Fig. 20.185i); spermatheca short, capsule narrowly tubular and club-shaped, apical invagination small and shallow, stem straight to base with small globular swelling (Fig. 20.185j). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018a). Habitat: In NB, this species occurs in various cobblestone habitats along river and lake margins. Adults usually occurred among cobblestones and gravel near water’s edge, often in areas with slowed current along fast flowing rivers where there was a buildup of fine debris and growth of algae on cobblestones. This species was also found along lake margins where wave action has created conditions similar to the flowing water of rivers. BC specimens were collected from a river margin

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Tribe Athetini Casey, 1910

391

among cobblestones and gravel. Collecting period: VI–VIII. Collecting method: Hand collecting from among cobblestones near river and lake margins.

aedeagus straight along entire length in lateral view (Fig. 20.187b).... Atheta (Lamiota) relicata Casey

(Subgenus Lamiota Casey, 1910) (Figs. 20.186a–h and 20.187a–e) (Casey 1910; Seevers 1978; Gusarov 2003a; Klimaszewski and Winchester 2002)

151. Atheta (Lamiota) (Fig. 20.186a–h)

Diagnosis. Large species, length 3.0–4.0 mm; body subparallel, flattened, broad, broadest at elytra; pronotum moderately densely punctate, punctation slightly asperate, pubescence on midline of disc directed posteriad basally and anteriad in apical half, on sides pubescence directed laterad and obliquely posteriad, several macrosetae distributed on lateral margins; antennomeres IV– V elongate or slightly transverse, VI subquadrate, slightly elongate, or transverse, VII–X subquadrate to slightly transverse; abdomen broad, arcuate laterally; male tergite VIII with or without teeth; apical part of internal sac of median lobe of aedeagus with two narrow, diverging apically, lateral projections; subapical part of median lobe rod-shaped in lateral view; spermatheca with small spherical capsule and stem thin, very long, sinuate, and narrowly looped posteriorly. Key to Species of Subgenus Atheta (Lamiota) 1. Antennomere IV slightly elongate, VII–X subquadrate to slightly transverse (Fig. 20.186a); elytra at suture slightly shorter than pronotum along midline (Fig. 20.186a); male tergite VIII with two large, curved lateral teeth and projected median part of apical margin, edge with some crenulation (Fig. 20.186d); tubus of median lobe of aedeagus straight basally and produced ventrad apically in lateral view (Fig. 20.186b)......Atheta (Lamiota) keeni Casey ............................................................................2 – Antennomere IV subquadrate, VII–X distinctly transverse (Fig. 20.187a); elytra at suture slightly longer than pronotum along midline (Fig. 20.187a); male tergite VIII truncate apically with two small lateral and two minute median teeth (Fig. 20.187d); tubus of median lobe of

keeni

Casey

Atheta (Lamiota) keeni Casey, 1910. Gusarov 2003a [synonymy], Klimaszewski and Winchester 2002 [as Atheta (Liogluta) vasta]. Diagnosis. Body moderately convex, broad, subparallel, length 3.7–4.0 mm, dark brown to nearly black, elytra, apical part of abdomen, and legs red-brown, antennae uniformly dark brown with antennomeres I–III paler (Fig. 20.186a); forebody moderately densely punctate and pubescent, pubescence long, punctation slightly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head elongate, round posteriorly, distinctly narrower than pronotum, postocular region shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; antennae slender, as long as pronotum and elytra combined, antennomere IV slightly elongate, VII–X subquadrate to slightly transverse, basal 4 antennomeres glossy and remaining matte; pronotum impressed mesobasally, transverse, approximately rectangular in shape, hypomeron visible for two thirds of pronotal length in lateral view, pubescence directed from midline of disc anteriad at apical third, on sides nearly horizontally laterad medially, and obliquely laterad posteriorly; elytra flattened, distinctly transverse, broader than pronotum, at suture as long as pronotum along midline, posterior margins feebly sinuate and with shallow, scarcely visible, lateral emarginations, pubescence directed straight or slightly obliquely posteriad; abdomen subparallel, strongly glossy, four first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII with two distinct arcuate lateral teeth and margin between teeth slightly produced and crenulate (Fig. 20.186d); sternite VIII parabolic apically

392

(Fig. 20.186e); bulbus of median lobe of aedeagus narrowly oval, tubus with two narrow lateral projections in dorsal view (Fig. 20.186c); tubus with ventral margin about straight basally, narrowly elongate apically and produced ventrad in lateral view (Fig. 20.186b). FEMALE. Tergite VIII broadly truncate and slightly emarginated medially (Fig. 20.186f): sternite VIII broadly rounded apically, with row of distinct setae on apical margin (Fig. 20.186g); spermatheca with capsule small, hemispherical apically and with elbowed neck posteriorly, stem thin, long, broadly U-shaped medially, and narrowly looped posteriorly with a single small coil at apex (Fig. 20.186h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, OR. New locality data: British Columbia, Queen Charlotte Is., 7.9 km NW Queen Charlotte City, 16-19.VII.1983, 83-19, dung and carrion traps, J.M. Campbell (CNC) 1 male, 3 females; Queen Charlotte Is., 1.4 km S Masset, 13.VII.1983, 83-3, sifting Alnus litter, J.M. Campbell (CNC) 1 male; Vancouver Is., Goldstream Park, 27. V.1968, B), A. Smetana (CNC) 1 male, 1 female; BCol., Vancouver Is., 3 km SW Brentwood Bay, 48 330 1200 N, 123 290 3600 W, 6.X.2011, moss, fern, and Amelanchier litter at edge of upland pond, A. Davies (CNC) 1 female; BCol., Kitsumkalum Lake at Hwy. 13, S of Rosswood, 54 47.340 N, 128 45.970 W, 30.VII.2012, 146 m, in moss and leaf litter at creek, A. Davies (CNC) 3 females COLLECTION AND HABITAT DATA. Habitat: in AK, found in forest litter, Alnus, Veratrum, ferns. In BC, found in transition zone of Sitka spruce, in dung and carrion, in Alnus litter, in moss, fern, and Amelanchier litter at edge of upland pond, and in in moss and leaf litter near creek. Collecting period: VII. Collecting method: sifting Alnus litter, moss and leaf litter, moss, fern, and Amelanchier litter; dung and carrion baited pitfall traps.

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Tribe Athetini Casey, 1910

Comments. This species was originally described from a specimen from Metlakatla, BC. 152. Atheta (Lamiota) relicata Casey [new subgeneric combination] (Fig. 20.187a–e) Atheta relicata Casey, 1911. Moore and Legner 1975. Lectotype (male): Canada, British Columbia: Metlakatla, B. Col., Keen; relicata—3, PARATYPE USNM 39272; Casey bequest 1935; lectotypus Atheta relicata Casey, V.I. Gusarov des. 2000. Our lectotype designation label (USNM). Present designation. Diagnosis (based on male lectotype). Body length 3.4 mm, head, pronotum and posterior abdomen dark brown, elytra, bases of antennae and legs yellow brown (Fig. 20.187a); microsculpture of forebody strong, sculpticells hexagonal, punctation moderately dense; head distinctly narrower than pronotum, postocular area shorter than diameter of eye seen from above; eyes large; antennomere IV subquadrate, VII–X moderately transverse; pronotum transverse, evenly arcuate laterally, truncate basal and slightly sinuate laterobasally, pubescence on midline directed mostly anteriad and on side laterad and obliquely laterad; elytra subparallel, at suture slightly longer than pronotum along midline; abdomen slightly arcuate laterally. MALE. Tergite VIII truncate apically with two small lateral and two minute median teeth (Fig. 20.187d); sternite VIII parabolic apically (Fig. 20.187e); tubus of median lobe of aedeagus narrow, rod-like, almost straight and rounded apically in lateral view (Fig. 20.187b), in dorsal view tubus has two narrow lateral diverging projections, characteristic for this subgenus (Fig. 20.187c); bulbus large; internal sac as illustrated (Fig. 20.187b, c). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

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Tribe Athetini Casey, 1910

393

COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. This species is known only from the male holotype. (Subgenus Metadimetrota Klimaszewski and Winchester, 2002) (Fig. 20.188a–h) (Klimaszewski and Winchester 2002) Diagnosis. Body narrowly elongate, broadest at elytra; pronotum sparsely punctate, punctation slightly asperate, pubescence on disc directed anteriad along most of midline, and apicolaterad along arcuate lines elsewhere, several macrosetae distributed on lateral margins; infraorbital carina present but diffused near eye; antennomeres IV– V elongate, VI quadrate to slightly elongate, VII– X slightly transverse; labrum transverse, shallowly emarginate medially; mandibles broad, median tooth of right mandible strong; maxillae with galeae extending slightly beyond apex of laciniae, maxillary palpi with four palpomeres, last one narrowly elongate; labial palpi with three palpomeres; ligula small, split apically to about half length; pronotal hypomera visible laterally for almost its entire length; process of mesoventrite narrow and acutely pointed, mesocoxae narrowly separated, contiguous at middle; elytra with posterior margins broadly arcuate and with scarcely visible latero-posterior emarginations, pubescence on disc directed obliquely posterolaterad; abdomen with strong, long, arcuate macrosetae particularly at apex; male tergite VIII entire apically and without lateral teeth; male sternite VIII bearing extremely long setae apically, which are less pronounced in female; tubus of the median lobe of aedeagus bearing lateral carinae. 153. Atheta (Metadimetrota) Klimaszewski (Fig. 20.188a–h)

Diagnosis. Body slightly flattened, length 3.0–3.5 mm, head, apical antennomeres, and median abdominal tergites dark brown, remainder of the body yellowish brown or entire body dark brown with paler pronotum and elytra (Fig. 20.188a); integument moderately glossy, especially on posterior of abdomen; forebody with punctation and pubescence moderately dense, punctation slightly asperate, microsculpture distinct, consisting of convex isodiametric sculpticells; head transverse, angular, slightly narrower than pronotum; pronotum transverse, about one fourth broader than long, broadest in apical third, only slightly broader than head and about 1/5 narrower than width of elytra, pubescence on disc directed laterad from midline; elytra transverse, at suture slightly longer than pronotum; abdomen narrowly elongate and tapering apically, bearing strong and long apico-ventral and lateral macrosetae in males. MALE. Tergite VIII rounded apically, antecostal suture arcuate, close to basal margin (Fig. 20.188d); sternite VIII produced apically and with extremely long, coarse macrosetae, antecostal suture arcuate (Fig. 20.188e); median lobe of aedeagus with narrow apex and angular base in lateral view (Fig. 20.188b), in dorsal view tubus narrowly triangular with angular lateral projections on each side (Fig. 20.188c). FEMALE. Tergite VIII truncate apically with shallow median emargination, antecostal suture sinuate (Fig. 20.188f); sternite VIII rounded apically and slightly produced medially, antecostal suture sinuate (Fig. 20.188g); spermatheca with capsule elongate and spherical apically, with a deep apical invagination, stem broadly looped and twisted posteriorly, end part swollen (Fig. 20.188h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

cheersae

Atheta (Metadimetrota) cheersae Klimaszewski, in Klimaszewski and Winchester, 2002.

New locality data: British Columbia, BColl., 13 km SW Parksville, Englishman River Falls, 20.X.2008, old-growth Douglass fir and maple litter, A. Davies (CNC) 4 males, 3 females, 2 sex undetermined; BColl., Duncan,

394

Mt. Tzouhalem, 48 470 N, 123 370 W, 19.X.2008, moss and maple litter by small pond in secondgrowth Douglas fir forest, A. Davies (CNC) 1 female. COLLECTION AND HABITAT DATA. Habitat: Sitka spruce forest, forest interior and transition zone, Carmanah Valley, and old-growth Douglas fir and maple forest litter, and in moss and maple litter by small pond in second-growth Douglas fir forest. Collecting period: IX–XI. Collecting method: sifting forest litter, pitfall traps and Malaise traps. Comments. There is an additional species of this subgenus, A. savardae Klimaszewski, occurring in eastern Canada (Klimaszewski et al. 2018) (Subgenus Microdota Mulsant and Rey, 1873a) (Figs. 20.189–20.194) (Klimaszewski et al. 2015c, 2018b; Lee and Ahn 2015) Diagnosis. Body subparallel, length 1.5–2.5 mm; antennomere I enlarged but not extremely swollen, longer than II, II longer than III, V–X slightly to distinctly transverse; median region of prementum very narrow, without pseudopores; labial palpi with three palpomeres; glossae split apically forming Y-shaped structure; maxillary palpi with three palpomeres, last one narrowly elongate; pronotum transverse, more than 1.2 times as wide as long, pubescence on disc directed anteriad along midline in most specimens, and laterad elsewhere; pronotal hypomera fully visible laterally; elytra in some species with pubescence forming wavy pattern in postero-sutural area of disc; abdomen subparallel; male tergite VIII in most species simple, truncate apically, without teeth, sometimes with minute crenulation and pairs of small teeth; median lobe of aedeagus in dorsal view with large bulbus, tubus triangularly shaped apically, internal sac with well-developed complex structures; spermatheca of variable shape, L- or S-shaped, capsule usually narrowly sac-shaped, or club-shaped, stem often sinuate. Some species of Microdota, due to small body size and some superficial external similarity, may

20

Tribe Athetini Casey, 1910

be confused in collections with members of the subgenus Datomicra Mulsant and Rey, from which they differ in having the pronotal hypomera fully exposed in lateral view, while they are only partially visible in Datomicra. Many species of Datomicra also have the forebody more densely and coarsely punctate than that of Microdota. Microdota may be distinguished from Dimetrota by the following combination of characters: body usually parallel-sided, on average 2 mm long (1.5–2.8 mm in Microdota; 1.8–3.8 mm in Dimetrota), glossae Y-shaped (deeply split in Dimetrota); pronotum with sparse to moderately dense and slightly asperate punctation (dense and coarsely asperate in Dimetrota); lateral margins of pronotum and elytra, and middle and hind tibiae with moderately coarse macrosetae (strong bristles in Dimetrota); hypomera fully visible in lateral view (partially to less often fully visible in Dimetrota); and male tergite VIII truncate apically, rarely crenulate, and usually without large lateral teeth (with two large lateral teeth and margin between them often distinctive or with pattern of smaller teeth in Dimetrota). Details on diagnostics of Microdota are provided by Brundin (1948) and Lee and Ahn (2015). Species of Nearctic Microdota need revision. Key to Species of Subgenus Atheta (Microdota) 1. Elytra at base slightly wider than maximum width of pronotum (Figs. 20.189a and 20.191a) ............................................................................2 – Elytra at base distinctly wider than maximum width of pronotum (Figs. 20.190a and 20.192a– 20.194a) .............................................................3 2. Antennomeres VII–X each about twice as wide as long (Fig. 20.189a); genae angular and longer than length of eye visible from above (Fig. 20.189a); median lobe of aedeagus and spermathecal as illustrated (Fig. 20.189b, g) ....... ......... Atheta (Microdota) copleyi Klimaszewski – Antennomeres VII–X each less than twice as wide as long (Fig. 20.191a); genae rounded and shorter than length of eye visible from above

20

Tribe Athetini Casey, 1910

(Fig. 20.191a); median lobe of aedeagus and spermathecal as illustrated (Fig. 20.191a, h) ....... ............. Atheta (Microdota) lucifera Bernhauer 3. Pronotal punctures fine, antennomeres VII–X slightly transverse (Fig. 20.190a); median lobe of aedeagus and spermathecal as illustrated (Fig. 20.190b, c, h)............................................... ......... Atheta (Microdota) holmbergi Bernhauer – Pronotal punctures course, antennomeres VII– X strongly transverse (Figs. 20.192a, 20.193a and 20.194a); median lobe of aedeagus and spermathecal not as above .................................4 4. Eyes not protruding from outline of head in dorsal view, each eye slightly longer than postocular region of head (Fig. 20.193a); pubescence of sutural region of elytra directed obliquely posteriad (Fig. 20.193a); median lobe of aedeagus and spermathecal as illustrated (Fig. 20.193b, c, h)............................................... ................Atheta (Microdota) ripariides Newton – Eyes protruding from outline of head in dorsal view, distinctly longer than postocular region of head (Figs. 20.192a and 20.194a); pubescence of sutural region of elytra forming wavy pattern (Figs. 20.192a and 20.194a); median lobe of aedeagus and spermathecal not as above (Figs. 20.192b, c, h and 20.194b, c) ..................5 5. Antennomeres VII–X strongly transverse (Fig. 20.192a); elytra slightly broader than base of pronotum (Fig. 20.192a); ventral margin of tubus of median lobe of aedeagus strongly sinuate medially in lateral view (Fig. 20.192b)................ .............. Atheta (Microdota) platonoffi Brundin – Antennomeres VII–X subquadrate or slightly transverse (Fig. 20.194a); elytra distinctly broader than base of pronotum (Fig. 20.194a); ventral margin of tubus of median lobe of aedeagus almost straight in lateral view (Fig. 20.194b)....................................................... ...................Atheta (Microdota) surgens (Casey)

154. Atheta (Microdota) copleyi Klimaszewski, sp. n. (Fig. 20.189a–g)

395

Holotype (male): Canada, British Columbia, Yoho National Park, Takakkaw falls, 51.4858116.4766, 1334 m, 9.VIII.2013, moss, Berlese, coll. R.G.B. Bennett and D.C. Copley (RBCM) 1 male. Paratypes: British Columbia, Yoho National Park, Takakkaw falls, 51.4858116.4766, 1334 m, 9.VIII.2013, moss, Berlese, coll. R.G.B. Bennett and D.C. Copley (RBCM) 4 females. Etymology. This species is dedicated to Mr. Darren C. Copley of RBCM, British Columbia, who collected the type series and many other specimens used in this study. Diagnosis. Body moderately convex, narrowly subparallel, strongly glossy, length 2.7–2.9 mm, dark brown, irregular spots on pronotum, elytra, end of abdomen, legs and antennae paler, red brown (Fig. 20.189a); forebody sparsely punctate and pubescent, pubescence fine, microsculpture meshy and scarcely visible on entire forebody, sculpticells flat; head subquadrate, angulate posteriad, as wide as pronotum, postocular region slightly longer than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; eyes moderate in size; antennae shorter than pronotum and elytra combined, antennomere IV transverse, VI–X strongly transverse, each about twice as broader as long, terminal antennomere as long as two preceding ones combined, basal 3 antennomeres glossy and remaining matte; pronotum moderately transverse, one fourth broader than long, approximately rectangular in shape, hypomeron visible for about three thirds of pronotal length in lateral view, pubescence directed obliquely laterad from midline of disc; elytra flattened, at suture shorter than pronotum along midline, slightly broader than pronotum, posterior margins almost straight and with minute lateral emarginations, pubescence directed straight or slightly obliquely posteriad with some wavy pattern; abdomen subparallel, at base almost as wide as elytra, strongly glossy, three first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII transverse, arcuate apically and without teeth

396

(Fig. 20.189c); sternite VIII transverse, and parabolic apically (Fig. 20.189d); tubus of median lobe of aedeagus sinuate basally and arcuate apically, apex pointed and produced ventrad in lateral view (Fig. 20.189b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 20.189e); sternite VIII parabolic apically, antecostal suture arcuate (Fig. 20.189f); spermatheca with club-shaped, spherical capsule and shallow apical invagination, stem strongly sinuate, and narrowly looped posteriorly, apex twisted (Fig. 20.189g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: Gary oak, litter/soil, moss. Collecting period: V. Collecting method: litter/soil sifting, processing moss through Berlese funnels. 155. Atheta (Microdota) holmbergi Bernhauer (Fig. 20.190a–h) Atheta (Microdota) holmbergi Bernhauer, 1907. Moore and Legner 1975, Klimaszewski and Winchester 2002. Lectotype (male): Alaska: Amer. R. Sikha; coll. Holmberg; Brh. Cotypus; Chicago NHMus. M. Bernhauer collection; Lectotypus male, Atheta holmbergi Bernhauer, V.I. Gusarov des. 2000 (FMNH). Lectotype present designation. Paralectotype (female): Alaska: Amer. R. Sitka; coll. Holmberg; Holmbergi Brh. Typus; NHMus. M. Bernhauer collection; Paralectotypus female Atheta holmbergi Bernhauer, V.I. Gusarov des. 2000 (FMNH). Examined. Diagnosis. Body moderately convex, strongly glossy, narrowly subparallel, length 2.5–3.3 mm, uniformly black, or black with central and posterior part of elytra red brown, legs paler rust-brown, antenna uniformly black (Fig. 20.190a); forebody moderately densely punctate and pubescent, punctation asperate, microsculpture meshy and well visible on entire forebody, sculpticells slightly convex; head

20

Tribe Athetini Casey, 1910

slightly elongate, round, nearly as wide as pronotum, postocular region slightly shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; eyes large and slightly protruding; antennae slender and nearly as long as pronotum and elytra combined, antennomere IV slightly elongate, VI–X transverse, each at most one third broader than long, terminal antennomere as long as two preceding ones combined, basal 4–5 antennomeres glossy and remaining matte; pronotum slightly transverse, approximately rectangular in shape, hypomeron visible for three thirds of pronotal length in lateral view, pubescence directed obliquely laterad from midline of disc; elytra moderately convex, at suture longer than pronotum along midline, distinctly broader than pronotum, posterior margins broadly arcuate and without shallow angular emarginations, pubescence directed straight or slightly obliquely posteriad, forming gently wavy lines posteriorly; abdomen subparallel, at base distinctly narrower than elytra, strongly glossy, three first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII arcuate apically and without teeth (Fig. 20.190d); sternite VIII rounded apically (Fig. 20.190e); median lobe of aedeagus with broadly oval and narrowed apically bulbus and short, broadly triangular tubus in dorsal view (Fig. 20.190c), in lateral view tubus short, arcuate basally and straight apically, apex rounded (Fig. 20.190b). FEMALE. Tergite VIII truncate apically (Fig. 20.190f); sternite VIII rounded apically, antecostal suture sinuate (Fig. 20.190g); spermatheca with tubular capsule and deep apical invagination, stem strongly sinuate, narrowly looped posteriorly, apex slightly swollen (Fig. 20.190h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. New locality data: British Columbia, Queen Charlotte Is., Gate Cr., 10.VIII.1983, 83-72, sifting wet moss on forest floor, coll. J.M. Campbell (CNC) 1 male;

20

Tribe Athetini Casey, 1910

Valemont, N of Packsaddle ck, 52.78119.164, 767 m, 3-9.VI.2014, pitfall, Kinbasket # 1028, R-Mature, coll. Charlene Wood (RBCM) 1 male, 2 females; BC, Log Lake, 65 km N of Prince George, 54.2-122.616, 780 m, OM3C2SX, VIII-IX. 2012, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 male, 1 female; Kootenay, Porcupine forest service rd., 49.2646117.1815, 820 m, 12.IX.2011, Berlese, moss, coll. R.G.B. Bennett and D.&C. Copley (RBCM) 1 male, 1 female; Mount Cain, 50.14 N, 126.21 W, 23.VI-7.VII.1997, 16.6 PIT 11, 13.VII-27.VII.1996, 16.6 L. 3, 27.VII-10. VIII, 1996, 16.6 P. 2, N. Winchester (LFC) 2 males, 5 females. COLLECTION AND HABITAT DATA. Habitat: transition zone in Sitka spruce forest, Carmanah Valley, Vancouver Island (Klimaszewski and Winchester 2002); wet moss on forest floor; sub-boreal spruce forest, moss. Collecting period: VI–IX. Collecting method: pitfall traps, processing moss through Berlese funnels. Comments. This species was previously recorded from the type locality in Sitka Is., Alaska (Bernhauer 1907), and Carmanah Valley on Vancouver Island (Klimaszewski and Winchester 2002). Here, we provide additional localities for this species in BC. We designate a Lectotype because Gusarov’s designation was never published. 156. Atheta (Microdota) lucifera Bernhauer (Fig. 20.191a–h) Atheta (Microdota) lucifera Bernhauer 1906. Moore and Legner 1975. Diagnosis. Body moderately convex, narrowly subparallel, strongly glossy, length 2.8–3.0 mm, head and posterior abdomen dark brown to black, remainder of body red-brown, legs and antennae paler rust-brown (Fig. 20.191a); forebody sparsely punctate and pubescent, microsculpture meshy and scarcely visible on entire forebody, sculpticells slightly convex; head subquadrate,

397

round, nearly as wide as pronotum, postocular region slightly shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong throughout; eyes large; antennae slender and shorter than pronotum and elytra combined, antennomere IV transverse, VI–X transverse, each at most one thirds broader than long, terminal antennomere as long as two preceding ones combined, basal 3–4 antennomeres glossy and remaining matte; pronotum slightly transverse, one fourth broader than long, approximately rectangular in shape, hypomeron visible for about three thirds of pronotal length in lateral view, pubescence directed obliquely laterad from midline of disc; elytra flattened, at suture about as long as pronotum along midline, slightly broader than pronotum, posterior margins broadly arcuate and without shallow angular emarginations, pubescence directed straight or slightly obliquely posteriad; abdomen subparallel, at base distinctly narrower than elytra, strongly glossy, three first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII transverse, arcuate apically and without teeth (Fig. 20.191d); sternite VIII transverse, and parabolic apically (Fig. 20.191e); median lobe of aedeagus with narrowly oval bulbus and long, narrowly triangular, constricted basally tubus, apex rounded in dorsal view (Fig. 20.191c), in lateral view tubus long, straight basally and arcuate apically, apex rounded and pointed ventrad (Fig. 20.191b). FEMALE. Tergite VIII sinuate apically (Fig. 20.191f); sternite VIII rounded apically, antecostal suture arcuate (Fig. 20.191g); spermatheca with tubular, clubshaped capsule and shallow apical invagination, stem strongly sinuate, and narrowly looped posteriorly (Fig. 20.191h). DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record]. USA: CA, WA. We report this species from BC for the first time here based on the following: British Columbia, Victoria, Fort Rodd Hill, 48.4344-123.455, 9 m, 23.V.2015, Berlese, moss, coll. Daren Copley (RBCM) 2 males, 3 females; Victoria,

398

Victoria, Fort Rodd Hill, 48.4348-123.4521, 10 m, 23.V.2015, Gary oak, soil sifting, coll. B. Godin and D. Horwood (BGC) 1 male, 1 female. COLLECTION AND HABITAT DATA. Habitat: Gary oak, litter/soil, moss. Collecting period: V. Collecting method: litter/soil sifting, processing moss through Berlese funnels. 157. Atheta (Microdota) platonoffi Brundin  (Fig. 20.192a–h) Atheta (Microdota) platonoffi Brundin, 1948. Smetana 2004, Klimaszewski et al. 2005, 2012, 2015a, c, 2018b, Majka and Klimaszewski 2008b. Diagnosis. Body small and narrow, subparallel, length 1.5–1.9 mm; colour dark brown to almost black, with legs and elytra except for scutellar region, often paler and yellowish- or reddishbrown (Fig. 20.192a); integument moderately glossy, forebody with fine, dense, slightly asperate punctation and meshed microsculpture; head almost as wide as pronotum, pronotum slightly narrower than elytra; eyes large, much longer than genae in dorsal view; antennae slender, slightly incrassate apically, antennomere IV subquadrate, V–X slightly to distinctly transverse; elytra slightly broader than pronotum, at suture longer than pronotum; abdomen subparallel, highly glossy. MALE. Tergite VIII truncate apically (Fig. 20.192d); sternite VIII with apical margin obliquely produced, arcuately rounded apically (Fig. 20.192e); median lobe of aedeagus with tubus strongly sinuate and strongly arcuate baso-medially, apical part and apex pointing dorsally in lateral view (Fig. 20.192b); in dorsal view bulbus large, subtriangular, and tubus short and triangular apically (Fig. 20.192c), internal sac split apically in dorsal view (Fig. 20.192c). FEMALE. Tergite VIII truncate apically (Fig. 20.192f); sternite VIII rounded apically (Fig. 20.192g); spermatheca highly sinuate, acutely bent at base of capsule, capsule tubular

20

Tribe Athetini Casey, 1910

with long and narrow apical invagination, stem narrowly looped posteriorly and slightly twisted at base (Fig. 20.192h). DISTRIBUTION. Origin: Holarctic. CANADA: AB, BC, LB, NB, NF, NS, ON, SK, YT. USA: AK. New locality data: British Columbia, Queen Charlotte Is., Louise Is., 1.5 km W Skedans, 6. VIII.1983, 83-60, ex moss along seepage, J.M. Campbell (CNC) 2 males; Queen Charlotte Is., 7.9 km NW Queen Charlotte City, 16-19. VII.1983, dung-carrion trap, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Louise Is., 4.7 km N Rennell, Sound Rd., Ghost Cr., 19. VII.1983, 83-24, 700 ft., ex gravel along edge of stream, J.M. Campbell (CNC) 2 sex undetermined; Queen Charlotte Is., Ghost Creek Drainage, Ghost Main Rd., 7.3 km NW Rennell Sound, 21.VIII.1983, 83-103, Berlese of moss ex under deciduous shrubs around base of cedar, J.M. Campbell (CNC) 2 sex undetermined; Queen Charlotte Is., Queen Charlotte City, 83-10, 16.VII.1983, sifting compost, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., Yakoun Riv., 26 km NW Queen Charlotte City, 21.VIII.1983, 83-101, sifting under mushrooms and moss, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., 11.1 km NW Queen Charlotte City, 83-9, 15. VII.1983, treading sandy river bank, J.M. Campbell (CNC) 2 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: this species occurs in decaying organic material (compost, rotting mushrooms, dung) and litter in forests and open habitats; in NF, specimens from pitfall carrion-baited and flight intercept traps in various mixed and coniferous forests; in NB, from unbaited pitfall traps in a red spruce forest, well rotted mushrooms in jack pine forest, in decaying corncobs (moldy) and cornhusks near a mixed forest (common), in grass debris on river margin, under dung on lake margin; a few flying adults collected in late afternoon in mixed forest opening. In BC, specimens

20

Tribe Athetini Casey, 1910

were found in moss along seepage, in gravel along edge of stream, in moss ex under deciduous shrubs around base of cedar, in moss under deciduous shrubs around base of cedar, and in mushrooms, moss, and compost. Collecting period: V–IX. Collecting method: sifting forest litter, compost, rotting mushrooms, and dung, unbaited and carrion-baited pitfall traps, flight intercept traps. 158. Atheta (Microdota) ripariides Newton (Fig. 20.193a–h) Atheta (Microdota) ripariides Newton, in Klimaszewski et al. 2012. Diagnosis. Body length 1.9–2.0 mm, narrow, subparallel; black, with tarsi reddish-brown; moderately glossy, with fine, dense punctation and meshed microsculpture on forebody (Fig. 20.193a); head approximately as wide as pronotum, depressed medially, eyes slightly protruding; antennae slender, slightly incrassate apicad, antennomeres IV–X slightly to strongly transverse; pronotum emarginate laterally; elytra broader and longer at suture than pronotum; head, pronotum and base of abdomen of the same width; sides of abdomen subparallel. MALE. Tergite VIII truncate apically and without teeth (Fig. 20.193d); sternite VIII parabolic apically (Fig. 20.193e); median lobe of aedeagus with small bulbus and long tubus, subapical part narrow, apex produced ventrad in lateral view (Fig. 20.193b), in dorsal view bulbus oval, narrowed apically, and tubus narrowly triangular (Fig. 20.193c). FEMALE. Tergite VIII truncate apically (Fig. 20.193f); sternite VIII broadly rounded apically (Fig. 20.193g); spermatheca with club-shaped capsule and posteriorly-twisted stem (Fig. 20.193h). This species differs from other Nearctic Microdota by the combination of body shape, strongly punctate surface and the shape of the median lobe of the aedeagus and spermatheca.

399

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], MB, SK, YT. USA: not recorded. We report this species from BC for the first time here based on the following: British Columbia, Admiral Isl. Grp., AD06, 128.271451 N, 52.023064 W, pitfall trap, interior, 21-24. V.2017, 100 Islands Project, C. Ernst (HAK-100ISL17-INV-22) (RBCM) 1 male; Stirling Isl. Grp., ST14, 128.065295 N, 51.783654 W, pitfall trap, North shoreline, 11-16.VI.2017, 100 Islands Project, C. Ernst (HAK-100ISL17-INV-1319) (RBCM) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2012). Habitat: litter in mixed aspen and white spruce forest, wet organic forest litter and mushrooms. In BC, one specimen was found in shoreline habitat and one inland. Collecting period: IX. Collecting method: sifting forest litter and mushrooms, pitfall traps. 159. Atheta (Microdota) surgens (Casey), [new subgeneric combination] (Fig. 20.194a–e) Datomicra surgens Casey 1910. Bousquet et al. 2013. Lectotype (male): Canada, Glenora, BC, Wickham; surgens-2, PARATYPE USNM 39174; lectotypus Datomicra surgens Casey, V.I. Gusarov det. 2000; our present lectotype designation label 2019 (USNM). Examined. Diagnosis. Body length 1.7–2.0 mm, stout, small and narrow, subparallel, broadest at elytra; colour uniformly dark brown with appendages paler or head and pronotum slightly darker, integument strongly glossy, forebody with coarse and dense asperate punctation and meshed microsculpture (Fig. 20.194a); head slightly narrower than pronotum, slightly transverse; antennae slender, antennomere IV subquadrate, and V–X subquadrate to slightly transverse; pronotum transverse, distinctly narrower than elytra at base; elytra distinctly broader than

400

pronotum, at suture slightly longer than pronotum, shoulders angular; abdomen distinctly narrower than elytra, subparallel. MALE. Tergite VIII truncate apically, without teeth (Fig. 20.194d); sternite VIII parabolic apically (Fig. 20.194e); tubus of median lobe of aedeagus almost straight ventrally (slightly sinuate), and with narrow and short triangular apical part in lateral view (Fig. 20.194b); in dorsal view tubus as illustrated (Fig. 20.194c). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. This species was originally described as Datomicra (Casey 1910), but it is here transferred to the subgenus Microdota of Atheta on the basis of its close morphological similarities to A. (M.). platonoffi. Externally, it is distinguishable from A. platonoffi by broader elytra and antennomeres VII–X less transverse (subquadrate to slightly transverse). The two species differ in the shape of the tubus of the median lobe of the aedeagus in lateral view. In A. platanoffi, the ventral margin of the tubus in lateral view is consistently strongly sinuate and strongly arcuate medially (slightly so, almost straight in A. surgens), and the apical triangular part is longer and more projecting upward in A. platonoffi (Fig. 20.192b), while it is almost straight and less projecting upward in A. surgens (Fig. 20.194b). These are clearly close but different species. We have formally designated a lectotype for A. surgens because Gusarov’s designation was not published. One male of the type series of Atheta wrangleri Casey from Alaska belongs to A. surgens, and has a straight ventral margin of the tubus as in the latter species. (Subgenus Pancota Casey, 1910) (Fig. 20.195a–h) (Gusarov 2003a; Seevers 1978; Klimaszewski et al. 2018b)

20

Tribe Athetini Casey, 1910

Diagnosis. Head with umbilicate punctation, pronotum and elytra densely and asperately punctate; pronotum 1.3–1.4 times as wide as long; pronotum with pubescence directed straight posteriad or obliquely posteriad along midline except anteriad in apical third, and obliquely laterad on sides of disc; elytra 1.3 times as long as pronotum; eyes large; infraorbital carinae strong basally and faint apically; antennomere V strongly transverse, VI–X strongly transverse, and highly incrassate; male tergite VIII slightly emarginate medially, bearing four small lateral triangular teeth; spermatheca S-shaped, capsule tubular, club-shaped with deep apical invagination, stem thick and sinuate, apex swollen. 160. Atheta (Pancota) (Fig. 20.195a–h)

nescia

Casey

Pseudota nescia Casey 1910. As Atheta (Pancota): Moore and Legner 1975. As Atheta (Datomicra) vancouveri Klimaszewski: Klimaszewski and Winchester 2002. Diagnosis. Body moderately convex, subparallel, length 2.2–2.9 mm, dark brown, legs paler, elytra reddish brown, antennomeres I–IV rust brown (Fig. 20.195a); forebody moderately densely punctate and pubescent, punctation slightly impressed, microsculpture present and particularly well visible on pronotum, sculpticells meshy and slightly convex; head elongate and slightly depressed basally, distinctly narrower than pronotum, postocular region elongate, slightly shorter than the maximum diameter of eye, tempora carinate, carinae strong basally and faint apically; antennae thick, shorter than pronotum and elytra combined, antennomeres IV–X transverse, and VI–X more than twice as broad as long, terminal antennomere as long as two preceding ones combined, basal 4 antennomeres glossy and the remaining matte; pronotum slightly impressed medially, transverse, approximately rectangular in shape, hypomeron visible across basal two-thirds pronotal length in lateral view, pubescence radiating from midline of disk obliquely laterad

20

Tribe Athetini Casey, 1910

and obliquely posteriad; elytra moderately convex, at suture longer than pronotum, broader than pronotum, posterior margins broadly arcuate and with small angular emarginations, pubescence directed straight and obliquely posteriad; abdomen subparallel, first four visible tergites impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII slightly emarginate medially, bearing four small lateral triangular teeth (Fig. 20.195d); sternite VIII parabolic apically (Fig. 20.195e); tubus of median lobe of aedeagus with large bulbus and moderately long tubus with narrowly triangular apex in lateral view (Fig. 20.195b); in dorsal view median lobe narrowly elongate and tubus triangular (Fig. 20.195c). FEMALE. Tergite VIII transverse and arcuate apically (Fig. 20.195f); sternite VIII broadly rounded apically (Fig. 20.195g); spermatheca S-shaped, capsule tubular, club-shaped with deep apical invagination, stem thick and sinuate, posterior part swollen (Fig. 20.195h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: transition zone of Sitka spruce forest and clearcut, Carmanah Valley, Vancouver Island. Collecting period: VI–X. Collecting method: pitfall traps. Comments. Originally described from Queen Charlotte Islands (Casey 1910). We have followed Moore and Legner’s (1975) classification of this species as Atheta (Pancota). However, the concept of this and many other athetine subgenera need revision and should be treated as tentative. (Subgenus Pseudota Casey, 1910) (Figs. 20.196a–h and 20.197a–h) (Gusarov 2003a; Klimaszewski et al. 2018b) Diagnosis. Pronotum and elytra asperately punctate; pronotum 1.3–1.4 times as wide as long; pronotum with pubescence directed straight posteriad or obliquely posteriad along midline

401

and obliquely posteriad elsewhere on disc; elytra 1.25–1.40 times as long as pronotum; eyes large; infraorbital carinae complete but feeble; antennomere V subquadrate to slightly transverse, VI–X transverse, and moderately to highly incrassate; isthmus between intercoxal processes long; male tergite VIII with four or with multiple teeth. Comments. We adopted the definition of Seevers (1978) for this subgenus but it is inadequate and needs additional studies. Key to Species of Subgenus Atheta (Pseudota) 1. Elytra uniformly yellow-brown (Fig. 20.196a); male tergite VIII with two scarcely visible lateral teeth and margin between them crenulate (Fig. 20.196d); male sternite VIII with oblique row of strong vertical macrosetae in apical half (Fig. 20.196e); spermathecal stem with two posterior coils (Fig. 20.196h)........................ ..................... Atheta (Pseudota) concessa Casey – Elytra dark brown with two paler, yellowbrown oblique spots from shoulder to lower sutural corner of disc (Fig. 20.197a); male tergite VIII with two distinct lateral teeth and margin between them with distinct smaller teeth (Fig. 20.197); male sternite VIII without oblique row of strong vertical macrosetae in apical half (Fig. 20.197e); spermathecal stem with multiple posterior coils (Fig. 20.197h) ................................. ..Atheta (Pseudota) pseudoklagesi Klimaszewski and Webster

161. Atheta (Pseudota) concessa Casey [new subgeneric placement] (Fig. 20.196a–h) Atheta (Lamiota) concessa Casey 1911. As Liogluta: Hatch 1957, Moore and Legner 1975. As Atheta: Klimaszewski and Winchester 2002. Atheta sumpta Casey, 1911. Synonymized by Gusarov (2003a). Diagnosis. Body moderately convex, broadly subparallel, length 2.5–2.9 mm, dark brown,

402

head and central part of abdomen sometimes nearly black, elytra yellowish brown, legs yellowish brown, antennae brown with antennomeres I–III yellowish brown (Fig. 20.196a); forebody densely punctate and pubescent, pubescence long, punctation asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head slightly elongate, round, a little narrower than pronotum, postocular region shorter than maximum diameter of eye, tempora carinate, carinae complete, strong throughout; antennae slender, slightly incrassate, antennomeres VI–X transverse, terminal one elongate and as long as two preceding antennomeres combined, basal 4 antennomeres glossy and remaining matte; pronotum impressed mesobasally, strongly transverse, approximately rectangular in shape, hypomeron visible for three quarters of pronotal length in lateral view, pubescence directed anteriad at one third of apical midline, and obliquely laterad across arcuate lines elsewhere; elytra moderately convex, slightly elongate, broader than pronotum, posterior margins broadly arcuate and with shallow lateral emarginations, pubescence directed straight or slightly obliquely posteriad; abdomen subparallel, strongly glossy, three first visible tergites deeply impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII with two short, broadly triangular lateral teeth, margin between teeth obtuse straight and slightly crenulate (Fig. 20.196d); sternite VIII rounded apically and slightly pointed medially, bearing oblique row of strong vertical macrosetae in apical half of disc (autapomorphy) (Fig. 20.196e); median lobe of aedeagus pear-shaped, bulbus broadly oval and narrowed apically, tubus short, triangular in dorsal view (Fig. 20.196c), in lateral view tubus elongate, arcuate ventrally and apex broad and rounded (Fig. 20.196b). FEMALE. Tergite VIII truncate apically (Fig. 20.196f); sternite eight broadly rounded apically (Fig. 20.196g); spermatheca with capsule pitcher-shaped, and round apically, stem narrowly elongate and sinuate medially, looped posteriorly with two small coils at apex (Fig. 20.196h).

20

Tribe Athetini Casey, 1910

DISTRIBUTION. Origin: Nearctic. CANADA: BC [Metlakatla, Carmanh Valley]. USA: not recorded. New locality data: British Columbia, Queen Charlotte Is., Lyall Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, coll. J.M. Campbell (CNC) 1 male. COLLECTION AND HABITAT DATA. Habitat: transition zone of Sitka spruce forest, Carmanah Valley, Vancouver Island; and in mushrooms and marten dung on thick moss, Queen Charlotte Is. Collecting period: VI–IX. Collecting method: pitfall traps. Comments. We have transferred this species, formerly listed in the subgenus Lamiota, because of its median lobe, spermatheca, and male tergite VIII similarities to the species listed in Pseudota (Klimaszewski et al. 2018b). 162. Atheta (Pseudota) pseudoklagesi Klimaszewski and Webster (Fig. 20.197a–h) Atheta (Pseudota) pseudoklagesi Klimaszewski and Webster, in Webster et al. 2016. Klimaszewski et al. 2016b, c, 2018b. Diagnosis. Body narrowly oval, length 2.9–3.5 mm; head, pronotum, and posterior part of abdomen dark brown to nearly black, elytra dark brown with oblique yellowish-brown band extending from shoulder to posterior part of suture, legs, bases of antennae and maxillary palpi yellow, basal part of abdomen dark brown to nearly black, light brown in some individuals (Fig. 20.197a); integument highly glossy, with meshed microsculpture; forebody with punctation and pubescence minute and dense, less so on head; head rounded posterolaterally, eyes moderately large, about as long as genae in dorsal view; antennomere IV subquadrate, V–X moderately to distinctly transverse; pronotum broadly rounded laterally, broadest just anterior of middle of its length, slightly transverse, distinctly wider than head, distinctly narrower than elytra, pubescence

20

Tribe Athetini Casey, 1910

on disc directed laterad from midline; elytra distinctly transverse, with pubescence on disc directed posterolaterad; abdomen subparallel, narrower than elytra. MALE. Tergite VIII truncate apically, with two larger sublateral rounded teeth and multiple smaller narrowly rounded ones between (Fig. 20.197d); sternite VIII rounded apically (Fig. 20.197e); median lobe of aedeagus in lateral view with tubus long, arcuately curved ventrad, with apical part elongate, dorsal margin sinuate to rounded apex (Fig. 20.197b); in dorsal view bulbus moderately broad, ovoid, tubus elongate, tapering to triangular apex (Fig. 20.197c), internal sac with weak structures (Fig. 20.197b, c). FEMALE. Tergite VIII with apical margin arcuate laterally, truncate medially (Fig. 20.197f); sternite VIII broadly rounded apically (Fig. 20.197g); spermatheca with capsule spherical with deep apical invagination, stem long, narrow, tightly coiled posteriorly (Fig. 20.197h). This is the sibling species of A. klagesi (not recorded in BC), and was confused with the latter in collections. It may be distinguished from A. klagesi by the following combination of characters: size slightly larger, body less glossy, legs, bases of antennae, maxillary palps and bands on elytra less intensely yellowish in coloration, body colour less contrasting overall; median lobe of aedeagus with tubus longer, with apex shaped slightly differently in lateral view; the shape of the spermatheca is very similar in the two species, and females may be difficult to identify without accompanying males. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], QC, NB, NF, SK, YT. USA: [all published records of A. klagesi need to be re-examined because they may contain mixed series with A. pseudoklagesi]. We report this species from BC for the first time here based on the following: British Columbia, Boya Lake, 28.VI.2015, 59.3839-129.1035, 698 m, mushrooms sifting, coll. B. Godin and D. Horwood (LFC) 1 male.

403

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, in mature mixed forest, old-growth and old white spruce and balsam fir forests, a mature red spruce forest, and in a wet alder swamp; adults in coral fungi on a Populus log, fleshy polypore fungi at base of a dead standing Populus, in decaying gilled mushrooms, in gilled mushrooms, and under bark of red spruce and dead aspen. In NF, from pitfall traps in boreal forests. In QC, in yellow birch/balsam fir forest, using pitfall traps, Luminoc pit/light traps, and intercept traps. In SK, in aspen woodland, in bracket/gilled fungi, under bark of dead aspen, and on fungus on dead lodgepole pine stump. In YT, in mushrooms. Collecting period: V–X. Collecting method: sifting mushrooms. Comments. In the past, the two sibling species were mixed together and identified as A. klagesi (see above). All material across Canada needs to be re-examined to understand the true distribution of the two species. So far, only AB, BC, NF, NB, QC, SK, and YT specimens have been re-evaluated. Subgenus Tetropla Mulsant and Rey, 1873b (Fig. 20.198a–j) (Gusarov 2003a; Klimaszewski et al. 2018b) Diagnosis. Body narrowly subparallel; pronotum and elytra asperately punctate; head rounded basally, eyes large, longer than genae in dorsal view; antennomere V subquadrate to slightly transverse, VIII–X 1.3 times as wide as long; pronotum about four-fifths as wide as elytra, pubescence on disc directed obliquely posteriad from midline; elytra at suture slightly longer than pronotum along midline, posterior margin truncate and straight, without lateral emargination; apical margin of male tergite VIII angulately produced in middle 2/3, sinuate medially; median lobe of aedeagus with two elongate structures in bulbus; spermatheca with capsule club-shaped, straight, with narrow and deep apical invagination, stem angularly looped, twisted and slightly swollen basally.

404

163. Atheta (Tetropla) frosti Bernhauer (Fig. 20.198a–j) Atheta (Atheta) frosti Bernhauer, 1909. Casey 1910, 1911 [as ten synonyms], Gusarov 2003a, Klimaszewski et al. 2005, 2011, 2016d, 2018b, Majka and Klimaszewski 2008b. Diagnosis. Body narrowly subparallel, moderately convex, length 2.0–2.8 mm; colour brown to dark brown, with antennomeres I–III, palps, legs, elytra and apical part of abdomen light brown to yellowish-brown, integument moderately glossy (Fig. 20.198a); forebody with pubescence fine and sparse, denser on elytra; head narrower than pronotum, genae shorter than eyes in dorsal view; antennae moderately thick, antennomere IV subquadrate to slightly transverse, V–X moderately transverse, each about 1.5 times as wide as long; pronotum distinctly transverse, widest at middle, slightly narrower than elytra; elytra at suture slightly longer than pronotum, posterior margin straight, without lateral emarginations; abdomen mostly subparallel. MALE. Apical margin of tergite VIII angulately produced in middle 2/3, sinuate medially (Fig. 20.198e); sternite VIII with sides almost evenly arcuate to slightly more rounded apex (Fig. 20.198f); median lobe of aedeagus in lateral view with tubus moderately long, arcuate ventrally, with apex narrow and rounded (Fig. 20.198b, c); in dorsal view bulbus oval, tubus broad and triangular apically (Fig. 20.198d); internal structures elongate (Fig. 20.198d). FEMALE. Tergite VIII with apical margin obtuse laterally, broadly truncate medially (Fig. 20.198g); sternite VIII semicircularly rounded from antecostal suture to apex (Fig. 20.198h); spermatheca with capsule club-shaped, straight, with narrow and deep apical invagination, stem angularly looped, twisted and slightly swollen basally (Fig. 20.198i, j). DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NS, ON, QC, SK. USA: MA, NC, NH, NY, PA, RI, VT, WV.

20

Tribe Athetini Casey, 1910

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, in various coniferous, mixed, and hardwood forests in decaying fungi where it is often common; white pine and spruce forest in decaying mushrooms, old-growth eastern hemlock forest in decaying mushrooms, red spruce and eastern white cedar forest in decaying mushrooms, red pine forest in decaying mushrooms, jack pine forest in decaying mushrooms, mixed forests in decaying mushrooms, red oak forest in gilled mushrooms, rich Appalachian hardwood forest in decaying/rotten fleshy fungi; one from pitfall trap in red spruce forest and some in decaying (moldy) corncobs and cornhusks near a mixed forest. Elsewhere, from pitfall traps in open spruce forest with sandy soil and Cladina lichen cover (LB), birch-dominated forests (QC), in polypore fungus in coniferous forest, and on forest floor in red oak and deciduous forests (NS). Collecting period: VII–X. Collecting method: sifting fungi and forest litter, pitfall traps. Comments. Before 2003, North American specimens identified as Atheta nigritula (Gravenhorst) were actually Atheta frosti (Gusarov 2003a).

Atheta Incertae Sedis 164. Atheta brunswickensis Klimaszewski (Fig. 20.199a–h) Atheta brunswickensis Klimaszewski, in Klimaszewski et al. 2005. Klimaszewski et al. 2007b, 2008a, 2018b. Diagnosis. Body narrowly subparallel, length 1.9–2.4 mm, colour yellowish brown or rustbrown, with head, antennae (except for basal two or three segments), and apical part of abdomen dark brown, or entire body dark brown with legs lighter, yellowish-brown (Fig. 20.199a); integument moderately glossy, forebody with

20

Tribe Athetini Casey, 1910

isodiametric microsculpture and fine punctation; pubescence moderately dense, sparser on pronotum than on elytra; head rounded posteriad, slightly transverse, broadest at level of eyes, strongly flattened medially; antennomere IV subquadrate, V–X slightly to moderately transverse, slightly incrassate apically, and terminal one elongate; pronotum convex, moderately transverse (wider in females than in males), broadest at middle, pubescence on disc directed laterad from midline; elytra transverse, at suture about as long as pronotum along midline, pubescence on disc directed slightly obliquely posteriad; abdomen subparallel. MALE. Tergite VIII truncate apically, with two small, rounded, apicolateral teeth (Fig. 20.199d); sternite VIII broadly rounded apically, with antecostal suture sinuate, widely separated from basal margin (Fig. 20.199e); median lobe of aedeagus in lateral view with crista apicalis of bulbus narrowly elongate, tubus narrowly elongate, broadly curved ventrally, apical part moderately broad and moderately produced ventrally (Fig. 20.199b); in dorsal view bulbus oval in outline and evenly coalescent with narrowly elongate tubus in dorsal view (Fig. 20.199c); internal sac with two inconspicuous round structures (Fig. 20.199c). FEMALE. Tergite VIII truncate apically (Fig. 20.199f); sternite VIII broadly rounded posteriorly, with apical fringe of microsetae, antecostal suture narrowly separate from basal margin (Fig. 20.199g); spermatheca with capsule club-shaped, stem thick, sinuate, narrowly looped and twisted posteriorly (Fig. 20.199h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, NS, ON, QC, YT. USA: not recorded. We report this species from BC for the first time here based on the following: BC, Mount Cain, 50.15 N, 126.24 W, 28.VII-10.VIII.1996, 17.4 P.2, N. Winchester (LFC) 1 female; Goose Isl. Grp., GS06, 51.92814 N, 128.4397 W, Yellow pan trap, Interior, 8-12.V.2015, 100 Islands Project, C. Ernst, HAK-100ISL15-INV-820 (RBCM) 1 female.

405

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, from various forest types; upland black spruce forest, red spruce and yellow birch forest, mature red spruce and eastern white cedar forest, balsam fir stand, red pine forest, jack pine forest, various mixed forests, red oak forest, and various hardwood forests. Adults in various kinds of mushrooms (mostly gilled mushroom species) in various stages of decay (fresh to well rotted) on forest floor; also found in forest litter (moldy conifer duff, leaf litter). Some from pitfall traps in regenerating red spruce forest and yellow birch forest. Others from decaying (moldy) corncobs and cornhusks near mixed forest. Collecting period: V–IX. Collecting method: sifting fungi and forest litter, pitfall traps, Yellow pan trap. Comments. Atheta brunswickensis bears certain characteristics common with A. irrupta: both species have similarly shaped male tergite VIII with two rounded lateral teeth, similar shape of median lobe of aedeagus in dorsal view, internal sac with weak structures, similarly shaped spermatheca (with compressed spherical capsule and large but shallow invagination and short neck, and identically shaped stem), and similar body form. These two species should probably form a group of their own. 165. Atheta formalis (Casey) Pseudota formalis Casey, 1911. Diagnosis (based on original description). Body slender, moderately convex, glossy, length 1.2 mm; colour black; punctation fine and slightly asperate; head with median part of disc broadly impressed; pronotum slightly broader than head, small, not impressed; elytra broad, one-fourth wider than pronotum, at suture three-fifth longer than pronotum; abdomen subparallel, much narrower than elytra, three basal tergites impressed basally. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

406

COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. Known only from the original specimen reported from Metlakatla, BC. The holotype specimen of this species is missing at USNM. The labels and pin are there, but the specimen has been lost undoubtedly at some point due to mishandling (F. Shockly, pers. com.). In the future, a neotype should be designated from the type locality if it can be matched with the original description, or the name should be considered as a nomen nudum and removed from the valid species list. 166. Atheta irrupta (Casey) (Fig. 20.200a–i) Pseudota irrupta Casey 1910. As Atheta (Pancota): Hatch 1957, Moore and Legner 1975. As Atheta (Pseudota): Klimaszewski and Winchester 2002. Diagnosis. Body moderately convex, subparallel, length 3.0–3.6 mm, light yellowish brown with some rust tinge (rarely entirely dark brown), head and central part of abdomen dark brown, legs yellowish brown, antennae dark brown with antennomeres I–III yellow-brown (Fig. 20.200a); forebody moderately densely punctate and pubescent, pubescence long, punctation slightly asperate, microsculpture meshy and particularly well visible on pronotum, sculpticells slightly convex; head slightly elongate, round, distinctly narrower than pronotum, postocular region about as long as the maximum diameter of eye, tempora carinate, the carinae complete, strong but weak apically; antennae slightly incrassate, shorter than pronotum and elytra combined, antennomere IV slightly transverse, VI–X transverse, each at most one-third wider than long, terminal antennomere as long as two preceding ones combined, basal 4 antennomeres glossy and remaining matte; pronotum strongly transverse, approximately rectangular in shape, hypomeron visible for one third of pronotal length in lateral view,

20

Tribe Athetini Casey, 1910

pubescence directed nearly horizontally laterad and obliquely laterad from midline of disc; elytra moderately convex, distinctly elongate, broader than pronotum, at suture slightly longer than pronotum along midline, posterior margins, broadly arcuate and with shallow lateral emarginations, pubescence directed straight or slightly obliquely posteriad; abdomen subparallel, strongly glossy, four first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII with two short, broadly triangular and rounded lateral teeth, margin between teeth obtuse and arcuate (Fig. 20.200e); sternite VIIII rounded apically (Fig. 20.200f); median lobe of aedeagus with narrowly oval and narrowed apically bulbus and triangular tubus in dorsal view (Fig. 20.200d), in lateral view tubus narrowly elongate apically and strongly produced ventrally (Fig. 20.200b, c). FEMALE. Tergite VIII transverse and emarginated at apex (Fig. 20.200g); sternite VIII broadly rounded apically (Fig. 20.200h); spermatheca with capsule hemispherical and posteriorly slightly pitcher-shaped, stem broadly arcuate medially, and looped posteriorly with 1–2 small coils at apex (Fig. 20.200i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: transition zone and forest interior of Sitka spruce forest, Carmanah Valley, Vancouver Island. Collecting period: VI–IX. Collecting method: pitfall traps. Comments. This species is known only from the type locality, Metlakatla, and from Vancouver Island (Klimaszewski and Winchester 2002). It bears a certain resemblance to A. brunswickensis in the similar shape of male tergite VIII and the female spermatheca. It remains to be seen, if these similarities are indicative of any close relationship. 167. Atheta pseudoschistoglossa Klimaszewski and Webster (Fig. 20.201a–h)

20

Tribe Athetini Casey, 1910

Atheta pseudoschistoglossa Klimaszewski and Webster, in Webster et al. 2016. Klimaszewski et al. 2016d, 2018b. Diagnosis. Body narrowly elongate, subparallel, length 2.9 mm; head, pronotum, and abdomen dark brown, elytra rust brown mottled with black, legs and antennae light brown (Fig. 20.201a); integument highly glossy; forebody with minute and sparse punctation and sparse pubescence; head rounded and slightly angular posterolaterally, eyes moderately large, shorter than genae in dorsal view; antennomeres V–X subquadrate to slightly transverse; pronotum rounded anterolaterally and posterolaterally, slightly transverse, insignificantly wider than head and slightly narrower than elytra, pubescence on disc directed laterad from midline; elytra slightly transverse, flattened, with pubescence on disc directed posterolaterad; abdomen subparallel medially, narrower than elytra. MALE. Tergite VIII arcuate apically (Fig. 20.201d); sternite VIII distinctly elongate and rounded apically, like in species of Philhygra (Fig. 20.201e); median lobe of aedeagus in lateral view with bulbus oval, tubus strongly produced ventrally, apex narrow and pointed ventrally (Fig. 20.201b); in dorsal view bulbus oval, tubus subparallel basally and triangular apically (Fig. 20.201c); internal sac with complex structures (Fig. 20.201c). FEMALE. Tergite VIII with apical margin truncate (Fig. 20.201f); sternite VIII evenly broadly rounded apically (Fig. 20.201g); spermatheca with capsule narrow, sac-shaped, with shallow apical invagination, stem sinuate, narrowly hooked posteriorly (Fig. 20.201h).

407

material, and flood debris along river margins, moist leaves along vernal pond margin in a silver maple swamp, in leaf litter and moss along brook margins in alder swamps, and in litter at base of red maple, in Carex hummock in Carex marshes, in leaf litter in a red oak forest near a flooded seasonally flooded marsh, in a salt marsh, in marsh litter in a Carex–sedge marsh, and in litter and sphagnum at the base of a tree in a marsh. A few individuals were captured in Lindgren funnel traps in a mixed forest and in a hardwood woodland near a seasonally flooded marsh. Collecting period: V–VIII. Collecting method: sifting various kinds of leaf litter, debris, and moss, Lindgren funnel traps. Comments. The subgeneric position of this species is unsettled. It bears a superficial resemblance to members of the genus Schistoglossa but does not have the apical parts of the mandibles split.

PALEARCTIC SPECIES NOT CONFIRMED IN BC AND EXCLUDED FROM THE LIST OF VALID SPECIES IN BC

Atheta basicornis (Mulsant and Rey, 1852) { (Fig. 20.202a–i) Atheta subrugosa (Märkel and Kiessenwetter, 1848) { (Fig. 20.203a–g)

DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, SK. USA: AK.

Comments. The records of these two species from BC by Bernhauer (1909: 520 and 1907: 394 respectively), represent most likely misidentifications. We have not found these species in any samples examined from BC. We provide, however, images for these species, based on European specimens, for future reference.

New locality data: USA, AK, 8-16 miles E of Willow, 7.VII.1988, evening sweeping gravel pit, leg. R. Baranowski (ZMLU) 1 male.

Genus Boreophilia Benick, (Fig. 20.204a–f) (Lohse et al. Klimaszewski et al. 2011, 2018b, 2019)

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in and near wetland habitats; among cobblestones, drift

Diagnosis. Body moderately narrow to broad, subparallel; average body length 3.0 mm; head

1973 1990;

408

20

narrower or nearly as broad as pronotum; eyes moderate in size, as long as or shorter than postocular region of head, tempora partially feebly carinate at base of head; antennae often reaching posterior margin of elytra, antennomeres V–X subquadrate, slightly transvers, or elongate; ligula divided into two separate and diverging lobes; mandibles simple (not bifid at apex as in Schistoglossa); maxillary palps with 4 palpomeres, terminal one needle-shaped; pronotum transverse, broadest in the middle or in apical fourth, hind angles rounded, hypomeron clearly visible in lateral view; integument smooth and moderately glossy, weak microsculpture present, punctation fine, pubescence on midline of disc directed anteriad at least in apical ¾, and elsewhere directed laterally; mesocoxae contiguous; median lobe of aedeagus broad, flattened latero-ventrally, bulbus enlarged and tubus narrow, short or moderately elongate, approximately triangularly shaped in dorsal view, crista apicalis narrowly elongate, internal sac with two large, elongate sclerites in bulbus, and additional smaller sclerites in apical part of internal sac; male tergite VIII entire, and not modified; spermatheca diverse in shape, with elongate tubular capsule and sinuate stem, capsule’s invagination small or absent; cold loving species occurring in arctic habitats, in temperate regions usually confined to fens and bogs. Comments. It is very likely that there are more species of Boreophilia in the northern parts of BC at higher elevations, which are not reported in this book due to insufficient collecting efforts. For any additional species of Boreophilia found in BC, readers should consult the recent revision of this genus (Klimaszewski et al. 2019). 168. Boreophilia 20.204a–f)

nomensis

(Casey) (Fig.

Dinaraea nomensis Casey, 1910. As Boreophilia: Lohse et al. 1990, Gusarov 2003a, Klimaszewski et al. 2019.

Tribe Athetini Casey, 1910

Diagnosis. Body narrow, subparallel, moderately glossy, abdomen slightly more so; length 3.0–3.8 mm; head, pronotum and abdomen dark brown, elytra reddish-brown, legs paler, yellowish-brown or specimens predominantly dark brown (Fig. 20.204a); antennomeres VIII– X subquadrate; pronotum longer than elytra at suture, maximum width of pronotum about the same as maximum width of elytra. MALE. Tergite VIII arcuate apically (Fig. 20.204e); sternite VIII elongate, parabolic apically (Fig. 20.204f); tubus of median lobe of aedeagus almost straight (slightly arcuate) in lateral view, apex triangular in shape, slightly pointed (Fig. 20.204b), bulbus broad in dorsal view and with two elongate narrow sclerites as illustrated (Figs. 20.204c, d). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: BC, YT. USA: AK. New locality data: USA, AK, near Hatcher Pass, 12-15 mi NW of Palmer, 600–800 m, 6. VIII.1988, under stone, open ground, leg. R. Baranowski (ZMLU) 1 male; Eureka Summit, 1000 m, Hwy 1, leg. R. Baranowski (ZMLU) 1 male. COLLECTION AND HABITAT DATA. Habitat: spruce and aspen forest with horsetail/ shrub/grass undergrowth; edge of snowfield; and under stone in open ground. Collecting period: VI–VIII. Collecting method: pitfall trap, hand collecting under rocks. Genus Boreostiba Lohse, 1990 (Figs. 20.205a– h and 20.206a–h) (Lohse et al. 1990; Klimaszewski et al. 2011, 2018b) Diagnosis. Body elongate, subparallel; integument brown to black, elytra dark brown to black or in some species reddish-brown; ligula divided into two lobes in apical half; antennomeres VII–X moderately to distinctly transverse; head with infraorbital carinae incomplete, never reaching hind border of eye; elytra at suture at most as long as pronotum along midline; mesocoxae

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Tribe Athetini Casey, 1910

narrowly separated; procoxal cavities separated from lateral margin of pronotum by long hypomera; median lobe of aedeagus in lateral view with tubus strongly produced ventrally, thin subapically; condylite of paramere greatly enlarged in most species; male tergite VIII truncate apically, with margin entire and without distinct teeth; spermatheca with capsule club- or pitcher-shaped, stem thin, sinuate, narrowly looped posteriorly. Comments. The taxonomic status of this genus is unsettled and in need of revision. Key to Species of Boreostiba 1. Ventral margin of tubus of median lobe of aedeagus straight basally in lateral view, apex moderately narrow (Fig. 20.206a); capsule of spermatheca narrowly cub-shaped, stem moderately thick and long, strongly sinuate and looped poseriorly (Fig. 20.206h) ..................................... .................. Boreostiba parvipennis (Bernhauer) – Ventral margin of tubus of median lobe of aedeagus sinuate basally in lateral view, apex very narrow (Fig. 20.205a); capsule of spermatheca large and broadly pitcher-shaped, stem mostly thin and moderately long, strongly sinuate and looped poseriorly (Fig. 20.205h)................... ......... Boreostiba pseudolaticollis Klimaszewski and Godin, sp. n.

169. Boreostiba pseudolaticollis Klimaszewski and Godin, sp. n. (Fig. 20.205a–h) Holotype (male): USA, Alaska Skagway, Dyea, 59.5301-135.3583, 112 m, 5.V.2013, Hemlock, sifting, coll. B. Godin and D. Horwood (CNC). Paratypes: Canada, British Columbia, Mt. Cain, 50.13 N, 126.21 W, 9.VI23.VI.1997, 17.4 PIT 2, coll. N. Winchester (LFC) 1 male, 1 female; Mt. Cain, 50.13 N, 126.21 W, 5.VI-27.VI.1996, 16.6 P. 1, coll. N. Winchester (LFC) 1 male, 1 female; Mt. Cain, 50.13 N, 126.21 W, 23.VI-7. VII.1997, 16.6 PIT 4, coll. N. Winchester (LFC, RBCM) 8 females; Mt. Cain, 50.13 N,

409

27.VI-13.VII.1996, 126.21 W, coll. N. Winchester (LFC) 1 male, 2 females; Mt. Cain, 50.13 N, 126.21 W, 7.VII-20. VII.1997, 16.6 PIT4, coll. N. Winchester (LFC, RBCM) 2males, 7 females; Mt. Cain, 50.13 N, 126.21 W, 20.VII-5.VIII.1997, 16/6 PIT 10, coll. N. Winchester (CNC, LFC) 1 male, 8 females; Mt. Cain, 50.13 N, 126.21 W, 10. VIII-23.VIII.1966, 16.1 PAN 2, coll. N. Winchester (LFC, RBCM) 2 males; Mt. Cain, 50.13 N, 126.21 W, 1.IX-19.IX.1997, 16.6 PIT 8, N. Winchester (LFC) 2 females; Mt. Cain, 50.13 N, 126.21 W, 28.IX-12. X.1996, 16.1 PAN 2, N. Winchester (LFC) 2 females. USA, Alaska: Skagway, Dyea, 59.5273-135.3525, 5.V.2013, Hemlock/alder, ck bank, sifting, coll.B. Godin and D. Horwood (BGC, CNC, LFC) 5 males, 8 females; same data except: 59.5301-135.3583, 112 m, Hemlock, sifting, coll. B. Godin and D. Horwood (BGC) 1 female; 29.III.2013, Hemlock, sifting, coll.B. Godin and D. Horwood (LFC) 2 females. Etymology. Prefix pseudo- added to specific name laticollis, in allusion to certain similarities of the spermatheca of this species to that of the Palearctic Atheta laticollis. Diagnosis. Body narrowly subparallel, length 3.2–3.4 mm; head and abdomen dark brown, pronotum, elytra, legs and basal parts of antennae paler, light brown or reddish brown (Fig. 20.205a); integument highly glossy, particularly on abdomen; forebody with minute and sparse punctation and sparse pubescence; head rounded, eyes moderately large, shorter than genae in dorsal view; antennae slender, antennomere IV elongate, V subquadrate, VI–X moderately transverse; pronotum rounded laterally and posterolaterally, slightly transverse, slightly wider than head and about as wide as elytra, pubescence on disc directed anteriad along midline and laterad from midline of disc; elytra strongly transverse, at suture shorter than pronotum along midline, flattened, with pubescence on disc directed posterolaterad; abdomen broad, arcuate laterally, at middle broader than elytra. MALE. Tergite VIII with apical margin

410

arcuate apically (Fig. 20.205d); sternite VIII parabolic apically and sinuate basally (Fig. 20.205e); median lobe of aedeagus in lateral view with bulbus suboval, tubus strongly produced ventrally, apex narrow and sharp, pointed ventrally (Fig. 20.205b); in dorsal view bulbus broadly oval, sides converging apically, tubus narrowly triangular (Fig. 20.205c); internal sac with two elongate sclerites in dorsal view (Fig. 20.205c). FEMALE. Tergite VIII with apical margin arcuate (Fig. 20.205f); sternite VIII broadly rounded apically (Fig. 20.205g); spermatheca with capsule broad, pitcher-shaped and elbowed-neck, stem thin and narrowly looped posteriorly (Fig. 20.205h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK. COLLECTION AND HABITAT DATA. Habitat: litter in hemlock forest, litter in hemlock/alder forest near creek bank. Collecting period: V–X. Collecting method: sifting forest litter, pitfall traps, Pan traps. 170. Boreostiba parvipennis (Bernhauer) (Fig. 20.206a–h) Atheta (Ousipalia) parvipennis Bernhauer, 1907. Lohse et al. 1990 [as B. hudsonica], Gusarov 2003a, Klimaszewski et al. 2008a, b, 2011, 2015a, 2018b. Diagnosis. Body narrowly elongate, subparallel, length 2.0–2.2 mm; colour dark brown to black, with antennae, elytra except for scutellar region, apex of abdomen and legs reddish-brown (Fig. 20.206a); integument moderately glossy, forebody with fine, dense punctation and meshed microsculpture; head subquadrate, almost as wide as pronotum and elytra, with eyes shorter than genae in dorsal view; antennae moderately robust, antennomere IV subquadrate to slightly elongate, V–X slightly to moderately transverse; pronotum narrowest at base, slightly widening apicad, arcuate laterally, all angles broadly rounded; elytra at suture slightly shorter than pronotum; abdomen

20

Tribe Athetini Casey, 1910

slighty widening apicad. MALE. Tergite VIII with apical margin arcuate, apex truncate, lacking teeth or denticles (Fig. 20.206d); sternite VIII parabolic apically (Fig. 20.206e); median lobe of aedeagus in lateral view with tubus straight basally and strongly produced ventrally at apex (Fig. 20.206b), in dorsal view tubus elongate, narrowly triangular and pointed at apex (Fig. 20.206c). FEMALE. Tergite VIII with apical margin arcuate, apex truncate (Fig. 20.206f); sternite VIII rounded apically (Fig. 20.206g); spermatheca with capsule club-shaped and stem highly sinuate and narrowly looped posteriorly (Fig. 20.206h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], LB, NF, NT, QC, YT. USA: AK, NH. We report this species from BC for the first time here based on the following: British Columbia, Atlin, McDonald Lk., 20.VI.2015, 59.7116133.5787, 939 m, Willow, soil sifting, coll. B. Godin and D. Horwood (BGC, LFC) 1 male, 3 females; Atlin, Pine Creek, 21. VI.2015, 59.7116-133.5787, 939 m, Alder soil sifting, coll. B. Godin and D. Horwood (BGC, LFC) 5 males; Boya Lake, 28.VI.2015, 59.3639129.1035, 698 m, mushroom, sifting, coll. B. Godin and D. Horwood (BGC, LFC) 2 males. COLLECTION AND HABITAT DATA. Habitat: most specimens in eastern Canada are from pitfall traps in fir and spruce forests; one from detritus on a sandy beach and another from a rotting mushroom (Klimaszewski et al. 2018a). In BC, specimens were collected from mushrooms and alder soils/litter. Collecting period: VI–VIII. Collecting method: sifting organic debris including mushrooms, pitfall traps. Comments. Gusarov (2003a) placed this species in Atheta sensu lato pending a generic revision of Boreostiba. We have tentatively retained this species in Boreostiba (as originally in Lohse et al. 1990), due to its greater external similarity to

20

Tribe Athetini Casey, 1910

411

other species of Boreostiba (elytra short, about as long as pronotum, abdomen broadening apically, antennal articles VIII–X slightly transverse) than to Nearctic Atheta. We have examined two paratypes of Boreostiba campbelliana Lohse (male, female) and they are very similar to B. parvipennis in terms of the external morphology and the shape of the median lobe of the aedeagus. We suspect this species may be conspecific with B. parvipennis but we need to examine the holotype specimen which is on loan to V. Gusarov and was not available for study (Klimaszewski et al. 2018b). Genus Clusiota, Casey 1910 (Fig. 20.207a–h) (Klimaszewski et al. 2015c, 2018b) Diagnosis. Body narrowly elongate, length 1.5–2.5 mm; antennomere I more-or-less swollen and longer than II, V–X distinctly transverse; labial palpi with three palpomeres; ligula narrow, deeply split forming V-shaped structure; maxillary palpi with three palpomeres, last one narrowly elongate; pronotum narrower than elytra; pronotal hypomera fully visible medially in lateral view; elytra at suture longer than pronotum along midline, flattened, anterior shoulders narrowly rounded, posterior margin truncate and without distinct lateral emargination; abdomen with sides arcuate, swollen in some; male and female tergite VIII shallowly emarginate medially and without teeth; median lobe of aedeagus with large bulbus, in lateral view tubus strongly curved ventrad at base, subapical part narrow, crista apicalis located on elevated part of bulbus; spermatheca S-shaped, with capsule pear-shaped, apical invagination small and short. 171. Clusiota impressicollis (Fig. 20.207a–h)

(Bernhauer)

Atheta (Microdota) impressicollis Bernhauer, 1907. As Clusiota: Casey 1910 [as C. claviventris], Gusarov 2003b, Klimaszewski et al. 2008a, 2011, 2015c, 2018b.

Diagnosis. Body narrowly elongate, flattened, length 1.9–2.0 mm; colour reddish brown to dark brown, with head, elytra and base of abdomen darker (Fig. 20.207a); integument moderately glossy, forebody densely punctate and densely pubescent, less so on head and abdomen, microsculpture of forebody fine, meshed with hexagonal sculpticells; head as wide as pronotum, slightly angular posteriorly, eyes large, prominent, as long as genae in dorsal view; pronotum arcuate laterally and basally, slightly transverse, narrower than elytra; elytra roughly quadrate, wider and longer than pronotum; abdomen swollen at middle. MALE. Tergite VIII broadly, obtusely emarginate apically (Fig. 20.207d); sternite VIII broadly rounded apically (Fig. 20.207e); median lobe of aedeagus in lateral view with bulbus swollen, globular, tubus curved obliquely ventrad basally, ventral margin sinuate, narrowly subparallel to narrowly rounded apex (Fig. 20.207b), in dorsal view bulbus oval, tubus narrowly elongate, triangular apically (Fig. 20.207c), internal sac structures pronounced (Fig. 20.207b, c). FEMALE. Tergite VIII with apical margin arcuate except for shallow apical median emargination (Fig. 20.207f); sternite VIII evenly broadly rounded apically (Fig. 20.207g); spermatheca elongate S-shaped, capsule broadly club-shaped with moderately deep median invagination, stem sinuate with short posterior loop (Fig. 20.207h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB, NF, ON. USA: AK, CA, WA. New collection data: British Columbia, Vancouver Island, Mt. Cain, 50.15 N, 126.25 W, 23.VI-7.VII.1997, 17.4 LIN 9, N. Winchester (LFC) 1 female. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Macrohabitat: in NF, specimens from flight intercept traps in a fir-deciduous forest; a specimen from NB under coyote dung on a gravel road through an old

412

growth white spruce and balsam fir forest. Collecting period: VII–VIII. Collecting method: sifting dung, flight intercept traps. Comments (Klimaszewski et al. 2018b). Atheta impressicollis (Bernhauer 1907) was replaced with Atheta (Microdota) nova Fenyes, 1920: 187, because at that time it was a junior secondary homonym of Atheta impressicollis (Mulsant and Rey 1852: 27). It was first published in the genus Clusiota by Gouix and Klimaszewski (2007), and has been cited thus several times since then (see above). According to ICZN Article 59.3, even though the junior homonym (C. impressicollis) was replaced before 1961, it is not to be rejected since the replacement name has not been in use since Moore and Legner (1975), and the two homonyms are not considered congeneric. Genus Dalotia Casey, 1910 (Fig. 20.208a–g) (Gusarov 2003a, Klimaszewski et al. 2013a, 2018b) Diagnosis. Body broad, narrowly oval, brown to black, glossy; head almost as wide as pronotum, infraorbital carina complete; epipharynx with a sensilla long; hypopharynx with long setae; ligula with narrow base and divided apical part bearing two long setae; labial palpi with setae α, β, γ, and δ present; pronotum 1.3 times as wide as long, microsetae on disc directed anteriad and laterad along midline, obliquely posteriad elsewhere, macrosetae long; pronotal hypomera fully visible in lateral view; process of mesoventrite narrow; elytra longer and wider than pronotum; abdomen broad, tapering apically; mesolateral macroseta of middle tibia coarse; hind tarsus with basal article as long as second one; male tergite VIII with two lateral teeth and distinctive broadly V-shaped apical emargination with several irregular in size small teeth; median lobe of aedeagus in lateral view with enlarged bulbus and very short tubus, internal sac sclerites complex; spermatheca sinuate, capsule with apical hemispherical extension, stem short and highly sinuate. Illustrations of

20

Tribe Athetini Casey, 1910

mouthparts are in Figs. 125 and 126 of Gusarov (2003a). 172. Dalotia coriaria (Kraatz) { (Fig. 20.208a–g) Homalota coriaria Kraatz, 1856. Bernhauer 1906, 1907, Casey 1910 [as Pseudota miscella, Dimetrota crucialis and D. pectorina], Criddle 1922, Frank 1981, Muona 1984, Klimaszewski and Peck 1986, Gusarov 2003a [list of synonyms], Gouix and Klimaszewski 2007, Klimaszewski et al. 2007a, 2009a, 2013a, 2018b, Webster et al. 2009, Pochubay and Grieshop 2012, Renkema et al. 2015. Diagnosis. Body length 2.1–2.8 mm; colour brown to almost black, with 2–3 basal antennomeres, legs or only tarsi paler, reddish brown (Fig. 20.208a); integument moderately glossy, forebody apparently without microsculpture, densely punctate and densely pubescent, less so on head and abdomen; head slightly narrower than pronotum, rounded posteriorly, eyes large, prominent, as long as genae in dorsal view; pronotum rounded laterally and basally, distinctly transverse; elytra wider and longer than pronotum; abdomen broad basally and slightly tapering apically. MALE. Apical margin of tergite VIII oblique laterally, with two larger triangular teeth bordering a broad, shallow, irregularly denticulate and crenulate medial emargination (Fig. 20.208c); sternite VIII with apical margin semicircularly rounded (Fig. 20.208d); median lobe of aedeagus with bulbus enlarged, tubus very short, with triangular apex, internal sac sclerites as illustrated (Fig. 20.208b). FEMALE. Tergite VIII transverse, truncate apically (Fig. 20.208e); sternite VIII arcuate apically (Fig. 20.208f); spermatheca sinuate, capsule with apical hemispherical extension to pitchershaped base, stem short and highly sinuate (Fig. 20.208g). DISTRIBUTION. Origin: Palearctic, now cosmopolitan, adventive in Canada and the USA. CANADA: AB, BC, NB, NS, ON, QC. USA:

20

Tribe Athetini Casey, 1910

CA, FL, LA, MA, NJ, NY. First North American record was before 1870 (Crotch 1870). COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Macrohabitat: in NB, common in compost (decaying corncobs and vegetables) near a mixed forest, under decaying seaweed on sea beach, and one from Lindgren funnel trap in a silver maple swamp. From other areas reported from rotting organic substances (grass heaps, carrion, mushrooms) and from forest litter. Collecting period: V, VII–IX. Collecting method: sifting litter and other decaying organic materials, pitfall traps, Lindgren funnel trap. Comments (Klimaszewski et al. 2018b). This species is a predator of small arthropods and it is used in North America as a biological control agent in greenhouses, and adults were observed invading breeder piles in commercial greenhouses and were likely consuming mites (Pochubay and Grieshop 2012). Drosophila suzukii Matsumura (Diptera: Drosophilidae) is a recent invasive fruit pest in North and South America and Europe, causing pre-harvest damage and significant economic losses to crops. Renkema et al. (2015) reported the predatory capability of D. coriaria Kraatz, a commercially available control agent against small dipteran pests, and developed molecular tools to assess predation. Dalotia coriaria showed a Type II functional response to all D. suzukii instars, with maximal consumption of 26 first, 15 second, and 6 third instars per beetle per day. No pupae were consumed. When raspberries with 11–21 larvae (2nd–3rd instar) per berry were exposed to six D. coriaria for 4.5 days, a 50% reduction in the Drosophila infestation occurred. Dalotia coriaria was also used for control of fungus gnats (Bradysia spp., Diptera: Sciaridae) in greenhouse crops, but larvae and adults may also prey on immature stages of thrips (Thysanoptera), shore flies (Scatella spp., Diptera: Ephydridae), cabbage root fly (Delia radicum L., Diptera: Anthomyiidae), carrot fly

413

(Chamaepsila rosae F., Diptera: Psilidae), and sap beetles (Nitidulidae) (Renkema et al. 2015).

Genus Dinaraea C.G. Thomson, 1858 (Figs. 20.209–20.212) (Klimaszewski et al. 2013b, 2018b) Diagnosis. Body subparallel, flattened, on average 3.0 mm long; integument with coarse meshed microsculpture, punctation distinct; head large, robust, subquadrate to slightly elongate, eyes usually shorter than genae in dorsal view, infraorbital carina absent; clypeus long, nearly horizontal, not depressed as in other genera of Athetini; mandibles strong, slightly curved apically, left one with margins smooth, inner margin of right one with small subapical tooth in apical one-third; maxillary palpi with four palpomeres, basal one small and apical one short and needle-shaped; laciniae and galeae moderately elongate, laciniae with fringe of long hairs subapically; labial palpi with three palpomeres; ligula V-shaped with each arm narrow; pronotum usually trapezoidal, widest in apical one-third in most species, disc with pubescence at midline directed posteriad in apical two-thirds, anteriad in basal one-third, and laterad elsewhere; tarsal formula 4-5-5, hypomera visible in lateral view; elytra short, at suture as long as pronotum or only slightly longer; first three visible abdominal tergites with distinct basal impression; apical margin of male tergite VIII without teeth or with 2–4 small teeth; median lobe of aedeagus of a simple form, with elongate structures in internal sac; spermatheca with capsule with subspherical apical part and very short, narrower basal part, stem long, narrow, coiled or looped posteriorly. Comments (Klimaszewski et al. 2018b). A number of species are known from the subcortical galleries of other insects but their diet is unknown. The adults and larvae may be either predatory on larvae of insects inhabiting galleries or feed on fungus growing inside galleries.

414

Key to Species of Dinaraea 1. Pronotum with posterior margin narrowly explanate, hind angles highly angulate (Fig. 20.212a); median lobe of aedeagus with enlarged bulbus, tubus thin apically and slightly produced ventrally at apex in lateral view (Fig. 20.212b), internal sac structures as illustrated (Fig. 20.212b); male tergite VIII without apical teeth (Fig. 20.212c) ............................. ................... Dinaraea subdepressa (Bernhauer) – Pronotum slightly impressed along posterior margin, not explanate, hind angles rounded (Figs. 20.209a–20.211a); genitalic structures different ............................................................ .2 2. Male tergite VIII with four large teeth (Fig. 20.209c); stem of spermatheca tightly coiled posteriorly (Fig. 20.209g) .................................... .........................Dinaraea angustula (Gyllenhal) – Male tergite VIII without or with minute teeth (Figs. 20.210c and 20.211c); stem of spermatheca broadly looped posteriorly (Figs. 20.211a)........3 3. Body length 2.2–2.5 mm; lateral margin of elytra straight (Fig. 20.211a); tubus of median lobe of aedeagus moderately long, highly sinuate ventrally in lateral view (Fig. 20.211c); male tergite VIII with minute apical teeth (Fig. 20.211c); spermathecal stem with broad posterior loop (Fig. 20.211e)....................................................... ....... Dinaraea pacei Klimaszewski and Langor – Body length 2.8 mm; lateral margin of elytra arcuate (Fig. 20.210a); tubus of median lobe of aedeagus moderately long, nearly straight ventrally in lateral view (Fig. 20.210b), internal sac structures as illustrated (Fig. 20.210b); male tergite VIII without apparent teeth, margin sinuate (Fig. 20.210c); female unknown.......................... ....... Dinaraea inexpectata Klimaszewski, sp. n.

173. Dinaraea angustula (Gyllenhal, 1810) { (Fig. 20.209a–g) Aleochara angustula Gyllenhal, 1810. Fauvel 1889, Muona 1984, Klimaszewski et al. 2007a, 2011, 2012, 2013a, b, 2016d, 2018b, Webster

20

et al. 2009, 2008b, 2010.

Majka

Tribe Athetini Casey, 1910

and

Klimaszewski

Diagnosis. Body length 3.3–3.7 mm; colour dark brown with legs, antennae at least basally, labial palpi and most of elytra yellowish- or reddish-brown, pronotum and abdomen lighter than head, sometimes entire body appears dark brown (Fig. 20.209a); head, pronotum and elytra slightly glossy, with dense microsculpture; abdominal microsculpture less dense and integument more glossy than that of pronotum and elytra; head almost as broad as pronotum, eyes slightly shorter than prominent genae in dorsal view; antennomeres VII–X moderately transverse; pronotum broadest at middle, slightly transverse, slightly longer than elytra at suture; elytra transverse, truncate posteriorly; abdomen subparallel. MALE. Tergite VIII with four small ridge-like teeth protruding slightly beyond apical margin and additional smaller subapical ridges on disc (Fig. 20.209c); sternite VIII parabolically rounded from base to more narrowly rounded apex (Fig. 20.209d); tubus of median lobe of aedeagus with ventral margin straight, apicodorsal part curved, slightly bent ventrad at apex in lateral view (Fig. 20.209b). FEMALE. Tergite VIII arcuate apically (Fig. 20.209e); sternite VIII broadly rounded apically, antecostal suture arcuate (Fig. 20.209f); spermatheca with short pearshaped asymmetrical capsule with small apical invagination, stem long, rectangularly bent at base of capsule, straight to tight posterior coil, base slightly swollen (Fig. 20.209g). DISTRIBUTION. Origin: Palearctic, adventive in Canada and the USA. CANADA: AB, BC [new provincial record], LB, NB, NF, NS, ON, PE, QC, SK, YT. USA: CA, NY, PA. We report this species from BC for the first time here based on the following: British Columbia, Ladner, Mathew road, 15.V.1961, coll. A.T. Wilkinson (UBC) 1 male, 1 female; Osoyoos IR1, 49.04N, 119.29W, 7.VII.1995/9. VIII.1995, pitfall trap, Purshia association, coll. G.G.E. Scudder (UBC) 1 male.

20

Tribe Athetini Casey, 1910

415

COLLECTION AND HABITAT DATA. Habitat: in variety of habitats such as open dry area with Purshia association (BC), agricultural fields and disturbed urban meadows, cattail and sedge marshes, mixed forests, coastal dunes, and lakeshores. Adults in compost (decaying corncobs and cornhusks and vegetables), soil and organic debris, amongst vegetation on a coastal sand dune, in litter in a cattail marsh, in leaf litter along a vernal pond, and in drift material along a lakeshore. Collecting period: IV–IX. Collecting method: sifting organic litter, pitfall traps. Comments. This Palaearctic species is adventive in North America (Klimaszewski et al. 2013a). The earliest Canadian records are: Elora, ON in 1975; St. Andrews, NB in 1978; Fairview, Alberta in 1982. The first North American records are: Buffalo, NY and New York, NY before 1889 (Fauvel 1889); Davis, CA before 1984 (Muona 1984). 174. Dinaraea inexpectata sp. n. (Fig. 20.210a–d)

Klimaszewski,

Holotype (male): Canada, British Columbia, Windermer Valley, 50 43.400 N, 116 07.320 W, 26.VII-17.IX.2000, pitfall #12, coll. R. Sargent (UBC). Etymology. The species name is derived from “inexpectatus”, a Latin word for unexpected. Diagnosis. Body length 2.8 mm; colour light brown with slightly darker head and abdomen (Fig. 20.210a); integument moderately glossy, with dense hexagonal microsculpture; abdominal microsculpture less dense and integument more glossy than pronotum and elytra; head rounded, slightly narrower than pronotum, eyes about as long as genae in dorsal view; antennomeres VII– X slightly transverse; pronotum arcuate laterally, broadest in middle, slightly transverse, as long as elytra at suture; elytra transverse, truncate posteriorly and sinuate laterally; abdomen subparallel. MALE. Apical margin of tergite VIII sinuate,

with two minute elevations medially, barely protruding at margin (Fig. 20.210c); apical margin of sternite VIII parabolic posteriorly (Fig. 20.210d); median lobe of aedeagus in lateral view with ventral part of tubus slightly sinuate (appearing almost straight), apical part broadly rounded (Fig. 20.210b); internal sac without distinct structures (Fig. 20.210b). FEMALE. Unknown. DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NF, QC, SK, YT. USA: AK. COLLECTION AND HABITAT DATA. Habitat: unspecified mountain habitat. Collecting period: VII–IX. Collecting method: pitfall trap. 175. Dinaraea pacei Klimaszewski Langor (Fig. 20.211a–g)

and

Dinaraea pacei Klimaszewski and Langor, in Klimaszewski et al. 2011. Klimaszewski et al. 2013b, 2015a, 2016d, 2018b. Diagnosis. Body length 2.2–2.5 mm; colour dark brown with legs, antennae (at least basally), palpi and elytra slightly paler and appearing reddish-brown, pronotum and abdomen lighter than head, sometimes entire body appears brown (Fig. 20.211a); head, pronotum and elytra slightly glossy, with dense microsculpture; abdominal microsculpture less dense and integument more glossy than pronotum and elytra; head about as large and broad as pronotum, eyes slightly shorter than genae in dorsal view; antennomeres VII–X moderately transverse; pronotum broadest in apical third, slightly transverse, slightly shorter than elytra at suture; elytra transverse, truncate posteriorly; abdomen subparallel. MALE. Apical margin of tergite VIII oblique laterally, obtusely angulate and slightly projecting on each side of broadly truncate apex, with two low ridge-like teeth (elevations) medially, barely protruding at margin (Fig. 20.211c); apical margin of sternite VIII arcuate, slightly truncate medially (Fig. 20.211d); median lobe of aedeagus in lateral

416

view with tubus short, sinuate basally, broadly arched ventrad at middle, apical half very narrow, slightly curled dorsad at apex (Fig. 20.211b); internal sac with elongate, sinuate structures (Fig. 20.211b). FEMALE: apical margin of tergite VIII arcuate, slightly emarginate medially (Fig. 20.211e), sternite VIII broadly rounded apically, antecostal suture slightly sinuate (Fig. 20.211f); spermatheca with short pearshaped capsule with small apical invagination, stem short, curved, with single broad loop posteriorly, slightly swollen at base (Fig. 20.211g). DISTRIBUTION. Origin: Nearctic. CANADA: BC, LB, NB, NF, QC, SK, YT. USA: AK. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NF and LB, from pitfall traps and flight intercept traps in various coniferous forest types; one specimen from under bark of red pine; in NB, adults under bark of large fallen spruce in old-growth eastern white cedar swamp, under tight bark of American elm in a silver maple forest, in fleshy polypore fungi at base of dead standing Populus sp. in a wet alder swamp, and in a group of Pholiota sp. at base dead standing Populus sp. in a mixed forest. Adults also from Lindgren funnel traps in old-growth white spruce and balsam fir forest, an old mixed forest, and a rich Appalachian hardwood forest with some conifers. In BC, from emergence traps attached to the trunks of lodgepole pine (Pinus contorta Dougl. ex Loud. latifolia Engelm.) infested by mountain pine beetle (Dendroctonus ponderosae Hopkins. Collecting period: III–IX. Collecting method: hand searching under bark, sifting fungi, Lindgren funnel traps, pitfall traps, flight intercept traps, emergence traps attached to tree trunks. Comments. Klimaszewski et al. (2016d) listed this species from AB, ON, PE, but we could not find voucher specimens representing these records and therefore we removed these provinces from the official distribution list for this species.

20

176. Dinaraea (Fig. 20.212a–g)

Tribe Athetini Casey, 1910

subdepressa

(Bernhauer)

Atheta (Dinaraea) subdepressa Bernhauer, 1907. Klimaszewski et al. 2013b, 2018b, Webster et al. 2016. Diagnosis. Body length 2.2–2.5 mm; colour dark brown to black, with legs, antennae (at least basally), palpi and edges and sutural parts of elytra slightly paler and appearing reddish-brown (Fig. 20.212a); head, pronotum and elytra slightly glossy, with dense microsculpture; abdominal microsculpture less dense and integument more glossy than pronotum and elytra; head as large and almost as broad as pronotum, eyes slightly shorter than genae in dorsal view; antennomeres VII–X moderately transverse; pronotum broadest in apical third, slightly transverse, slightly shorter than elytra at suture, posterolateral angles of the very angulate; elytra transverse, truncate posteriorly; abdomen arcuate laterally, broadest in middle. MALE. Tergite VIII truncate apically, without apical teeth (Fig. 20.212c); sternite VIII arcuate apically and slightly pointed medially (Fig. 20.212d); median lobe of aedeagus with enlarged bulbus and short and sinuate tubus, narrow subapically, apex pointed ventrally in lateral view (Fig. 20.212b); internal sac with large elongate structures (Fig. 20.212b). FEMALE. Tergite VIII arcuate apically (Fig. 20.212e), sternite VIII rounded apically, antecostal suture slightly sinuate (Fig. 20.212f); spermatheca with short pearshaped capsule with small apical invagination, and stem short and looped posteriorly, with slightly swollen apical part (Fig. 20.212g). This species is externally similar to D. curtipenis and D. pacei but differs in having the posterolateral angles of the pronotum very angulate, with the posterior margin narrowly explanate. In D. curtipenis, the posterior angle is rounded and the posterior margin is not explanate. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], NB, ON [new provincial record]. USA: NH.

20

Tribe Athetini Casey, 1910

We report this species from BC and ON for the first time here based on the following: British Columbia, Monashee Mtn., near Cherryville, 1400–1600 m, 9.VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 male; British Columbia, Creston, near Kootenay Pass, 1500–1800 m, 19. VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 male, 1 female; 15 km W Creston, 18. VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 1 female. Ontario, Algoma Co., Lake Superior Provincial Park, 2.IX.1980, leg. R. Baranowski, BWRS (ZMLU) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: all specimens from NB were captured in Lindgren funnel traps in the following forest types: a red pine forest, an old jack pine forest, an old-growth eastern white cedar forest, an old black spruce forest, mixed forests, a red oak forest, and an old balsam poplar forest near a river. Nothing else is known about its biology and microhabitat requirements but it probably lives in subcortical habitats like some other members of the genus. Collecting method: V–VI, VIII–IX. Collecting method: Lindgren funnel traps. Genus Dochmonota C.G. Thomson, 1859 (Fig. 20.213a–h) (Gusarov 2003a; Klimaszewski et al. 2011, 2018) Diagnosis. Body elongate, moderately broad; head subquadrate, eyes large, much longer than genae in dorsal view; antennomere II as long as III, IV subquadrate, V–X moderately transverse, last antennomere as long as three preceding ones combined; mandibles broad, medial tooth of right mandible inconspicuous; ligula with wide base and split into two lobes in apical one-third; pronotum transverse and widest at middle, with microsetae directed posteriad along midline of disc and slightly obliquely posteriad elsewhere, pronotal macrosetae moderately long, pronotal hypomera fully visible in lateral view; mesocoxae broadly separated; mesolateral macroseta of middle tibia inconspicuous, shorter than tibial width;

417

hind tarsi with article I slightly longer than II, each tarsus with one empodial seta; first three visible abdominal tergites each with moderately transverse basal impression, tergite VII as long as tergite VI; punctation on tergites VI–VII sparser than on tergites III–V, tergite VII with white palisade fringe; median lobe of aedeagus with apical process straight or slightly bent ventrally in lateral view; spermatheca with capsule mushroom-shaped, stem thick and tightly coiled posteriorly. 177. Dochmonota rudiventris (Eppelsheim) { (Fig. 20.213a–h) Homalota rudiventris Eppelsheim, 1886. Casey, 1910 [as Dimetrota revoluta and Datomicra vaciva], Muona 1984, Gusarov 2003a [synonyms], Webster et al. 2009, Majka and Klimaszewski 2008b, Klimaszewski et al. 2011, 2013a, 2016d, 2018. Diagnosis. Body narrowly elongate, subparallel, length 2.2–2.5 mm; colour dark brown to black, often with antennae, legs and posterior elytra reddish-brown (Fig. 20.213a), moderately glossy, punctation fine, pubescence brownish, microsculpture present on forebody; head round, distinctly narrower than pronotum, eyes moderately large; antennomeres IV–X subquadrate to slightly transverse; pronotum slightly transverse, widest at middle; elytra distinctly wider than pronotum, at suture as long as pronotum; abdomen broadly arcuate laterally. MALE. Tergite VIII truncate apically (Fig. 20.213d); sternite VIII produced apically and pointed medially (Fig. 20.213e); in lateral view median lobe of aedeagus with large bulbus and short tubus, tubus sinuate basally and narrowed apically in lateral view, crista apicalis enlarged (Fig. 20.213b), in dorsal view bulbus long, narrowly oval, tubus short and wide (Fig. 20.213c). FEMALE. Tergite VIII truncate apically and with small apical emargination (Fig. 20.213f); sternite VIII rounded and with minute apical emargination (Fig. 20.213g); spermatheca with small

418

mushroom-shaped capsule, apical invagination deep and narrow, stem thick and tightly coiled posteriorly (Fig. 20.213h). DISTRIBUTION. Origin: Possibly Holarctic. CANADA: BC [new provincial record], NB, NF, NT QC, YT. USA: ID, MA. We report this species from BC for the first time here based on the following: British Columbia, Smithers perimeter trail wetland, S of Hwy 16, 490 m, 54 46.220 N, 127 09.130 W, 31. VII.2012, in moss in swamp, A. Davies (CNC) 1 female. COLLECTION AND HABITAT DATA. Habitat: mixed forests and silver maple swamps (flood plain forests) in moist leaf litter near the margins of vernal ponds and in flood debris along rivers, in moss in swamp; similar to habitats reported from Europe. Collecting period: IV– VIII. Collecting method: sifting leaf litter. Comments. In view of its wide distribution in the Palearctic region (Smetana 2004), its wide distribution in Canada (BC, NF, NB, QC, west to NT and YT), and its occurrence in several natural wetland habitats, which are not typical of adventive species, it is likely that this is a Holarctic species. There are three additional species of Dochmonota described from Saskatchewan, not reported from BC (Klimaszewski et al. 2016d).

Genus Earota Mulsant and Rey, 1873b (Fig. 20.214a–h) (Gusarov 2002a; Klimaszewski et al. 2018b) Diagnosis. Body elongate, moderately broad, length 3.0–4.5 mm (average 3.0 mm); head subquadrate, eyes large, longer than genae in dorsal view; antennomere II as long as III, IV subquadrate, V–X moderately transverse, last antennomere as long as three preceding ones combined; mandibles broad, medial tooth of right mandible inconspicuous; ligula with wide base and split into two lobes in apical one-third;

20

Tribe Athetini Casey, 1910

infraorbital carina present; pronotum transverse and widest at middle, with pubescence short and dense, appearing silky, directed posteriad along midline of disc and slightly obliquely posteriad elsewhere, pubescence, pronotal hypomera fully visible in lateral view; mesocoxae broadly separated; mesolateral macroseta of middle tibia inconspicuous, shorter than tibial width; hind tarsus with article I slightly longer than II, each tarsus with one empodial seta; first three visible abdominal tergites with moderately transverse basal impression, tergite VII as long as tergite VI; punctation on tergites VI–VII sparser than on tergites III–V, tergite VII with white palisade fringe; median lobe of aedeagus with apical process straight or slightly bent ventrally in lateral view; spermatheca with dorso-ventrally compressed capsule, stem sinuate and highly swollen posteriad. For illustrations of mouthparts see Gusarov (2002a). 178. Earota 20.214a–h)

dentata

(Bernhauer) (Fig.

Atheta (Liogluta) dentata Bernhauer, 1906. Casey 1906 [as Macroterma alutacea and M. borealis], Casey 1910 [as Atheta (Macroterma) iowensis], Gusarov 2002a; Klimaszewski and Winchester 2002, Klimaszewski et al. 2005, 2007b, 2008a, b, 2011, 2016d, 2018b, Webster et al. 2009, Majka and Klimaszewski 2008b, Bernh 1907. Diagnosis. Body narrowly elongate, subparallel, slightly flattened, length 3.0–4.5 mm; colour dark brown, elytra except for scutellar region, legs, and antennomeres I–II rusty-brown (Fig. 20.214a); integument of forebody moderately glossy, with fine, sparse punctation, short pubescence and meshed microsculpture; head subquadrate, distinctly narrower than pronotum; antennae slender and long, antennomere IV subquadrate, V–X moderately transverse, last antennomere as long as three preceding ones combined; pronotum impressed medially, moderately transverse, widest at middle; elytra wider than pronotum, at suture slightly longer than pronotum, with no

20

Tribe Athetini Casey, 1910

apparent emargination near angular posterior angles; abdomen subparallel, tergites glossy and with fine punctation and microsculpture. MALE. Tergite VII with short medial carina before posterior margin; apical margin of tergite VIII obtusely angulate, with shallow notch halfway to apex and a shallow angular notch at apex (Fig. 20.214d); sternite VIII parabolically rounded from base to apex (Fig. 20.214e); median lobe of aedeagus in lateral view with tubus evenly arcuately curved ventrad, narrow apically, apex narrowly rounded (Fig. 20.214b), in ventral view bulbus oval and tubus triangular apically (Fig. 20.214c). FEMALE. Tergite VIII arcuate apically (Fig. 20.214f); apical margin of sternite VIII semicircularly rounded (Fig. 20.214g); spermatheca with capsule broad, flattened, stem short, looped and distinctly swollen posteriad (Fig. 20.214h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NF, NS, ON, QC, SK, YT. USA: AK, AL, CA, CO, DC, IA, ID, IL, KS, IN, MA, ME, MD, MT, NC, NH, NJ, NM, NV, NY, OH, OR, PA, VA, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: occurs in various habitats; in NB, adults from leaf litter near brook margin in red maple swamp, moss and leaf litter along small shaded brooks in mixed forests, under sea wrack on sea beach, among decaying corncobs and cornhusks (common); flying adults collected with butterfly net late in afternoon (16:00–18:00 h) early in spring (April), and some from pitfall traps in red spruce forest. Elsewhere, it has been reported from leaf litter, river debris, and pocket gopher (Geomys sp.) burrows. Collecting period: IV–IX. Collecting method: sifting organic debris, pitfall traps.

Genus Geostiba C.G. Thomson, 1858 (Fig. 20.215a–h) (Lohse and Smetana 1988; Gusarov 2002b; Klimaszewski et al. 2018b; Assing 2001)

419

Diagnosis. Body length 1.7–3.2 mm, subparallel, dark brown to yellow; head subquadrate, eyes reduced and much shorter than genae in dorsal view (Fig. 20.215a), greatly reduced in most species; mandibles symmetrical and broad, right mandible with a small medial tooth, ventral molar area without patches of denticle; ligula divided into two separate contiguous lobes; antennomere II longer than III, IV subquadrate, V–X transverse, XI as long as two preceding antennomeres combined; pronotum with microsetae directed posteriad along midline of disc, macrosetae short, pronotal hypomera fully visible in lateral view; most species wingless, with shortened elytra; mesocoxae contiguous; process of mesoventrite long and wide, extended to half length of mesocoxal cavities; middle tibia with mesolateral macroseta short, erect, hind tarsus with basal article longer than second one, tarsi each with one empodial seta. For illustrations of mouthparts, see Gusarov (2002b). Comments. Geostiba differs from the species of Atheta with similar pronotal pubescence and short elytra in having the ligula divided into two separate lobes (Gusarov 2002b). Geostiba is similar to Ousipalia in having the ligula divided into two separate lobes, but in Geostiba these lobes are contiguous while in Ousipalia they are widely separated (Gusarov 2002b). (Subgenus Geostiba C.G. Thomson, 1858) (Gusarov 2002b) Diagnosis. Male pronotum with median tubercle at posterior margin; each elytron with tubercle near scutellum; tergite VII with one short medial carina in front of posterior margin. 179. Geostiba horwoodae Klimaszewski and Godin, sp. n. (Fig. 20.215a–h) Holotype (male): Canada, British Columbia, Gulf Island, Saltspring Island, 48.8678-123.4739, 19.III.2012, moss/douglas fir, Berlese, coll. Darren Copley (RBCM). Paratypes: Canada, British Columbia, Gulf Island, Saltspring Island,

420

48.8678-123.4739, 19.III.2012, moss/douglas fir, Berlese, coll. Darren Copley (RBCM) 3 females; BC, Nanaimo, Yellowpoint lodge, 49.0219123.451, 22 m, 34.IV.2012, Moss/fir litter, Berlese, coll. Darren and Claudia Copley (RBCM) 2 females; BC, East Sooke Park, 48.3254-123.6434, 79 m, Berlese, moss, coll. Darren Copley (RBCM) 1 male; BC, Mechosin, Camosun College, 48.3789-123.603, 160 m, Berlese, moss, coll. Darren Copley (RBCM) 1 female. Excluded from paratypes: BC, 13 km N Topley, 54.616-126.3, 1132 m, VIII-IX.2012, pitfall # 389, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 1 female. Etymology. This species name is dedicated to Mrs. D. Horwood, wife of Mr. B. Godin, for her dedication to collect aleocharine beetles in western Canada. Her specimens lead to discovery of many new provincial records and new species. Diagnosis. Body subparallel, length 2.0–2.4 mm; head and most of abdomen dark brown, and pronotum, elytra and appendages yellowish brown (Fig. 20.215a); surface of forebody with fine isodiametric microsculpture, punctation very fine, sparse, punctures separated by 3–4 times their diameter; head subquadrate; eyes about half length of genae in dorsal view; antennomere II longer than III, and IV–X strongly transverse, XI as long as IX and X combined; pronotum slightly transverse, slightly wider than head; elytra at suture shorter or about as long as pronotum along midline of disc, wider than long (1.2), with fine isodiametric microsculpture and fine, somewhat asperate punctation, punctures separated by 2–3 times their diameter; abdomen subparallel, abdominal terga with fine, scarcely visible, microsculpture of transverse meshes, with fine and sparse punctation, punctation becoming finer towards abdomen apex. MALE. Tergite VIII with apical margin arcuate, entire (Fig. 20.215d); sternite VIII parabolic apically (Fig. 20.215e); median lobe of aedeagus in lateral view with tubus arched ventrad near apical fourth of ventral margin, apical part moderately broad, apex rounded (Fig. 20.215b); in dorsal view bulbus narrowly oval, tubus triangular apically

20

Tribe Athetini Casey, 1910

(Fig. 20.215c). FEMALE. Tergite VIII with posterior margin arcuate (Fig. 20.215f); sternite VIII broadly rounded apically (Fig. 20.215g); spermatheca with club-shaped capsule, stem double coiled and swollen posteriorly (Fig. 20.215h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: moss in Douglas fir forest, moss/fir litter. Collecting period: III, IV, VIII, IX Collecting method: sifting moss and forest litter and processing through Berlese funnel, pitfall traps. Comments. One female, missing the spermatheca and from 13 km N Topley was excluded from the paratype series.

Genus Goniusa Casey, 1906 (Fig. 20.216a–g) (Casey 1906; Fenyes 1918; Gusarov 2003b; Maruyama and Klimaszewski 2006) Diagnosis. Body length 3.2–4.2 mm, body broad; antennomeres V–X slightly elongate or slightly transverse; ligula split apically; labial palpus with setae α, β, γ and δ present; pronotum with basal angles of disc strongly angulate and basal margin arcuate, strongly transverse, at least two times as wide as long, with microsetae directed posteriorly along the midline and in lateral parts of the disc microsetae directed posteriorly and obliquely laterally; pronotal macrosetae short; pronotal hypomera fully visible in lateral view; medial macroseta of mesotibia inconspicuous, shorter than tibial width; tarsal formula 4-55; basal metatarsomere slightly shorter than second; one empodial seta present; abdominal sterna with numerous semierect macrosetae; sternite VIII with 30–60 macrosetae; male pronotum with broad medial impression that is half as wide as pronotum, deeper in the posterior half, postero-lateral portions of the impression with less dense microsculpture, without punctation

20

Tribe Athetini Casey, 1910

and pubescence; male sternite VIII with broad apical emargination; median lobe of aedeagus with narrow but blunt apex in dorsal view; medial sclerites of internal sac absent; flagellum troughshaped, with pointed apex; posterior portion of spermatheca with 2–3 coils. According to Gusarov (2003b), Goniusa can be distinguished from similar-looking genera of the tribe Lomechusini by having longer and narrower mesothoracic process, by shorter galea with its apical lobe only slightly projecting beyond the apex of lacinia, and by stronger setae on its internal margin.

421

V-shaped emarginated apically (Fig. 20.216d); tubus of median lobe of aedeagus truncate apically in dorsal view, in lateral view tubus sinuate ventrally and narrowly triangular apically (Fig. 20.216b). FEMALE. Tergite VIII rounded apically (Fig. 20.216e); sternite VIII truncate apically with shallow median emargination (Fig. 20.216f); spermatheca with capsule clubshaped, apical invagination deep, stem sinuate and narrowly multiple coiled posteriorly (Fig. 20.216g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB. USA: DC, MA, NH, NJ, NY, TX.

180. Goniusa caseyi Gusarov (Fig. 20.216a–g) Goniusa caseyi Gusarov, 2003b. Fenyes 1920, Maruyama and Klimaszewski 2006. Diagnosis. Length 3.2–4.2 mm; head darker than the rest of the body, brownish black to brown, pronotum, elytra and abdomen light brown to reddish brown, elytra and abdominal segments III–V often paler than pronotum and abdominal segments V–VII, legs, antennae and mouthparts brownish red (Fig. 20.216a); in females head surface glossy, with weak isodiametric microsculpture, with strong punctation, distance between punctures equals ½–1 times their diameter, and in males head surface matte, with strong isodiametric microsculpture, with weak, sparse and poorly visible punctation; temple length to eye length ratio 0.6–1.0; antennomeres IV–X slightly elongate; pronotum strongly transverse, with subparallel lateral sides, reaching maximum width in front of the middle, 1.6 times as wide as head; elytra slightly wider and longer than pronotum (pronotal length to elytral length ratio 0.9), 1.5 times wider than long, surface glossy, with fine and weak isodiametric microsculpture, and with fine and slightly asperate punctation, distance between punctures equal to their diameter; abdominal tergites glossy, with fine and often poorly visible transverse microsculpture, with fine punctation. MALE. Tergite VIII slightly emarginated apically, margin crenulate (Fig. 20.216c); sternite VIII deeply,

The only known locality data in AB and BC: Tp. 20, Rge. 3, W. 5 Mer., Alberta, 19.VI.1964, Lot 1, B.F. & J.L. Carr (LFC) 1 male, DNA voucher specimen CCDB-28936-G08; Gleneden, BC, 24.III.1978, Lot 4, B.F. & J.L. Carr (LFC) 1 female. COLLECTION AND HABITAT DATA. Habitat: Goniusa caseyi is associated with ants of the genus Formica (e.g., Formica integra (Nylander), F. rufa L.), some taken from ant nests under rocks. Collecting period: IV–VIII. Collecting method: aspirating from ant nests.

Genus Liogluta C.G. Thomson, 1858 (Figs. 20.217–20.223) (Klimaszewski et al. 2016a, 2018b) Diagnosis. Body narrowly subparallel, moderately flattened, length 2.8–5.4 mm; colour uniformly dark brown or reddish-brown with head and posterior part of abdomen dark brown to almost black; forebody with coarse meshed microsculpture, elytra often granulose; head with incomplete infraorbital carinae, genae moderately long to long; ligula apically split into two lobes; antennae long with antennomeres VI–X subquadrate, slightly transverse or rarely slightly elongate; pronotum with hypomera fully visible in lateral view; pubescence on disc of pronotum

422

directed posteriad on midline and lateroposteriad to laterad elsewhere; elytra and abdomen wider than head and pronotum; mesocoxae narrowly separated, process of metaventrite short and acute; legs long, three basal tarsomeres of hind tarsi highly elongate in most species, tarsal formula 5-5-5; apical margin of male tergite VIII often with broad, variably-shaped rectangular projection, with edge entire or bearing crenulation or variably shaped structures, with two lateral teeth and sometimes with additional median tooth; integument of disc often with broadly distributed asperate punctation near apex; female sternite VIII with apical margin rounded or medially emarginate, antecostal suture arcuate, or slightly to distinctly sinuate; median lobe of aedeagus simple with apical part variably shaped in lateral view; spermatheca with capsule narrowly club-shaped or tubular, with apical invagination moderate to deep, stem long, sinuate, and looped or twisted posteriorly. Key to Species of Liogluta 1. Antennae strongly to moderately swollen, antennomeres thick (Figs. 20.219a and 20.223a) ............................................................................2 – Antennae and antennomeres not as above ....3 2. Body narrowly subparallel, head dark brown to balck contrasting with uniformly red-yellow pronotum (Fig. 20.219a); median lobe of aedeagus strongly sinuate ventrally in lateral view (Fig. 20.219b); male tergite VIII traingularly produced and pointed medially (Fig. 20.219c); female sternite VIII distinctly emarginate medially at apex (Fig. 20.219f); spermathecal stem not twisted posteriorly (Fig. 20.219g)........................ ..............................Liogluta quadricollis (Casey) – Body broadly subparallel, head dark brown and not contrasting with uniformly red-brown pronotum (Fig. 20.223a); median lobe of aedeagus evenly arcuate ventrally in lateral view (Fig. 20.223b); male tergite VIII truncate apically (Fig. 20.223c); female sternite VIII with scarcely visible median emargination at apex

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Tribe Athetini Casey, 1910

(Fig. 20.221f); spermathecal stem twisted posteriorly (Fig. 20.223g)............................................. ................................. Liogluta wickhami (Casey) 3. Eyes small, each shorter than length of postocular region of head visible in dorsal view (Fig. 20.221a)..........Liogluta terminalis (Casey) – Eyes modertely large to large, each longer or equal to length of postoclur region of head visible in dorsal view.....................................................4 4. Body narrow (Fig. 20.222a); abdomen subparallel, distinctly narrower at base than elytra (Fig. 20.222a); male tergite VIII truncate apically, with two lateral teeth and serrate apical margin (Fig. 20.222d); bulbus of median lobe moderately large, tubus long with moderately broad apical part in lateral view (Fig. 20.222b, c); spermatheca with broad capsule and small and shallow apical invagination, stem sinuate and with moderately long posterior loop (Fig. 20.222h) ....................... ................................ Liogluta trapezicollis Lohse – Body and other characters not as above........ 5 5. Elytra broad, about 25% wider than maximum width of pronotum, eyes protruding laterally (Fig. 20.218a), integument strongly glossy; male tergite VIII truncate apically, not or moderately produced apically (Fig. 20.218c); spermatheca absent .......................... Liogluta nitens (Mäklin) – Elytra narrow, at most 20% wider than maximum width of pronotum, eyes not protruding laterally; integument in most species moderately glossy; male tergite VIII usually with welldefined lateral teeth, and with or without crenulation on apical margin ......................................... 6 6. Body length 2.8 mm; broadly subparallel, slightly glossy (Fig. 20.217a); male tergite VIII truncate apically and evenly serrated, not pointed medially (Fig. 20.217c); median lobe of aedeagus as illustrated (Fig. 20.217b); female undescribed ................................ Liogluta atriventris (Casey) – Body length 3.7–4.0 mm; narrowly subparallel, strongly glossy (Fig. 20.220a); male tergite VIII broadly rounded apically (Fig. 20.220c);

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Tribe Athetini Casey, 1910

423

median lobe of aedeagus as illustrated (Fig. 20.220b)....................................................... ............. Liogluta scudderi Klimaszewski, sp. n.

Microgranulosa Species Group (Klimaszewski et al. 2016a) Diagnosis. This group of species is characterized by a medium- to large-sized subparallel body (length 2.8–5.4 mm), large and bulging eyes, diameter of eye about as long as postocular area of head in dorsal view; integument of forebody moderately glossy; elytra at suture at least as long as pronotum, and elytra about 20–24% broader than pronotum; elytra in most species are densely and finely granulose; male tergite VIII truncate apically and apical margin straight, entire, serrate or pointed medially; tubus of median lobe of aedeagus more-or-less arched, and in lateral view narrow to moderately wide apically; spermatheca S-shaped with capsule bearing a deep or short invagination. 181. Liogluta 20.217a–d)

atriventris

(Casey) (Fig.

Athetota atriventris Casey 1906: 336. As Liogluta: Moore and Legner 1975, Klimaszewski et al. 2016a. Diagnosis (based on male lectotype). This species may be distinguished by the following combination of characters: small body size, length 2.8 mm; head, and abdomen dark brown, and pronotum, elytra and legs reddish yellow (Fig. 20.217a); integument strongly glossy with weak meshy microsculpture; moderately dense punctation and pubescence on pronotum and elytra, elytra with very fine micro-granulation; head subquadrate, slightly narrower than pronotum; large eyes, each about as long as postocular region in dorsal view; antennae of the lectotype are partially damaged and cannot be described; pronotum slightly transverse, widest near the apical third; elytra wider and slightly longer than pronotum; abdomen subparallel, about as wide as elytra. MALE. Tergite VIII

with apical margin dentate (Fig. 20.217c); sternite VIII broadly rounded apically (Fig. 20.217d); median lobe of aedeagus with arcuate ventral margin of tubus and narrowly elongate apical part in lateral view (Fig. 20.217b). FEMALE. Unknown. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. This species is known only from a damaged lectotype male specimen (Victoria, Vancouver Island). More specimens, including females, are needed for study to confirm the status of this species. It is similar to L. trapezicollis and may be conspecific with it. The only available type male specimen of L. atriventris is broader and more yellowish brown with longer elytra than those of L. trapezicollis from BC. An additional comparative DNA study of the type of L. atriventris and available specimens of L. trapezicollis from BC and AK are needed to unravel the true identity of this species. Nigropolita Species Group (Klimaszewski et al. 2016a) Diagnosis. This group of species has a body shape non-typical for Liogluta and it rather resembles some of the Atheta (Dimetrota) species with elytra distinctly wider than head and pronotum. This group is characterized by large and bulging eyes, diameter of eye about as long as postocular area of head in dorsal view; integument of forebody strongly glossy; elytra at suture at least as long as pronotum, and elytra about one-third broader than pronotum; male tergite VIII truncate apically and apical margin entire or slightly produced with rounded lateral teeth; tubus of the median lobe of the aedeagus is arched, or almost straight and in lateral view moderately narrow apically; spermatheca with spherical capsule with moderately long apical

424

invagination and sinuate stem. The spermatheca of L. nitens was not found and may be very small, not sclerotized, or absent. 182. Liogluta nitens (Mäklin) (Fig. 20.218a–f) Homalota nitens Mäklin, 1852: 307. Hatch 1957, Moore and Legner 1975, Lohse and Smetana 1985, Klimaszewski and Winchester 2002, Klimaszewski et al. 2016a. Diagnosis. Body length 2.8–3.3 mm, narrowly elongate, Atheta-like, moderately robust, dark brown to black with elytra, tarsi and tibiae often yellow- or red-brown (Fig. 20.218a); integument of forebody with moderately pronounced meshed microsculpture, surface strongly glossy; head about one-eighth narrower than maximum width of pronotum; pronotum transverse, about evenly wide until basal third of its length then becoming strongly narrowed apically (Fig. 20.218a); elytra at suture about as long as pronotum; basal four metatarsomeres about the same length and each shorter than fifth one. MALE. Tergite VIII truncate apically with apical part slightly produced and with rounded lateral teeth, apical margin smooth or with micro-crenulation (Fig. 20.218c); sternite VIII rparabolic apically (Fig. 20.218d); median lobe of aedeagus with tubus slightly sinuate ventrally and with moderately narrow and rounded apex (Fig. 20.218b). FEMALE. Tergite VIII arcuate apically (Fig. 20.218e); sternite VIII rounded apically and with sinuate antecostal suture (Fig. 20.218f). Spermatheca not found. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, YT. USA: AK, OR, WA. New locality data: BC, Queen Charlotte Is., Skowkona Cr., 7.9 mi NW Q.C. City, 29.VII-4. VIII.1983, 83-55A, ex rotting pile of mushrooms, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., Lyall Is., Gate Cr., 10. VIII.1983, 83-72, sifting wet moss on forest floor, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., 14 km E Masset, 13. VII.1983, 83-5, treading Carex marsh and sifting

20

Tribe Athetini Casey, 1910

Alnus, J.M. Campbell (CNC) 1 sex undetermined; Queen Charlotte Is., Moresby Is., Tekakia Lk., 23.VII.1983, 83-33, 21000 , sifting moss ex forest floor, J.M. Campbell (CNC) 3 sex undetermined; Queen Charlotte Is., 1 km S Moresby Camp, 20. VIII.1983, sifting moss at edge of old road, J.M. Campbell (CNC) 3 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: in BC, adults were found in the interior and transition zones of a Sitka spruce forest, Carmanah Valley, Vancouver Island (Klimaszewski and Winchester 2002), and in moss on forest floor and edge of ald road, in mushrooms, and in Carex marsh and Alnus litter on Queen Charlotte Is. Several adults were collected from moss at the edge of an old road in the Queen Charlotte Islands, British Columbia. Adults in Alaska were collected in a wide variety of habitats spanning lowland forests to alpine zones: alpine meadow litter, lowland forest clearcuts, floodplain meadows with Athyrium, Caltha, and Rubus, under rocks, in krummholz alpine habitats of Tsuga mertensiana, near bear dung in alpine habitats, old growth temperate rain coniferous forests, alpine heath with Empetrum, and Vaccinium, subalpine habitats with Salix, and Veratrum (Klimaszewski et al. 2016a). Collecting period: VI–IX. Collecting method: pitfall traps, sifting moss and mushrooms, and treading Carex marsh and sifting Alnus. Comments. There is considerable variation in length and width of the elytra in specimens from Vancouver Island, Oregon (having broader and longer elytra), and those with narrow and shorter elytra from the Queen Charlotte Islands, Alberta, and Alaska. The genitalic features were the same in those of the typical form with the longer and broader elytra, and those with narrower and shorter elytra. Therefore, we consider this as an intraspecific variation. Additional studies, including DNA comparison, are needed to reveal the relationship between these two morphotypes. Two UAM Alaskan specimens (UAM: Ento:152502, UAM:Ento:232546) were DNA barcoded (UAMIC2665-15, UAMIC2701-15)

20

Tribe Athetini Casey, 1910

425

and they cluster closely with two specimens of this species. DNA barcoded from Alberta, Canada (Klimaszewski et al. 2016a). Terminalis Species Group (Klimaszewski et al. 2016a, 2018b) Diagnosis. Eyes small, distinctly shorter than genae in dorsal view; integument of forebody moderately to highly glossy; elytra short, at suture about as long as pronotum; male tergite VIII truncate apically, with or without minute crenulations; median lobe of aedeagus with tubus usually arched and moderately narrow apically in lateral view; capsule of spermatheca with apical part compressed, with apical invagination small and short. 183. Liogluta 20.219a–g)

quadricollis

(Casey) (Fig.

Anepsiota quadricollis Casey 1893. Klimaszewski et al. 2016a. Holotype (female): Canada, British Columbia, Vancouver Island, Anepsiota quadricollis; Type USNM 39471 (USNM). Examined. Diagnosis. Body length 4.1–4.3 mm, subparallel, slender, bicoloured, pronotum, base and apex of abdone, 2–3 basal antennomeres, and legs orange and remainder of body dark brown to red-brown (Fig. 20.219a); integument of forebody with weak meshed microsculpture, surface strongly glossy; head slightly narrower than pronotum; pronotum subquadrate; antennae enlarged and black to brown, antennomeres VII– X slightly transverse; elytra about as wide as pronotum and at suture shorter or as long as pronotum; basal two metatarsomeres strongly elongate, co-equal in length, each longer than third one. MALE. Tergite VIII broadly traingularly produced apically and pointed medially (Fig. 20.219c); sternite VIII elongate, parabolic apically and with wide distance between basal margin and arcuate antecostal suture (Fig. 20.219d); median lobe of aedeagus with tubus short and strongly sinuate, its subapical

part narrowly elongate in lateral view, internal sac structures as illustrated (Fig. 20.219b). FEMALE. Tergite VIII slightly produced medially (Fig. 20.219e); sternite VIII shallowly emarginated medially (Fig. 20.219f); spermatheca with club-shaped capsule and short apical invagination, stem long and strongly sinuate and looped posteriorly (Fig. 20.219g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: one male was found under a rock in a poplar stand. Collecting period: IX. Collecting method: not recorded. 184. Liogluta scudderi Klimaszewski, sp. n. (Fig. 20.220a–g) Holotype (male): Canada, British Columbia, Valemont, Yellowjacket ck., 24-29.VI.2014, pitfall, Kinbasket # 1168, 52.703-119.049, 771 m, R-Mature, coll. Charlene Wood (RBCM). Paratypes: Canada, British Columbia, Valemont, Yellowjacket ck., 24-29.VI.2014, pitfall, Kinbasket # 1168, 52.703-119.049, 771 m, R-Mature, coll. Charlene Wood (RBCM) 1 female; Valemont, Peatland, 23-28. VI.2014, pitfall, Kinbasket # 1091, 52.762-119.164, 762 m, R-Mature, coll. Charlene Wood (RBCM) 2 females; Atlin, Pine creek, 21.VI.2015, 59.7116-133.5787, 939 m, Alder soil sifting, coll. B. Godin and D. Horwood (LFC) 1 male; Atlin, McDonald Lake, 20.VI.2015, 59.7116133.5787, 939 m, willow, soil sifting (BGC, LFC) 1 male, 1 female. Etymology. This species is dedicated to Prof. Geoff G.E. Scudder, emeritus professor of the University of British Columbia, who supported this study by providing aleocharine specimens from British Columbia. Diagnosis. Body length 3.7–4.0 mm, narrowly subparallel, colour dark brown to almost black, elytra slightly and legs reddish-brown

426

(Fig. 20.220a); integument of entire body strongly glossy, with meshed microsculpture; head elongate, about one-quarter narrower than maximum width of pronotum; antennomere IV subquadrate, VII–X distinctly transverse, last antennomere as long as two preceding ones combined; pronotum strongly margined, trapezoidal in shape, slightly transverse, narrower at base and widest in apical third; elytra at suture about as long as pronotum; abdomen broad and arcuate laterally; hind tarsus with four basal tarsomeres about the same length. MALE. Tergite VIII with apical margin arcuate and smooth (Fig. 20.220c); sternite VIII rounded apically and truncate medially (Fig. 20.220d); tubus of median lobe of aedeagus in lateral view straight basally and arcuate apically, apex narrow and rounded (Fig. 20.220b). FEMALE. Tergite VIII arcuate apically (Fig. 20.220e); sternite VIII arcuate apically and truncate medially (Fig. 20.220f); spermatheca with club-shaped narrow capsule and arcuate neck, stem narrow, strongly sinuate, narrowly looped and twisted posteriorly (Fig. 20.220g). This species is distinct by its narrow and strongly glossy body, short elytra, by the arcuate apical margin of male tergite VIII, by the shape of the median lobe and the strongly sinuate stem of the spermatheca. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: mature unspecified forest and peatland. Collecting period: VI. Collecting method: pitfall traps. Comments. The specimens from Atlin, BC were much paler in colour than the remaining dark brown to near black specimens, but the genitalic features were the same. We consider the paler colour as intraspecific variation and included the pale specimens in the paratype series. 185. Liogluta terminalis (Casey) (Fig. 20.221a–g)

20

Tribe Athetini Casey, 1910

Anepsiota terminalis Casey, 1906. As L. aloconotoides: Lohse et al. 1990, Majka and Klimaszewski 2008b, Klimaszewski et al. 2008b, 2011, 2015a, Webster et al. 2012, Stefani et al. 2016. As L. terminalis: Klimaszewski et al. 2016a, 2018b. Diagnosis. Body length 3.9–4.5 mm, subparallel, colour entirely reddish-brown or with head and posterior abdomen chestnut brown (Fig. 20.221a); integument of forebody moderately glossy, with meshed microsculpture; head about one-quarter narrower than maximum width of pronotum; last antennomere, at least in males, as long as two preceding ones combined; pronotum trapezoidal in shape, slightly transverse, narrower at base and widest in apical third; elytra at suture about as long as pronotum; hind tarsus with three basal tarsomeres elongate, first longest, second about as long as third, fourth shorter than either of preceding ones. MALE. Tergite VIII with apical margin with broad, short, truncate projection with rounded lateral angles, apical margin smooth or slightly crenulate (Fig. 20.221c); sternite VIII parabolic apically (Fig. 20.221d); median lobe of aedeagus in lateral view arcuate ventrally, subapical part narrow, apex narrowly rounded (Fig. 20.221b). FEMALE. Tergite VIII arcuate apically (Fig. 20.221e); sternite VIII slightly emarginate medially at apex (Fig. 20.221f); spermatheca with broad club-shaped capsule, neck arcuate, stem long, looped and twisted posteriorly (Fig. 20.221g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, YT. USA: MT, NH. New locality data: British Columbia, 160 km W of Watson Lake, 27.VI.2015, 59.9056131.4924, 872 m, Firekill, deciduous, litter sifting, coll. B. Godin and D. Horwood (BGC, LFC) 1 male, 1 female; BColl., Driftwood Canyon Prov. Park, 54 49.650 N, 127 01.410 W, 752 m, 27.VII.2012, alder, poplar litter and moss along creek, A. Davies (CNC) 1 male, 1 female.

20

Tribe Athetini Casey, 1910

427

COLLECTION AND HABITAT DATA. Habitat: in NF, collected from pitfall traps in various forest types such as birch-dominated forests, spruce-fir forests, black spruce/lichen forests, fir forests, a recently burned forest, and from coastal limestone barrens; in NB, from dung in a coastal red spruce forest, Lindgren funnel trap in a rich Appalachian hardwood forest with some conifers, and treaded from sedges on lake margin (Klimaszewski et al. 2018b). In AB, some reared from well-decayed white spruce logs. In BC, poplar litter and moss along creek. Collecting period: VII–IX. Collecting method: pitfall traps, treading sedges along a small lake margin, Lindgren funnel trap. Comments. This is the most commonly encountered species of Liogluta in forest samples in central and eastern Canada. 186. Liogluta 20.222a–h)

trapezicollis

Lohse (Fig.

Liogluta (Anepsiota) trapezicollis Lohse, in Lohse et al. 1990. Klimaszewski et al. 2016a. Diagnosis. Body length 3.8–4.4 mm, subparallel, slender, dark brown to black with pronotum brown and paler than head, elytra with yellowbrown irregular spots (Fig. 20.222a); integument of forebody with meshed microsculpture moderately pronounced, surface moderately strongly glossy; head about one-quarter narrower than maximum width of pronotum; pronotum transverse, narrow at base and widest at middle (width of pronotum is variable, some specimens have pronotum markedly narrower than base of elytra and some have pronotum nearly as wide as base of elytra); elytra at suture slightly shorter than pronotum; basal three metatarsomeres elongate, equal in length and each slightly longer than fourth one. MALE. Tergite VIII truncate apically with two short lateral teeth, and apical margin variably sculptured and ranging from smooth to crenulate, or denticulate (Fig. 20.222d); sternite VIII parabolic apically (Fig. 20.222e); tubus of median lobe of aedeagus arcuate laterally and

with narrow apical part pointed at apex in lateral view (Fig. 20.222b, c). FEMALE. Tergite VIII rounded apically (Fig. 20.222f); sternite VIII truncate and slightly emarginate medially (Fig. 20.222g); spermatheca with club-shaped capsule bearing shallow invagination, stem narrow and sinuate, narrowly looped and twisted posteriorly (Fig. 20.222h). DISTRIBUTION. Origin: Nearctic. CANADA: BC [NPR], YT. USA: AK. We report this species from BC for the first time here based on the following [it was listed from BC by Klimaszewski et al. 2016a, without records]: British Columbia, Kootenay, Porcupine forest service road, 49.2646-117.1815, 820 m, 12. IX.2011, Berlese, moss, coll. R.G.B. Bennet, D. &C. Copley (RBCM) 1 male, 2 females; Mechosin, Pearson College, 48.3469-123.567, 10 m, 24.V.2014, Berlese, moss, coll. Darren and Claudia Copley (RBCM) 2 males, 2 females; 13 km N Topley, 54.616-126.3, 1132 m, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 2 males, 1 female; Log Lake, 65 km N Prince George, 54.2122.616, 780 m, VII-IX.2012, OM-COSX, sub-boreal spruce, pitfall # 26, 208, coll. J.M. Kranabetter (RBCM) 2 males; NE Genoa Bay, 48.7817-123.568.99, Berlese, moss, coll. Darren and Claudia Copley (RBCM) 1 male; Gulf Island, Galiano Is., 48.921-123.4449, 110 m, 9.V.2012, Berlese, moss, coll. R.G.B. Bennet (RBCM) 1 male. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2016a). Habitat: the BC specimens were collected from moss, and some from boreal spruce forest; the AK specimens were collected from alpine zones between 453 and 1071 m elevation, none were collected in lowland forests. Habitats include alpine flood meadows, under rocks, herbaceous heath with Luetkea, Cassiopes, and Lupinus, low rocky tundra with Dryas, meadow heath with Phyllodoce, Senecio, and Luetkea, shrubby krummholz with Elliottia and Tsuga, wet meadows with Carex, Petasites,

428

20

Senecio and Ranunculus. Collecting period: V, VII–IX. Collecting method: pitfall traps, treading sedges along a small lake margin, Lindgren funnel trap, processing moss and forest debris through Berlese funnels. Comments. We were not able to compare types of L. trapezicollis Lohse with the specimens we examined and our determinations are based on the published description by Lohse in Lohse et al. (1990). The types of L. trapezicollis housed in the Canadian National Collection of Insects were borrowed several years ago and our consistent efforts to have these specimens returned to Canada have failed. Five specimens in UAM were successfully DNA barcoded (UAM GUID, BOLD Process ID): UAM:Ento:145576, UAMIC2696-15; UAM:Ento:145623, UAMIC2740-15; UAM: Ento:152467, UAMIC2750-15; UAM:Ento: 232527, UAMIC2677-15; UAM:Ento:232696, UAMIC2753-15. The DNA sequences for these specimens are all very similar (maximum distance of 0.32%, nearest neighbor of 3.13%) and fall within the same BIN (Barcode Index Number (Ratnasingham and Hebert (2013), BOLD: ACU9772, which is not shared by any other species. There is a possibility that L. trapezicollis is conspecific with L. vasta described form Sitka Is., AK. The DNA analysis of the type specimens of the two species is essential to unravel the true identity of the two species. 187. Liogluta wickhami (Casey) (Fig. 20.223a–g) Anepsiota wickhami Casey Klimaszewski et al. 2016a.

1893:

Tribe Athetini Casey, 1910

robust, antennomere IV slightly elonagte, VII–X elonagte or subquadrate; pronotum more or less evenly arcuate laterally; elytra at suture about as long as pronotum; abdomen broad, at base as broad as elytra, sides arcuatebasal metatarsomere the longerst, the following two slightly shorter, and fourth sharter than penultimate one. MALE. Tergite VIII truncate apically and with faint crenulation (Fig. 20.223c); sternite VIII parabolic apically (Fig. 20.223d); tubus of median lobe of aedeagus short, arcuate ventrally, with apex narrow and rounded (Fig. 20.223b). FEMALE. Tergite VIII arcuate apically and truncate medially (Fig. 20.223e); sternite VIII slightly emarginate apically (Fig. 20.223f); spermatheca with tubular capsule and deep apical invagination, stem thin, long, and strongly sinuate, looped posteriorly and twisted at apex (Fig. 20.223g). This species is somewhat similar to L. terminalis but has thick dark brown antennae, head and pronotum (antennae, head, and pronotum are uniformly reddish brown or only slightly darker than remaining parts of the body in L. terminalis). Spermatheca is differently shaped in each species; L. wickhami has smaller and differently shaped capsule with a deep apical invagination and has a shorter and differently looped posterior stem. Liogluta wickhami is also very similar to L. vasta but can be distinguished by the shape of the pronotum which has evenly arcuate sides and is broadest at the middle, while it is trapezoidal in shape and is broadest in the apical third in the latter species. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded.

331.

Diagnosis. Body length length 4.0–4.2 mm, broadly subparallel; pronotum, elytra, legs and basal antennopmeres red-brown, head and most of abdomen chestnut brown (Fig. 20.223a); integument of forebody with meshed microsculpture, moderately glossy; head about one-third narrower than maximum width of pronotum; antennae

The female holotype was captured in the Stickeen River Valley of British Columbia (Casey 1893), and subsequently one male was found in Three Guardsmen Pass, British Columbia. New locality data: British Columbia, 13 km N Topley, VIII-IX.2012, pitfall # 389, 54.616126.3, 1132 m, OM2C2SX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 6 males, 9 females; Log Lk., 65 km N Prince George,

20

Tribe Athetini Casey, 1910

VIII-IX.2012, pitfall # 26, # 204, # 208, 54.2122.616, 780 m, OM-COSX, sub-boreal spruce, coll. J.M. Kranabetter (RBCM) 5 males, 5 females. COLLECTION AND HABITAT DATA. Habitat: sub-boreal spruce forest. Collecting period: VII, IX. Collecting method: pitfall traps.

Genus Lypoglossa Fenyes, 1918 (Figs. 20.224a–h and 20.225a–i) (Hoebeke 1992; Gusarov 2004; Klimaszewski et al. 2018b) Diagnosis. Body broad, ovoid, length 3.2–4.6 mm; colour uniformly brown to dark brown with appendages paler, pronotum and elytra reddish-brown in some specimens; head transverse, infraorbital carinae complete; a sensillum of epipharynx long; ligula with broad base, split apically into two lobes; labial palpi with setae α β, γ and δ present; antennomere II as long III, IV elongate, V elongate to subquadrate, VI–X elongate to transverse, apical antennomere without coeloconic sensilla and as long as IX and X combined; pronotum exceptionally broad, 1.5 times as wide as long, disc with microsetae directed anteriad along midline, laterad and posteriad elsewhere, macrosetae short and thin, hypomera incompletely visible or totally obscured in lateral view; posterior margin of elytra near posterolateral angle straight or slightly concave; mesocoxae narrowly separated; process of mesoventrite long and slender, acute at apex; hind tarsi with basal article much longer than second one, each tarsus with single empodial seta shorter than claws, mesolateral macroseta of middle tibia thin, much shorter than tibial width; first three visible abdominal tergites with weak transverse basal impression, densely and finely punctate with short, silky, dense apressed pubescence, tergites VI–VIII sparsely punctate and pubescent; median lobe of aedeagus broad in dorsal view with base of tubus about as wide as

429

bulbus; spermatheca S-shaped, capsule with large apical invagination, stem long and sinuate, narrowly looped posteriad. For illustrations of mouthparts and terminology see Hoebeke (1992) and Gusarov (2004). Species of Lypoglossa share an overall resemblance to Devia prospera (Oxypodini) but differ in having a 4-5-5 tarsal formula (5-5-5 in Devia). Comments. Members of Lypoglossa are restricted to the boreal forest zone and northern coniferous forests and coastal areas of western North America (Hoebeke 1992; Gusarov 2004). Key to Species of Lypoglossa 1. Pronotal width to length ratio 1.2–1.3 (Fig. 20.225a); median lobe of aedeagus with apex of tubus broadest at the base, not constricted in ventral view, and without projections at the base of the apex (Fig. 20.225c); spermatheca larger, posterior loop of stem wide, almost circular (Fig. 20.225h) ................................................. ..................... Lypoglossa franclemonti Hoebeke – Pronotal width to length ratio 1.3–1.4 (Fig. 20.224a); median lobe of aedeagus with apex of tubus constricted basally in dorsal/ventral view, and with dentiform projection on each side of the base of apex (Fig. 20.224c); spermatheca smaller, with posterior loop of stem narrow (Fig. 20.224h).....Lypoglossa angularis (Mäklin)

188. Lypoglossa 20.224a–i)

angularis

(Mäklin) (Fig.

Euryusa obtusa Mäklin, 1853. Hoebeke 1992 [as L. angularis, ex parte], Gusarov 2004, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, Webster et al. 2012. Diagnosis. Body robust, wide, oval, length 3.8–4.2 mm; head and abdomen brown to dark brown and pronotum, elytra, legs and antennae light rusty-brown; integument moderately glossy,

430

forebody with fine and dense punctation, short pubescence and meshed microsculpture (Fig. 20.224a); head transverse, distinctly narrower than pronotum; antennae thin, antennomeres IV–VIII elongate, IX–X subquadrate; pronotum widely oval, transverse, widest at basal third; elytra as wide as pronotum, at suture about as long as pronotum, posterior margin arcuate; abdomen subparallel, as wide as elytra and pronotum. MALE. Tergite VIII arcuate apically (Fig. 20.224e); sternite VIII parabolic apically (Fig. 20.224f); median lobe of aedeagus has wide bridge in lateral view (Fig. 20.224b), and two lateral rounded apical projections and two basal processes in ventral view (Fig. 20.224c, d). FEMALE: tergite VIII arcuate apically (Fig. 20.224g); sternite VIII rounded apically (Fig. 20.224h); spermatheca with posterior loop moderately narrow (Fig. 20.224i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NF, QC, YT. USA: AK, AZ, CA, CO, MA, ME, MT, NH, NM, OR, WA. COLLECTION AND HABITAT DATA. Habitat: gopher burrows; moss at edge of forest, forest litter in Betula, Picea and Alnus, Veratrum, ferns forest litter; treeline subalpine meadow (Gusarov 2004). Collecting period: IV–VIII. Collecting method: sifting forest litter and moss, pitfall traps. Comments. Gusarov (2004) recognized three subspecies of L. angularis, and distinguished them by morphology of the median lobe of the aedeagus. These differences relate to the wider or narrower athetine bridge and width of the apex of the median lobe in ventral view. By definition, the subspecies should be geographically isolated as closed gene pools but it is not clear in the case of these subspecies. These genital differences are not pronounced and may represent only geographical variations. In this book, we do not subdivide this species into subspecies, but BC specimens of angularis belong to the subspecies angularis according to Gusarov (2004).

20

189. Lypoglossa (Fig. 20.225a–i)

Tribe Athetini Casey, 1910

franclemonti

Hoebeke

Lypoglossa franclemonti Hoebeke, 1992. Gusarov 2004, Webster et al. 2009, Majka and Klimaszewski 2010, Klimaszewski et al. 2011, 2012, 2015a, Stefani et al. 2016. Diagnosis. Body robust, wide, oval, length 3.8–4.6 mm; head and abdomen dark brown, pronotum, elytra, legs and antennae light rustybrown (Fig. 20.225a); integument moderately glossy, forebody with fine and dense punctation, short pubescence and meshed microsculpture; head transverse, distinctly narrower than pronotum; antennae thin, antennomeres IV–VIII elongate (more elongate than in L. angularis), IX–X subquadrate; pronotum widely oval, transverse, widest at basal third (less transverse than in L. angularis); elytra as wide as pronotum, at suture about as long as pronotum, posterior margin arcuate; abdomen subparallel, as wide as elytra and pronotum. MALE. Tergite VIII arcuate apically (Fig. 20.225e); sternite VIII parabolic (Fig. 20.225f); median lobe of aedeagus narrowly oval, widest medially, not constricted, and without large projections at the base of tubus in dorsal view (Fig. 20.225c, d); athetine bridge narrow in lateral view (Fig. 20.225b). FEMALE. Tergite VIII arcuate apically (Fig. 20.225g); sternite VIII rounded apically (Fig. 20.225h); spermatheca with stem forming broader posterior loop (Fig. 20.225i). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], MB, NB, NF, NS, NT, ON, QC, SK, YT. USA: ME, NH, NY, VT. We report this species from BC for the first time here based on the following: British Columbia, Valemont, Yellowjacket ck., 3-8.VI.2014, pitfall, Kinbasket # 1053, 52.703-119.049, 771 m, R-Mature, coll. Charlene Wood (RBCM) 4 males, 11 females; same date except: Kinbasket # 1054 (RBCM) 1 female, Kinbasket # 1061 (RBCM) 3 females, Kinbasket # 1074 (RBCM)

20

Tribe Athetini Casey, 1910

1 female; 24-29.VI.24, pitfall, Kinbasket # 11683, 2 females (RBCM). COLLECTION AND HABITAT DATA. Habitat: coniferous and deciduous forests, balsam fir stand, spruce bogs, birch bogs; adults in forest litter, in moss and lichens; in NB, this species was found under sea wrack under alders on sea beach in May (4 individuals) and in gilled mushrooms in an adjacent red spruce and yellow birch forest in August and September (4 individuals). Collecting period: V–IX. Collecting method: sifting sea wrack and mushrooms, pitfall traps.

Genus Mocyta Mulsant and Rey, 1873b (Figs. 20.226–20.228) (Klimaszewski et al. 2015b, 2018b) Diagnosis. Antennae very thin and pale, in most specimens contrasting with body colour, antennomeres V–X subquadrate or slightly transverse; pronotum glossy, moderately convex, broad and shield-shaped, widest at or near middle, at least as broad as the base of the elytra but in most specimens slightly broader, pubescence on disc directed posteriad along midline (or throughout central section) and posterolaterad elsewhere, hypomera not visible in lateral view; male tergite VIII truncate apically, without teeth and other secondary sexual characters; male sternite VIII with longer macrosetae than those of females; median lobe of aedeagus of a simple form, tubus about half length of median lobe, narrowly tapering and rounded apically in dorsal view, straight and narrow apically in lateral view, internal sac structures inconspicuous, usually elongate and not pronounced; spermatheca with capsule hemispherical, or elongate and sac-shaped, usually with small apical invagination and short neck, stem thin and regularly or irregularly coiled posteriorly. Key to Species of Mocyta 1. Pronotum distinctly broader and larger than elytra (Fig. 20.228a); elytra at suture shorter

431

than pronotum along midline (Fig. 20.228a) ....... ...................................... Mocyta discreta (Casey) – Pronotum as broad as elytra and at most as large as elytra (Figs. 20.226a and 20.227a); elytra at suture at least as long as pronotum along midline (Figs. 20.226a and 20.227a) .......................2 2. Body robust and broad (Fig. 20.227a); elytra variable in length but distinctly longer than pronotum in most [some specimens have elytra as long as pronotum] (Fig. 20.227a); pronotum approximately rectangular, gradually narrowed anteriad over apical one-third of length (Fig. 20.227a); spermatheca capsule pear-shaped, rounded apically and gradually narrowed posteriorly, apical invagination small and shallow, stem thin and irregularly twisted posteriorly (Fig. 20.227f–j); only females are known in North America.......Mocyta fungi (Gravenhorst) – Body slender (Fig. 20.226a); elytra as long as pronotum or slightly shorter (Fig. 20.226a); pronotum evenly rounded on both sides of disc (Fig. 20.226a); spermatheca with capsule pitchershaped, flat apically with elongate apical invagination, stem broadly coiled posteriorly (Fig. 20.226h); median lobe of aedeagus as illustrated (Fig. 20.226b, c).................................. ..............................Mocyta breviuscula (Mäklin)

Amblystegii Species et al. 2015b, 2018b)

Group (Klimaszewski

Diagnosis. Body entirely dark brown to black except for light-coloured appendages; pronotum almost twice as wide as long, approximately as broad as elytra or slightly broader, sides arcuate, pubescence on disc directed posteriad along midline and obliquely posteriad elsewhere; elytra in most specimens longer than pronotum, except for M. breviuscula; spermatheca and median lobe of aedeagus as illustrated (Fig. 20.226b, c, h). 190. Mocyta 20.226a–h)

breviuscula

(Mäklin) (Fig.

432

Homalota breviuscula Mäklin, 1852. Bernhauer 1907 [as Atheta fungi breviuscula]; Casey 1910 [as seven synonyms in Acrotona], Lohse and Smetana 1985 [as Atheta (Mocyta) breviuscula], Gusarov 2003a, Klimaszewski et al. 2007b, 2008a, b, 2011, 2015b, 2016d, 2018b, Majka and Klimaszewski 2008b, Webster et al. 2009, Brunke et al. 2012. Diagnosis. Body narrowly oval, length 2.4–3.0 mm; colour uniformly dark brown to almost black and often with reddish tinge, appendages yellowish to reddish-brown (Fig. 20.226a); antennomeres I–IV elongate and V–X subquadrate; pronotum transverse, arcuate laterally and arcuate basally; elytra transverse and nearly as long as pronotum; abdomen broadly arcuate laterally. MALE. Tergite VIII broadly arcuate apically (Fig. 20.226d); sternite VIII parabolic apically with broad distance between base of the disc and antecostal suture, the suture more or less sinuate (Fig. 20.226e); median lobe of aedeagus in lateral view with ventral margin of tubus straight, apex narrowly rounded (Fig. 20.226b); in dorsal view bulbus narrowly oval and tubus narrowly triangular apically, apex rounded (Fig. 20.226c). FEMALE. Tergite and sternite VIII broadly arcuate apically (Fig. 20.226f, g); spermatheca with capsule pitcher-shaped and flat apically, apical invagination deep, stem broadly coiled posteriorly (Fig. 20.226h). The combination of uniform body colour, elytra no longer than pronotum, distinct shape of spermatheca with deep capsular invagination, and shape of male sternite VIII with broad space between base of disc and antecostal suture, can distinguish M. breviuscula from the remaining Nearctic congeners. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB, NF, NS, ON, QC, SK, YT. USA: AK, CA, NV, OR. New locality data: British Columbia, Kamloops, 8.X.2008, aspen litter beside high pond, A.Davies (CNC) 2 males, 1 female. BCol., W of Terrace, Shames Mtn., road to ski

20

Tribe Athetini Casey, 1910

area, 54 28.140 N, 128 56.050 W, in moss and alder litter at cold stream, A. Davies (CNC) 1 sex undetermined; BCol., Hwy 16, rest stop at Cluculz Creek, 53 54.520 N, 123 35.220 W, 776 m, 31.VII.2012, sedges at edge of creek, A. Davies (CNC) 1 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: in NF, frequently captured in pitfall traps in various forest types such as birch, spruce-lichen, spruce-poplar, fir; also in vegetation on coastal dunes, in disturbed fields among grass and weeds, and on shrubby limestone barrens (Klimaszewski et al. 2018b). In QC, from pitfall traps in yellow birch/balsam fir forests, and sifted from litter in an oak savanna/ alvar in ON (Klimaszewski et al. 2018b). One individual from sphagnum and litter in an eastern white cedar swamp and another captured in Lindgren funnel trap in an old red pine forest in NB (Klimaszewski et al. 2018b). In BC, specimens were found in aspen litter beside high pond, in moss and alder litter at cold stream, and in sedges at edge of creek. Collecting period: IV–X. Collecting method: pitfall traps, Lindgren funnel trap, sifting forest litter. 191. Mocyta { (Fig. 20.227a–j)

fungi

(Gravenhorst)

Aleochara fungi Gravenhorst, 1806. Casey 1893, 1910 [as four synonyms], Bernhauer 1907 [in Atheta (Acrotona)], Clark 1949 [in Acrotona], Muona 1984 [in Atheta (Xenota)], Gusarov 2003a [synonymy], Smetana 2004 [synonymy], Klimaszewski et al. 2007a, b, 2008a, b, 2011, 2015b, 2018b, Majka and Klimaszewski 2008a, b, McLean et al. 2009. Diagnosis. Body oval, length 2.4–3.0 mm; colour uniformly dark brown to black, in some specimens body black and posterior or central part of elytra with reddish tinge, appendages light brown (Fig. 20.227a); antennomeres I–IV elongate and V–X subquadrate or slightly transverse; pronotum broad, transverse, rounded laterally and arcuate basally; elytra transverse, variable in length, and ca. as long as pronotum

20

Tribe Athetini Casey, 1910

or longer; abdomen broadly arcuate laterally. MALE. Median lobe of aedeagus as illustrated (Fig. 20.227b, c) [males absent in North America]. FEMALE. Tergite VIII broadly arcuate apically (Fig. 20.227d); sternite VIII truncate apically with fringe of microsetae, distance between antecostal suture and base of disc narrow, antecostal suture sinuate (Fig. 20.227e); spermatheca variable in shape, with capsule pear-shaped, and stem variably twisted posteriorly (Fig. 20.227f–j). This species is externally very similar to M. amblystegii and may be identified with certainty only by the shape of the spermatheca. The presence of males in Canadian populations of M. amblystegii and lack of males in Canadian populations of M. fungi may also aid in the identification of these species. DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: AB, BC, LB, NB, NF, NS, NU, ON, PE, QC, SK, YT. USA: AK, CA, ME, MA, MN, NH, NY, OR, RI, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: found in various forest types and wetlands, usually in litter. In NB, in silver maple swamps in leaf litter, in leaf litter in eastern white cedar swamps and forests, under alders in leaf litter and moss in a marsh, treading vegetation in boggy marsh and a cattail marsh, river margin on moist clay soil among tall grasses, a sea beach in decaying sea wrack, in pitfall traps in a red spruce forest. In NF, from pitfall traps in cut and burned balsam fir, birch, spruce-poplar and riparian forests, in agricultural fields and among vegetation on coastal sand dunes. Elsewhere, in mixed forests and yellow birch forests. Collecting period: V–IX. Collecting method: pitfall traps, sifting forest litter, treading marsh vegetation. Discreta Species Group (Klimaszewski et al. 2015b, 2018b) Diagnosis. Pronotum large, transversely orbicular with rounded lateral edges and hind angles,

433

usually much broader than elytra, pubescence on disc directed posteriad along midline and obliquely posteriad elsewhere; elytra short, as long as pronotum or shorter; median lobe of aedeagus and spermatheca as illustrated (Fig. 20.228b, g). 192. Mocyta discreta (Casey) (Fig. 20.228a–g) Eurypronota discreta Casey, Klimaszewski et al. 2015b, 2018b.

1893.

Diagnosis. Body broadly oval, length 2.4–2.8 mm; head and entire abdomen or its basal part brown to almost black, pronotum and basal half of abdomen in most specimens light brown, testaceous or reddish-brown, elytra yellowish to reddish-brown with some irregular small dark brown spots and darker than pronotum, legs and palps yellowish to reddishbrown and antennae either uniformly yellowish to light brown (Fig. 20.228a); antennomeres I–IV elongate and V–X variable in length from subquadrate to slightly elongate; pronotum transverse, usually very large but variable in width, from slightly to 1/7 broader than elytra [pronotum usually broader in females than in males], highly rounded laterally, and arcuate basally; elytra transverse and shorter than pronotum; abdomen broadly arcuate laterally and with very strong macrosetae apically. MALE. Tergite VIII truncate apically with very numerous long macrosetae (Fig. 20.228c); sternite VIII parabolic apically and with numerous long macrosetae in apical part of disc, space between base of disc and antecostal suture broad, antecostal suture sinuate medially (Fig. 20.228d); median lobe of aedeagus with narrowly oval bulbus and moderately long tubus, ventral margin straight, subapical part narrowly elongate, apex rounded in lateral view (Fig. 20.228b). FEMALE. Tergite and sternite VIII arcuate apically (Fig. 20.228e, f); spermatheca pear-shaped with small and shallow apical invagination, stem thin and straight anteriorly and coiled posteriorly (Fig. 20.228g). This species differs from other Mocyta species in having the body bicoloured, the pronotum

434

large and wider than the elytra, very strong macrosetae on the apical part of the abdomen, and antennomeres V–X subquadrate to slightly elongate. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], ON, QC, SK. USA: IA, MN, WA [new state record]. We report this species from BC and WA for the first time here based on the following: British Columbia, Inkaneep, Osoyoos IR1, 49.09 N, 119.32 W, 6.V.1994/31.V.1994, pitfall, Purshia association, coll. G.G.E. Scudder (UBC) 1female; near Mud Lk., Osoyoos IR1, 49.13 N, 119.31 W, 2.VI.1994/7.VII.1994, pitfall, Purshia association, coll. G.G.E. Scudder (UBC) 1 male; Osoyoos IR1, 49.09 N, 119.32 W, 6.V.1994/31. V.1994, pitfall, Purshia association, coll. G.G.E. Scudder (UBC) 1 male; Vasseux Cr., CWS bench, 49.16 N, 119.30 W, 10.V.1994/3. VI.1994, 3.VI.1994/8.VII.1994, 3.VI.1994/6. VII.1994, pitfall trap, Purshia association, coll. G.G.E. Scudder (UBC, LFC) 1 male, 4 females; Vasseux Cr., 49.16 N, 119.80 W, 10.V.1994/3. VI.1994, 3.VI.1994/8.VII.1994, 3.VI.1994/6. VII.1994, pitfall trap, Purshia association, coll. G.G.E. Scudder (UBC) 1 female. USA: Washington, Oroville E., Osoyoos Lk., 48.58 N, 119.25 W, 5.VII.1994/2.VIII.1994, pitfall, Purshia association, coll. G.G.E. Scudder (UBC) 1 male. COLLECTION AND HABITAT DATA. Habitat: associated with hardwood/deciduous forests; adults in forest litter, deciduous leaf mold, in maple/oak forest litter and other unspecified deciduous forest litter. In BC specimens were found in association with Purshia sp. Collecting period: IV–X. Collecting method: pitfall traps, sifting forest litter.

Genus Nehemitropia Lohse, 1971 (Fig. 20.229a–g) (Klimaszewski et al. 2007a, 2011, 2018b)

20

Tribe Athetini Casey, 1910

Diagnosis. Pronotum and elytra of about same length, pronotum with lateral margins strongly converging apicad, abdomen tapering with strong lateral macrosetae, superficially similar to Oxypoda species; head strongly narrowed basally, eyes moderately large, about as long as genae in dorsal view; antennomeres I–III distinctly elongate, IV–X subquadrate, XI as long as two preceding antennomeres combined; pronotum slightly transverse, pubescence on disc directed apicad along midline and obliquely posteriad elsewhere; elytra subparallel, pubescence on disc directed posteriad and slightly obliquely posteriad; abdomen strongly tapering apicad in apical half; median lobe of a simple form, venter of tubus sinuate in lateral view; tergite VIII without lateral teeth, slightly emarginate medially at apex; apical tergites and sternites with strong macrosetae like in Oxypoda species; spermatheca with tubular capsule lacking apical invagination, stem wide and arcuately bent posteriorly. 193. Nehemitropia lividipennis (Mannerheim) { (Fig. 20.229a–g) Oxypoda lividipennis Mannerheim, 1830. Hamilton 1889, Muona 1984 [as Acrotona sordida], Moore and Legner 1975 [as Atheta (Coprothassa) sordida], Klimaszewski et al. 2007a, 2011, 2016d, 2018b, Michaud et al. 2010. Diagnosis. Body narrowly oval, length 3.0–3.8 mm; colour dark brown, with elytra (except for scutellar region), legs, and antennae light brown to yellowish-brown (Fig. 20.229a); integument moderately glossy, forebody with fine, sparse punctation, short pubescence and meshed microsculpture; head slightly elongate, narrower than pronotum; antennae slender and long, antennomere IV subquadrate, V–VII subquadrate or slightly elongate, VIII–X slightly transverse; pronotum moderately transverse, widest basally and strongly narrowing apically; elytra as broad as maximum width of pronotum, at suture about as long as pronotum, with lateral posterior emarginations; abdomen tapering

20

Tribe Athetini Casey, 1910

apically. MALE. Tergite VIII truncate apically and slightly emarginate medially, macrosetae stout (Fig. 20.229c); sternite VIII rounded apically, macrosetae stout (Fig. 20.229d); median lobe of aedeagus with venter of tubus sinuate in lateral view (Fig. 20.229b). FEMALE. Tergite VIII truncate apically and slightly emarginated medially, macrosetae stout (Fig. 20.229e); sternite VIII with apical V-shaped emargination, macrosetae stout (Fig. 20.229f); spermatheca S-shaped, capsule tubular without apical invagination, stem broad and sinuate (Fig. 20.229g). Superficially, N. lividipennis is similar to some Oxypoda species (e.g., O. convergens), and Atheta longicornis, and it is often misidentified as such in various collections. DISTRIBUTION (Klimaszewski et al. 2018b). Origin: Palearctic, adventive in North America, considered cosmopolitan by Smetana (2004). CANADA: BC [new provincial record], NB, NF, NS, ON, PE, QC, SK. USA: CA, LA, MA, MI, MN, NE, NJ, NM, NY, PA, TX, VT. We report this species from BC for the first time here based on the following: British Columbia, Queen Charlotte Is., Q.C. City, 16.X.1983, sifting compost, J.M. Campbell (CNC) 1 sex undetermined. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: open fields and pastures, open marsh, maple/beech forest, the edge of an oak forest, lake margin, and jack pine forest. Adults in organic debris, including dead grass, in caribou, horse and cow dung, compost (common in decaying corncobs and cornhusks), rotting Boletus mushroom, and in nest of Microtus pennsylvanicus (Ord). One individual from cobblestones on lakeshore and some flying adults collected with butterfly net in evening (15:00–18:00 h). Collecting period: IV–IX. Collecting method: sifting compost, organic debris, decaying mushrooms, pitfall traps, collecting flying adults with butterfly net during evening.

435

Genus Ousipalia Gozis, 1886 (Fig. 20.230a–g) (Casey 1910; Moore and Legner 1975 (as Evanystes Gistel); Lohse and Smetana 1988; Ashe 2001) Diagnosis. Body length 2.0–2.1 mm, subparallel, brown to yellow with head and abdomen darker, integument glossy; head subquadrate, eyes small, genae about two diameter of eye in dorsal view; mandibles symmetrical and broad, right mandible with a small medial tooth, ventral molar area without patches of denticle; ligula divided into two widely separate lobes; antennomere II about as long as III, IV subquadrate, V–X transverse, XI as long as two preceding antennomeres combined; pronotum not distinctly margined laterally, microsetae directed posteriad along midline of disc, pubescence short, pronotal hypomera fully visible in lateral view; elytra broadening posteriorly, shorter than pronotum along midline; mesocoxae contiguous; process of mesoventrite long and wide, extended to half length of mesocoxal cavities; hind tarsus with basal tarsomere shorter than second one, tarsi each with one empodial seta; capsule of spermatheca narrowly pitcher-shaped, apical invagination moderately large, stem sinuate and narrowly looped posteriad but not swollen like in Geostiba. Comments. Ousipalia is similar to Geostiba in having the ligula divided into two separate lobes, but in Geostiba these lobes are contiguous while in Ousipalia they are widely separated (Gusarov 2002b). 194. Ousipalia pacifica Casey (Fig. 20.230a–g) Ousipalia pacifica Casey, 1910. Lectotype (female): USA: Cal. [ifornia]; pacifica Casey; Type USNM 39026; Casey bequest 1925 (USNM). Present designation. Diagnosis. Body subparallel, length 2.0–2.1 mm; head and most of abdomen dark brown, and pronotum, elytra and appendages

436

yellowish brown (Fig. 20.230a); surface of forebody with fine isodiametric microsculpture, punctation very fine, sparse, punctures separated by 3–4 times their diameter; head subquadrate; eyes about half length of genae in dorsal view; antennomere II about as long as III, and IV–X moderately transverse, XI as long as IX and X combined; pronotum slightly transverse, slightly wider than head; elytra at suture shorter or about as long as pronotum along midline of disc, wider than long (1.2), with fine isodiametric microsculpture and fine, somewhat asperate punctation, punctures separated by 2–3 times their diameter; abdomen broad, arcuate laterally, abdominal terga with fine, scarcely visible, microsculpture of transverse meshes, with fine and sparse punctation, punctation becoming finer towards abdomen apex, MALE. Tergite VIII with apical margin arcuate, entire (Fig. 20.230c); sternite VIII rounded apically (Fig. 20.230d); median lobe of aedeagus in lateral view with tubus arched ventrad near apical fourth of ventral margin, apical part moderately broad, apex rounded (Fig. 20.230b). FEMALE. Tergite VIII with apical margin arcuate (Fig. 20.230e); sternite VIII broadly rounded apically (Fig. 20.230f); spermatheca with club-shaped capsule, stem narrowly looped and not swollen posteriorly (Fig. 20.230g). DISTRIBUTION. Origin: Nearctic. CANADA: BC, SK [new provincial record]. USA: CA, WA [new state record]. New locality data for BC, new provincial record for SK, and new state records for WA in USA: Canada: British Columbia, Vancouver Is., BCol., Duncan, Mt. Tzouhalem, 48 470 N, 123 370 W, 19.X.2008, second-growth Douglas fir and maple litter with mushrooms, A. Davies (CNC) 1 female; 4 sex undetermined; Vancouver Is., NE Victoria, Mt. Douglas, 48 290 3200 N, 123 200 2600 W, 14.X.2011, red cedar and maple litter at base of rock face, A. Davies (CNC) 1 sex undetermined; Vancouver Is., BCol., French Beach Provincial Park, 48 23’N, 123 560 W, 6. VII.2010, in beach wrack, A. Davies (CNC)

20

Tribe Athetini Casey, 1910

1 female; Vancouver Is., BCol., 4 km SW Bamberton, Spectacle Lake, 48 340 4100 N, 123 340 1900 W, 7.X.2011, moss on Douglass fir logs in second-growth forest, A. Davies (CNC) 2 females; Victoria, Fort Rodd Hill, 48.4344-123.455, 9 m, 23.V.2015, Berlese, moss, coll. Darren Copley (RBCM) 1 female; BC, Victoria, Oak Bay, 48.4207-123.3033, 3 m, 25.V.2015, compost, soil sifting, coll. B. Godin and D. Horwood (RBCM) 1 female; BC, Lilloet, 50.7286-121.6181, 835 m, 23, 24.V.2014, moss, Berlese, coll. Darren and Claudia Copley (RBCM) 3 females; BC, Mechosin, Devonian Regional PK., 48.3645-123.537, 20 m, Berlese, moss, coll. Darren and Claudia Copley (RBCM) 1 female; BC, 25 km W Creston, 18.VIII.1982, leg. R. Baranowski, BWRS (ZMLU) 2 females; Vaseux Cr., 49.15 N, 119.31 W, 2.V.1995/8. VI.1995, Purshia association, pitfall traps, coll. G.G.E. Scudder (UBC, LFC) 9 females; same data except: Kennedy flats, 1.V.1995/8.VI.1995, 2.V.1995/8.VI.1995, 9.V.1984/3.VI.1983 (UBC, LFC) 9 females; Osoyoos, I.R.I., Brights Winery, 49.13 N, 119.32 W, 31.V.1994/5.VII.1994, 6. V.1994/2.VI.1994, 2.VI.1994/7.VII.1994, 7, VII.1994/4.VIII.1994, Purshia association, pitfall traps, coll. G.G.E. Scudder (UBC, LFC) 7 females, 1 sex undetermined; Oliver, I.R.I., 49.10 N, 119.31 W, 6.VII.1995/11.VIII.1995, Purshia association, pitfall traps, coll. G.G.E. Scudder (UBC, LFC) 1 male; Saskatchewan, Prince Albert, 53.9789-106.2859, 534 m, 13.IX.2014, Mushrooms, sifting, coll. B. Godin and D. Horwood (BGC, RBCM) 2 males, 2 females. USA, Washington, Oroville E, Osoyoos Lake, 48.58 N, 119.25 W, 2.VIII.1994/6.IX.1994, Purshia association, pitfall trap, coll. G.G.E. Scudder (LFC) 2 females. COLLECTION AND HABITAT DATA. Habitat: moss, mushrooms, compost; secondgrowth Douglas fir and maple litter with mushrooms; moss on Douglas fir logs in second-growth forest, red cedar and maple litter at base of rock face; also occurs in in beach wrack, and dry habitats with Purshia association. Collecting period: V–VIII, X. Collecting

20

Tribe Athetini Casey, 1910

method: sifting forest litter, moss and mushrooms, processing moss through Berlese funnels, pitfall traps.

Genus Paraleptonia Klimaszewski, 2002 (Fig. 20.231a–g) (Klimaszewski and Winchester 2002) Diagnosis. Paraleptonia can be distinguished from other athetine genera by the combination of the following characters: subparallel-sided body; average body length 3.0 mm; head with two fully marked infraorbital carinae, postocular region shorter than the maximum diameter of eye; pronotum transverse with microsetae at midline of disc directed mostly anteriad, and approximately horizontally laterad elsewhere, macrosetae present and distinct laterally, hypomera broad and visible for two thirds of the pronotal length in lateral view; elytron virtually lacking emargination at the posterolateral angle; mesosternum not carinate, mesosternal process slender and acutely pointed apically, reaching two thirds length of the mesocoxal cavities; metasternal process extremely short and wide, virtually non-existent; mesotibia with 3 long macrosetae (longer than tibial width); metatarsal basal segment only slightly longer than the second segment; mandibles broadly triangular basally and slender sickle-shaped apically and without distinct lateral dents; ligula reduced and not apparent; labial palpi with three palpomeres; maxillary palpi with four palpomeres, apical one needle-shaped and about two thirds length of the penultimate palpomere; maxilla with galea about as long as lacinia, without larger setae, internal edge of lacinia entire and without spines; male tergite VIII truncate apically and without teeth; median lobe of aedeagus narrowly elongate, bulbus only slightly broader than tubus in dorsal view; tubus strongly narrowed in apical third and projecting ventrally, athetine bridge scarcely visible at the base of tubus, internal sac sclerites small and

437

inconspicuous, flagellum not apparent; apical lobe of paramere short and slightly broadened apically with large subapical macrosetae and small one on the apex; male tergite eight simple and without sexually dimorphic structures. 195. Paraleptonia (Fig. 20.231a–g)

pacei

Klimaszewski

Paraleptonia pacei Klimaszewski, Klimaszewski and Winchester, 2002.

in

Diagnosis. Body length 3.2–3.5 mm, robust, moderately convex, narrowly oval, moderately glossy on forebody and strongly so on abdomen, light yellowish brown with rust tinge, head, median part of abdomen and antennae, except for three basal segments, dark brown (Fig. 20.231a); forebody sparsely punctate and pubescent, pubescence fine, punctation slightly asperate on elytra, microsculpture meshy, sculpticells irregularly hexagonal and slightly convex; head round, as long as wide, distinctly narrower than pronotum, depressed medially, postocular region shorter than the maximum diameter of eye, tempora carinate, carinae complete, strong basally and weak apically (scarcely visible); maxillary palpus with four palpomeres, last one needle-shaped and about two thirds of the length of the penultimate; labial palpus with three palpomeres; antenna slender and long, filiform, as long as the pronotum and elytra combined, antennomere IV slightly elongate, V–X slightly transverse and broadening apically, terminal elongate and as long as one and a half of the two preceding antennomeres combined, basal 4 antennomeres glossy and remaining matte; pronotum impressed mesally, strongly transverse, broadly rounded laterally and basally and straight apically, hypomeron visible for two thirds of the pronotal length in lateral view, pubescence directed anteriad along most of the midline, and apicolaterad along arcuate lines on sides of the disc; elytra moderately convex, slightly transverse, broader than pronotum, posterior margins

438

broadly arcuate and sinuate laterally, with shallow lateral emarginations, pubescence directed straight and slightly obliquely posteriad; abdomen subparallel, strongly glossy, three first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII transverse, truncate apically, bearing four large setae on each side (Fig. 20.231f); sternite VIII elongate, rounded apically, bearing 9–10 long and stout setae on each side posteriorly (Fig. 20.231g); median lobe of aedeagus in dorsal view with narrowly oval bulbus of median lobe of aedeagus and very long subparallel tubus in basal two-thirds, apical part narrowly triangular (Fig. 20.231d, e), in lateral view tubus very long and narrow, thin apically and pointed ventrad, apex pointed (Fig. 20.231b, c). FEMALE. Unknown. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. New locality data: British Columbia, Vancouver Island, Mt. Cain, 50.15 N, 126.25 W, 23.VIII-14.IX.1996, 17.5 PAN 1, coll. N. Winchester (LFC) 1 male; Mt. Cain, 50.15 N, 126.25 W, 1.IX-19.IX.1997, 17.4 PIT 10, coll. N. Winchester (LFC) 1 male. Mt. Cain, 50.15 N, 126.25 W, 23.VIII-14.IX.1996, 17.5 PAN 1, coll. N. Winchester (LFC) 1 male; COLLECTION AND HABITAT DATA. Habitat: transition zone of Sitka spruce forest in Carmanah Valley, Vancouver Is. Collecting period: VIII–IX. Collecting method: Malaise traps, pitfall trap, Pan trap.

Genus Pelioptera Kraatz, 1857 (¼Tropimenelytron Pace, 1983) (Fig. 20.232a–h) (Gusarov 2002c; Klimaszewski et al. 2018b) Diagnosis. Body parallel-sided, glength 2.5–3.7 mm; most of body uniformly brownishyellow to reddish-brown with head and abdominal segment VI dark brown, and appendages brown; head slightly longer than wide, eyes

20

Tribe Athetini Casey, 1910

small, genae 1.5–3.0 times as long as eyes in dorsal view, infraorbital carinae short, not reaching posterior margin of eye; antennomere II longer than III, IV–X transverse; ligula with wide base and split apically; pronotum subquadrate, margined laterally, microsetae directed posteriad along the midline, macrosetae moderately long, hypomera fully visible in lateral view; mesocoxae separated; mesolateral macroseta of middle tibia as long as tibial width, hind tarsus with basal article about as long as second, each tarsus with one empodial seta, tarsal formula 4-5-5; median lobe of aedeagus with tubus straight or slightly bent ventrally at apex in lateral view; spermatheca without apical invagination. Comments (Klimaszewski et al. 2018b). In reporting Tropimenelytron from North America for the first time, Gusarov (2002c: 3) treated Tropimenelytron as separate from Pelioptera at the generic level. Pace (2006: 349) explained that the ligula of Tropimenelytron is identical to that of Pelioptera, and the separation of the mesocoxae, another character used to separate the two genera, is very variable. This leaves only the shape of the distal bulb of the stem of the spermatheca which is asymmetrical in Tropimenelytron, as a diagnostic character to justify placing this taxon at the genus level. Pace (2006) pointed out that this character is sufficient only to separate taxa at the subgeneric level in athetines (e.g., as for the subgenera of Atheta), and he therefore considered Tropimenelytron as a subgenus of Pelioptera. We follow his classification. 196. Pelioptera 20.232a–h)

robusta

(Gusarov) (Fig.

Tropimenelytron robustum Gusarov, 2002c. Diagnosis. Body length 3.4–3.7 mm; reddish brown or brown, elytra and legs and parts of abdomen slightly paler (Fig. 20.232a); head surface with strong isodiametric microsculpture, making hard to see fine and slightly asperate

20

Tribe Athetini Casey, 1910

punctation, distance between punctures equal to 2–3 times their diameter; temples 2.5–2.7 times as long as eyes; antenna with second antennal article 1.1 times as long as third, article IV slightly transverse (ratio 1.6), V–X strongly transverse (ratio 1.8–2.0), last article longer than two preceding articles combined; pronotum subquadrate, length to width ratio 0.97, microsculpture and punctation as on head; elytra wider than pronotum (0.67–0.73 mm) and at suture slightly shorter than length of pronotum along midline, surface glossy, with isodiametric microsculpture, meshes larger than on pronotum, punctation fine and asperate, distance between punctures equal to their diameter; abdominal terga glossy, with fine and weak isodiametric microsculpture, with fine asperate punctation, on terga 3–5 distance between punctures equals 2–4 times their diameter, on terga 6–7 punctation finer, distance between punctures equals 3–6 times their diameter. MALE. Tergite VIII truncate apically and slightly crenulate (Fig. 20.232d); sternite VIII rounded apically (Fig. 20.232e); median lobe of aedeagus with oval bulbus and triangular tubus in dorsal/ventral view (Fig. 20.232c); in lateral view tubus narrow and about straight ventrally, apex narrowly rounded (Fig. 20.232b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 20.232f), sternum VIII truncate apically (Fig. 20.232g); spermatheca with spherical capsule and stem sinuate and narrowly twisted posteriorly (Fig. 20.232h). According to Gusarov (2002c), Pelioptera robusta differs from T. californicum in having a longer body and a broader apex of the median lobe of the aedeagus in ventral view (Figs. 50, 52 and 60, 62 in Gusarov 2002c). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: OR. New locality data: British Columbia, Vancouver Is., Mt. Cain, 50 13N, 126 22W, 9. VI-23.VI.1997, 16.2 PIT1, N. Winchester (LFC) 1 sex undetermined; same data except: 23.V-9. VI.1997, 16.2 PIT 4, coll. N. Winchester (LFC) 1 female; 23.VI-7.VII.1997 (LFC) 1 female.

439

COLLECTION AND HABITAT DATA. Habitat: mixed conifer forest, litter. Collecting period: VI–VII. Collecting method: pitfall traps.

Genus Philhygra Mulsant and Rey, 1873a (Figs. 20.233–20.239) (Lohse et al. 1990; Klimaszewski et al. 2011) Diagnosis. Body elongate, subparallel; colour dark brown to black; head orbicular or subquadrate, more-or-less narrower than pronotum; antennae long and slender, antennomeres V–X elongate to slightly transverse; infraorbital carina partial or complete; pronotum at most one quarter wider than long, broadest at middle of its length, sides arcuate, hind angles rounded, pubescence on disc directed anteriad on apical two-thirds of midline and posteriad on rest of midline, laterad elsewhere; elytra at suture about as long as pronotum or longer; abdomen subparallel; apex of process of mesoventrite rounded; coxae nearly contiguous; male tergite VIII truncate apically and without secondary sexual characters; male sternite VIII very elongate, often projecting posteriorly under pygidium; median lobe of aedeagus with tubus enlarged and of complex forms, bulbus large, internal sac often with dentate structures; spermatheca vestigial in most species and difficult to find; female pygidium with different membranous structures. Comments. This genus is rich in species in Canada and many species remain undescribed, including in British Columbia. Species are affiliated with riparian and wetland habitats and exceptionally with fungi. Key to Species of Philhygra 1. Antennomeres VII–X clearly elongate (Fig. 20.234a); body broad (Fig. 20.234a); median lobe of aedeagus with tubus highly sinuate subapically in lateral view (Fig. 20.234b);

440

female pygidium as illustrated (Fig. 20.234h) ..... ........................Philhygra botanicarum (Muona) – Antennomeres VII–X subquadrate or transverse; other characters not as above...................2 2. Antennomeres VII–X subquadrate to slightly transverse............................................................3 – Antennomeres VII–X clearly transverse .......5 3. Body narrow and uniformly dark brown (Fig. 20.238a); tubus of median lobe of aedeagus broadly elongate, apex broadly rounded in lateral view (Fig. 20.238b); female pygidium with heavily sclerotized narrow structure (Fig. 20.238g)....................................................... .. Philhygra terrestris Klimaszewski and Godin – Body broad, dark brown with paler, red or yellow-brown elytra; other characters not as above ..................................................................4 4. Body narrowly subparallel (Fig. 20.237a); tubus of median lobe of aedeagus narrowly elongate, apex triangular in shape and pointed in lateral view (Fig. 20.237b); female pygidium without heavily sclerotized structures (Fig. 20.237h, i) ................................................... ............................ Philhygra laevicollis (Mäklin) – Body broadly subparallel (Fig. 20.235a); tubus of median lobe of aedeagus broadly elongate, apex triangular in shape and hooked in lateral view (Fig. 20.235b); female pygidium without heavily sclerotized structures (Fig. 20.235h) ....... ........ Philhygra charlottae Klimaszewski, sp. n. 5. Body small, length 1.8–2.2 mm, narrowly subparallel (Fig. 20.236a); tubus of median lobe of aedeagus with serrate ventral margin in lateral view (Fig. 20.236b); female terminalia as illustrated (Fig. 20.236e–g).................................. ................................. Philhygra clemens (Casey) – Body large, length 2.6–3.2 mm; narrowly oval; other characters not as above ....................6 6. Head, pronotum and abdomen dark brown to black and elytra paler reddish brown (Fig. 20.233a); tubus of median lobe of aedeagus hooked apically and pointed ventrad in lateral view (Fig. 20.233b); female terminalia as

20

Tribe Athetini Casey, 1910

illustrated (Fig. 20.233e–g).................................. ................ Philhygra angusticauda (Bernhauer) – Head dark brown almost black, and pronotum, elytra and abdomen paler reddish brown (Fig. 20.239a); tubus of median lobe of aedeagus broadly subparallel apically and truncate apically in lateral view (Fig. 20.239b); female terminalia as illustrated (Fig. 20.239e–g) ............................. .......... Philhygra terrivaga Klimaszewski, sp. n.

197. Philhygra angusticauda (Bernhauer) (Fig. 20.233a–g) Atheta (Metaxya) angusticauda Bernhauer, 1909. Hatch 1957 [cited as Atheta (Brundinia)], Seevers 1978, Klimaszewski and Winchester 2002, Webster et al. 2009, Klimaszewski et al. 2018b. Diagnosis. Body moderately flattened, narrowly subparallel, length 3.0–3.2 mm; colour uniformly dark brown to nearly black, elytra with rust tinge, legs usually slightly paler and reddish brown, antenna uniformly dark brown or reddish brown (Fig. 20.233a); integument moderately glossy, forebody moderately densely punctate and pubescent, pubescence long, punctation fine and slightly asperate, entire forebody with distinct microsculpture, sculpticells consist of round and slightly convex meshes; head slightly elongate, round, approximately as wide as pronotum, eyes about as long as genae in dorsal view, strong basally, diffused posteriorly; antennae slender and long, about as long as the pronotum and elytra combined, basal 3 antennomeres distinctly elongate, IV–V moderately elongate, VI–VII quadrate, VIII–X slightly transverse, terminal one elongate and as long as two preceding antennomeres combined; pronotum impressed mesally, slightly transverse, emarginate laterally and posteriorly, approximately rectangular in shape, broadest at apical third, hypomeron visible for nearly entire length of pronotum in lateral view, pubescence on disc directed anteriad and anterolaterad along midline, laterad basally, and obliquely laterad along arcuate lines elsewhere; elytra moderately convex, slightly elongate, and

20

Tribe Athetini Casey, 1910

insignificantly broader than pronotum, posterior margins broadly arcuate and with shallow lateral emarginations, pubescence on disc directed slightly obliquely posterolaterad to nearly straight laterad at base; abdomen subparallel, slightly glossy, three first visible tergites deeply impressed at base; hind tarsus with basal article as long as following one. MALE. Tergite VIII transverse, broadly truncate apically (Fig. 20.233c); sternite VIII highly elongate, parabolic apically, distance between basal margin and antecostal suture wide (Fig. 20.233d); median lobe of aedeagus with tubus narrowly elongate, apex hooked and sharply pointed ventrad in lateral view (Fig. 20.233b), internal sac sclerites as illustrated in lateral view (Fig. 20.233b), in dorsal view median lobe elongate, with broadly oval bulbus and narrow tubus. FEMALE. Tergite VIII truncate apically (Fig. 20.233e); sternite VIII rounded apically and slightly produced medially, distance between apical margin and antecostal suture wide (Fig. 20.233f); pygidium as illustrated (Fig. 20.233g); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short and narrow stem. DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB. USA: AK, NH. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: in NB, specimens collected from gravel on gravel bars along shaded brooks in a northern hardwood forest (common), a boreal forest (white spruce, black spruce, balsam fir), a mature mixed forest, and in an alder swamp in debris on muddy soil near a brook flowing into a river; in BC, from pitfall traps in clearcut and transition zone of Sitka spruce forest. Collecting period: VI–VIII. Collecting method: splashing gravel and turning gravel on gravel bars along brooks, sifting litter on brook margin, pitfall traps. 198. Philhygra botanicarum (Muona)  (Fig. 20.234a–h)

441

Atheta (Philhygra) botanicarum Muona, 1983. Muona 1984 [as Atheta (Philhygra)], Klimaszewski et al. 2008a, 2011, 2015a, 2018b, Majka and Klimaszewski 2008a, b, Webster et al. 2009. Diagnosis. Body narrowly oval, broadest at elytra, length 3.8–4.5 mm; colour dark brown to piceous, with bases of antennae, legs, elytra except for scutellar region, and tip of abdomen rusty-brown or brownish yellow (Fig. 20.234a); integument moderately glossy, forebody with fine, dense punctation and meshed microsculpture; head round, slightly narrower than pronotum; eyes longer than genae in dorsal view; antennae slender with all antennomeres elongate (more elongate in females); pronotum slightly transverse and almost as wide as elytra; elytra at suture slightly longer than pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically and shallowly, widely emarginate (Fig. 20.234d); sternite VIII highly elongate and truncate apically (Fig. 20.234e); median lobe of aedeagus with tubus broadening apically and with hooked apex in lateral view (Fig. 20.234b); in dorsal view bulbus broadly oval, tubus broad, triangular apically (Fig. 20.234c); internal sac structures complex as illustrated (Fig. 20.234b, c). FEMALE. Tergite VIII truncate apically and shallowly emarginated medially (Fig. 20.234f); sternite VIII parabolic apically, apex produced and rounded (Fig. 20.234g); pygidium as illustrated (Fig. 20.234h); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short and narrow stem. DISTRIBUTION. Origin: Holarctic. CANADA: BC, LB, NB, NF, NS, ON, SK, YT. USA: not recorded. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018a). Habitat: typically found in moist leaves (often on muddy soils) near margins of vernal ponds or small brooks in alder swamps, mixed forests, northern hardwood forests, a coniferous forest, a spruce forest, and a wet willow stand; also in drift material on a

442

20

lakeshore, in moist leaves under a sap flow of yellow birch, and collected by treading vegetation in a seasonally flooded marsh on pond margin. Collecting period: V–VIII. Collecting method: sifting moist leaf litter and drift material, treading marsh vegetation, flight intercept traps. 199. Philhygra charlottae sp. n. (Fig. 20.235a–h)

Klimaszewski,

Holotype (male): Canada, British Columbia, Queen Charlotte Is., Yakoun R., 26 km NW Queen Charlotte City, 21.VIII.1983, 83-101, sifting under mushrooms and moss, coll. J.M. Campbell (CNC). Paratypes: Queen Charlotte Is., 11.1 km NW Queen Charlotte City, 15. VII.1983, 83-9, treading sandy river bank, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., 14 km E Masset, 13.VII.1983, 83-5, treading Carex marsh and sifting Alnus litter, coll. J.M. Campbell (CNC) 2 males, 2 females; Queen Charlotte Is., 1.4 km S Masset, 13. VII.1983, 83-3, sifting Alnus litter, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., 8.8 km SW Tow Hill, 22.VIII.1983, 83-108, treading Sphagnum and Carex at edge of marsh, coll. J.M. Campbell (CBC) 3 females; Queen Charlotte Is., Lyall Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, coll. J.M. Campbell (CNC) 1 female; Vancouver Is., Goldstream Pk., 27.V.1968, A. Smetana (CNC) 1 male. Etymology. Derived from the name of Queen Charlotte Is., where the type series was captured. Diagnosis. Body broad, subparallel, broadest at elytra, length 4.0–4.4 mm; colour dark brown to piceous, with bases of antennae, legs, elytra except for scutellar region, and tip of abdomen red-brown (Fig. 20.235a); integument moderately glossy, forebody with fine, dense punctation and meshed microsculpture; head round, slightly narrower than pronotum; eyes longer than genae in dorsal view; antennae slender with all antennomeres elongate, except IX–X subquadrate; pronotum slightly transverse,

Tribe Athetini Casey, 1910

distinctly narrower than elytra at base; elytra at suture slightly longer than pronotum; abdomen subparallel. MALE. Tergite VIII truncate apically (Fig. 20.235d); sternite VIII highly elongate and truncate apically (Fig. 20.235e); median lobe of aedeagus with tubus triangular apically and with sinuate apex in lateral view (Fig. 20.235b); in dorsal view bulbus broadly oval, tubus broad, triangular apically (Fig. 20.235c); internal sac structures complex as illustrated (Fig. 20.235b, c). FEMALE. Tergite VIII truncate apically and shallowly emarginated medially (Fig. 20.235f); sternite VIII parabolic apically, apex triangularly produced and rounded (Fig. 20.235g); pygidium as illustrated (Fig. 20.235h); spermatheca not found. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: specimens collected from mushrooms and moss, sandy river bank, Sphagnum and Carex marsh and Alnus litter, and mushrooms and marten dung on thick moss. Collecting period: V, VII, VIII. Collecting method: sifting mushrooms and moss, treading sandy river bank and Sphagnum and Carex at edge of marsh, and sifting mushrooms and marten dung on thick moss. Comments. This species is similar to P. botanicarum but may be distinguished by subquadrate antennomeres VII–X and by the genital characters. 200. Philhygra clemens (Casey) (Fig. 20.236a– g) Noverota clemens Casey, 1910. Gusarov 2002a [as Philhygra], Klimaszewski et al. 2005, 2007b, 2015a, 2018b, Majka and Klimaszewski, 2008b, Brunke et al. 2012. Diagnosis. Body narrowly subparallel, length 1.8–2.2 mm; colour dark brown with elytra, legs and tip of abdomen yellowish brown, or head and

20

Tribe Athetini Casey, 1910

abdomen except for apex dark brown, pronotum light reddish brown, and elytra, basal antennomeres, legs and palps yellowish (Fig. 20.236a); integument moderately glossy, forebody with fine, dense punctation and meshed microsculpture; head round, about as wide as pronotum; eyes about as long as genae in dorsal view; antennae slender, antennomere IV slightly elongate, V–X moderately transverse; pronotum slightly transverse and as wide as elytra, broadest in apical third; elytra at suture about as long as pronotum; abdomen subparallel. MALE. Tergite VIII broadly arcuate apically (Fig. 20.236c); sternite VIII elongate, rounded apically (Fig. 20.236d); median lobe of aedeagus with tubus highly narrowly elongate and pointed apically, its ventral margin bearing small denticles medially in lateral view (Fig. 20.236b); in dorsal view bulbus large and tubus narrowly elongate; internal sac structures as illustrated (Fig. 20.236b). FEMALE. Tergite VIII rounded apically (Fig. 20.236e); sternite VIII produced apically and slightly pointed medially (Fig. 20.236f); pygidium as illustrated (Fig. 20.236g); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short, looped, narrow stem. DISTRIBUTION. Origin: Nearctic. CANADA: BC, MB, NB, NS, ON, QC, YT. USA: OH, WI. New locality data: British Columbia, Queen Charlotte Is., 11.1 km NW Queen Charlotte City, 15.VII.1983, 83-9, sandy river bank, coll. J.M. Campbell (CNC) 1 male, 1 female. COLLECTION AND HABITAT DATA. Habitat: in NB, usually in riparian habitats or near lakeshores where it can be common; among gravel on gravel bars along shaded brooks in northern hardwood forest and a balsam popular forest near river, in moss near splash zone of waterfalls, in moss on rocks in middle of river, river margins in drift material, silver maple swamp near lake in moist leaves on vernal pond margins, in drift material (maple seeds and other kinds of plant debris) on lake margins; also

443

collected from pitfall traps in a red spruce regenerating forest and flight intercept traps in maple beech forest (Klimaszewski et al. 2018b). In BC, two adults were captured from sandy river bank Collecting period: IV–IX. Collecting method: sifting litter and moss, splashing moss and gravel, river bank, collected with butterfly net during late afternoon, flight intercept traps, pitfall traps. 201. Philhygra 20.237a–i)

laevicollis

(Mäklin) (Fig.

Homalota laevicollis Mäklin, 1852. Bernhauer 1907, Casey 1911 [as Atheta mordax and Metaxya prognata], Hatch 1957, Lohse and Smetana, 1985, Klimaszewski and Winchester 2002, Klimaszewski et al. 2005, 2018b, Majka and Klimaszewski 2008b, Brunke et al. 2012. Diagnosis. Body narrowly subparallel, moderately flattened, length 3.0–4.2 mm; colour dark brown, elytra except for scutellar area, legs, and basal antennomeres rust-brown (Fig. 20.237a); integument moderately glossy, forebody sparsely punctate and pubescent, with pubescence long, punctation fine, microsculpture distinct and consisting of round and slightly convex meshes; head slightly elongate, round, about as wide as pronotum, eyes as long as genae in dorsal view; antennae slender, at least as long as pronotum and elytra combined, antennomeres I–VII elongate, VIII–X subquadrate and terminal one about as long as two preceding antennomeres combined; pronotum slightly transverse, impressed mesally, emarginate laterally and basally, pubescence on disc reduced to scattered hairs; hypomeron visible almost for entire length of pronotum; elytra slightly transverse, broader than pronotum; abdomen subparallel. MALE. Tergite VIII arcuate apically (Fig. 20.237d); sternite VIII highly elongate, rounded apically and with wide distance between antecostal suture and base of disc (Fig. 20.237e); median lobe of aedeagus with moderate-sized bulbus, tubus short, triangular at apex in lateral view (Fig. 20.237b); in dorsal view bulbus roughly oval, tubus short, triangular

444

apically in dorsal view (Fig. 20.237c); internal sac sclerites as illustrated (Fig. 20.237b). FEMALE. Tergite VIII truncate apically and slightly emarginated medially (Fig. 20.237f); sternite VIII rounded apically, distance between antecostal suture and base of disc wide (Fig. 20.237g); pygidium as illustrated (Fig. 20.237h, i); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination and with short narrow stem. DISTRIBUTION. Origin: Nearctic. CANADA: BC, NB?, NS?, ON?. USA: AK, WA. New locality data: British Columbia, Queen Charlotte Is., Lyall Is., Gate Cr., 10.VIII.1983, 83-70, sifting mushrooms and marten dung on thick moss, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., Skowkona Cr., 7.9 km NW Queen Charlotte City, 29.VII-4.VIII.1983, 83-55, ex rotting pile of mushrooms, coll. J.M. Campbell (CNC) 1 male, 1 female; Queen Charlotte Is., 11.1 km NW Queen Charlotte City, 15. VII.1983, 83-9, treading sandy river bank, coll. J.M. Campbell (CNC) 1 male; Queen Charlotte Is., 7.9 km NW Queen Charlotte City, 16-19. VII.1983, 83-19, dung and carrion trap, coll. J.M. Campbell (CNC) 1 female; Queen Charlotte Is., Tanu Is., Tanu, 8.VIII.1983, sifting wet moss on weeping wall and at seepage area, coll. J.M. Campbell (CNC) 1 male; Kitsumkalum Lake at Hwy 13, S of Rosswood, 54 470 3400 , 128 450 9700 W, 146 m, 3/2.VII.2012, in moss at creek, A. Davies (CNC) 1 male, 3 females; BCol., Hwy 16 rest stop at Cluculz Creek, 53 540 5200 N, 123 350 2200 W, 776 m, 31.VII.2012, in Salix and Alnus litter with moss, 10 m from Lake, A. Davies (CNC) 1 male; BCol., Driftwood Canyon Prov. Pk., 54 490 6500 N, 127 010 4100 W, 752 m, 27. VII.2012, alder, poplar litter and moss along creek, A. Davies (CNC) 1 male, 2 females. COLLECTION AND HABITAT DATA. Habitat: in BC, adults were collected in clearcut and transition zone of Sitka spruce forest (Vancouver Island, Carmanah Valley); in

20

Tribe Athetini Casey, 1910

mushrooms and marten dung on thick moss, sandy river bank, wet moss along creek and on weeping wall and at seepage area and in alder and poplar litter. Collecting period: VII, VIII. Collecting method: sifting leaf litter (alder, poplar), wet moss along creek and moss on weeping wall and at seepage area; turning gravel along streams, treading sandy river bank, dung and carrion trap, pitfall traps. Comments (Klimaszewski et al. 2018b). Specimens of P. laevicollis from eastern Canada agree in most external and genital characters with those from BC and AK, but there are slight differences in the length of antennomeres VII–X and the internal sac structures of the median lobe of the aedeagus. Additional studies are needed to decide whether these differences represent geographical variation within one species or whether the two populations are closely related cryptic species. 202. Philhygra terrestris Klimaszewski and Godin (Fig. 20.238a–g) Philhygra terrestris Klimaszewski and Godin, in Klimaszewski et al. 2012. Klimaszewski et al. 2015a, 2018a, Webster et al. 2016. Diagnosis. Body narrowly subparallel, length 2.9–3.0 mm; head and abdomen dark brown to black, pronotum and particularly elytra slightly paler, basal antennomeres and legs yellowish (Fig. 20.238a); integument highly glossy, forebody with fine, dense punctation and meshed microsculpture; head round, slightly narrower than pronotum; eyes as long as genae in dorsal view; antennae slender, antennomeres IV–VI elongate, VII–X subquadrate to slightly transverse; pronotum slightly transverse and almost as wide as elytra; elytra at suture longer than pronotum; abdomen long, subparallel. MALE. Tergite VIII arcuate apically (Fig. 20.238c); sternite VIII elongate and parabolic apically (Fig. 20.238d); median lobe of aedeagus with tubus broadly elongate, constricted in basal half,

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Tribe Athetini Casey, 1910

subapical part broadly compressed oval in lateral view (Fig. 20.238b). FEMALE. Tergite VIII rounded apically (Fig. 20.238e); sternite VIII rounded apically and slightly produced medially at apex (Fig. 20.238f); pygidium as illustrated (Fig. 20.238g); spermatheca not illustrated, minute with short sac-shaped capsule without apical invagination, stem short, narrow. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], NB, SK, YT. USA: AK [new state record]. We report this species from BC and AK for the first time here based on the following: Canada, BC, Queen Charlotte Is., 4.7 km N Rennell Sound Rd., Ghost Cr., 19.VII.1983, 83-24, 700 ft., ex gravel along edge of stream, J.M. Campbell (CNC) 1 male; Valemont, Yellowjacket ck., 3-8. VI.2014, pitfall, Kinbasket # 1046, 52.702119.05, 758 m, debris, coll. Charlene Wood (RBCM) 1 male; 1 female; Vancouver Is., Goldstream Pk., 27.V.1968, B), A. Smetana (CNC) 3 males, 2 sex undetrmined; BCol., Toboggan Creek at Johnson Rd., N of Toboggan Lk., 54 530 3200 N, 127 150 7800 W, 480 m, 27. VII.2012, in sedges along creek, A. Davies (CNC) 1male; BCol., Kitsumkalum Lk. at Hwy 13, S of Rosswood, 54 470 3400 N, 128 550 9700 W, 146 m, 31.VII.2012, in moss at creek, A. Davies (CNC) 1 female; BCol., Smithers perimeter trail wetland S of Hwy 16 540 2200 N, 127 090 1300 W, 490 m, 31.VII.2012, moss in swamp, A. Davies (CNC) 1 female. USA: AK, Taylor Hwy, mi 82, 8.VII.1968, A. Smetana (CNC) 1 female. COLLECTION AND HABITAT DATA. Habitat: riparian; in NB, in gravel/clay on margin of partially shaded brooks in old-growth mixed forests and near lake, in drift material (mostly small sticks and conifer buds) on river margin, and in clay/sand under alders on river margin; YT specimen found in litter in a mixed forest, BC specimen found in gravel along edge of stream, in sedges along a creek, and in moss at creek and in swamp. Collecting period: V–VII.

445

Collecting method: splashing clay and fine sand on sand bars along shaded brooks, and splashing drift material along a river margin, sifting litter, pitfall traps. 203. Philhygra terrivaga Klimaszewski, sp. n. (Fig. 20.239a–g) Philhygra terrivaga Gusarov, Klimaszewski and Winchester, 2002.

in

Holotype (male): USA, Oregon, Hood River Co., 1.5 mi E Cascade Locks, Herman Creek, 10.VII.1978, L. and N. Herman (AMNH), present designation. Paratypes: Canada, British Columbia, Upper Carmanah Valley, UTM 10U CK 803005, 21.VI-3.VII.1991, CC.MT2/3, N. Winchester (LFC) 2 females; Upper Carmanah Valley, UTM 10U CK 803005, 4.VII-15. VII.1991, CC.PT4, N. Winchester (LFC) 1 female; Upper Carmanah Valley, UTM 10U CJ 803006, 12.VIII-27.VIII.1991, TZ.MT2, N. Winchester (LFC) 1 male, present designation. Diagnosis. Body moderately flattened, moderately glossy, narrowly subparallel, length 2.6–3.2 mm, light brown, head and median part of abdomen dark brown, legs usually yellowish, antenna uniformly light brown with basal antennomeres slightly paler (Fig. 20.239a); forebody moderately densely punctate and pubescent, pubescence long, punctation fine and slightly asperate, microsculpture well visible on entire forebody, sculpticells consist of round and slightly convex meshes; head slightly elongate, round, approximately as wide as pronotum, postocular region approximately as long as the maximum diameter of eye, tempora carinate, carinae almost complete, strong basally and weak apically; antenna slender, shorter than the pronotum and elytra combined, antennomeres IV–V slightly elongate or quadrate, VI–X moderately transverse, terminal one elongate and as long as two preceding antennomeres combined, basal IV–V glossy and remaining matte; pronotum impressed mesally, slightly transverse, emarginate laterally and posteriorly,

446

approximately rectangular in shape, broadest at apical third, hypomeron visible for nearly entire length of pronotum in lateral view, pubescence directed anteriad and antero-laterad along midline of disc, obliquely laterad along arcuate lines on sides and horizontally at base; elytra moderately convex, at suture as long as pronotum along midline, posterior margin broadly arcuate and with shallow lateral emarginations, pubescence directed slightly obliquely postero-laterad to nearly horizontally-laterad at base; abdomen subparallel, slightly glossy, three first visible tergites strongly impressed at base; basal metatarsomere as long as the following one. MALE. Tergite VIII transverse, broadly arcuate apically and slightly produced medially (Fig. 20.239c); sternite VIII strongly elongate, rounded apically, and bearing wide distance between basal margin and antecostal suture (Fig. 20.239d); median lobe of aedeagus elongate, with broadly oval bulbus in dorsal view; in lateral view tubus extremely narrowly elongate, subapical part angulate dorsally, apex truncate (Fig. 20.239b), internal sac sclerites as illustrated (Fig. 20.239b). FEMALE. Tergite VIII transverse and broadly arcuate apically (Fig. 20.239e); sternite VIII slightly produced, rounded apically, distance between apical margin and antecostal suture wide (Fig. 20.239f); pygidium as illustrated (Fig. 20.239g); spermatheca not found. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: OR, WA. COLLECTION AND HABITAT DATA. Habitat: transition zone of Sitka spruce forest in Carmanah Valley, Vancouver Is. Collecting period: VI–VIII. Collecting method: Malaise trap, pitfall trap. Comments. Gusarov never formally published Ph. terrivaga as anticipated (Klimaszewski and Winchester (2002). Klimaszewski and Winchester (2002: 44) cited this species as Philhygra terrivaga “Gusarov (in press)”, and they provided a description and designated a holotype; however the name was never published by Gusarov, and it

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was never cited explicitly as a “new species” (Article 16.1), and so the name was not made available until now.

Genus Pontomalota Casey, 1885 (Fig. 20.240a–i) (LeConte 1863; Casey 1910; Ahn and Ashe 1992) Diagnosis. Body length 2.6–5.5 mm, robust, heavily sclerotized, narrow, flattened, parallel sided, colour variable from light brown to dark brown or black, elytra pale appearing often creamy white, microsculpture strong, mostly hexagonal in shape; integument of forebody granulose due to a fine-meshed reticulation; pubescence sparse and uniform; head transverse, slightly deflexed, tempora subparallel, neck absent; eyes as long as postocular region of head; infraorbital carina partial; antenna slightly incrassate; labrum transverse, slightly sinuate anteriorly; mandibles asymmetrical, apex blunt and turned downward, incisor tooth absent, and molar tooth present on right mandible but absent on left one, molar region with rows of teeth, protheca well developed; lacinia and galea elongate; maxillary palpus with four palpomeres, terminal one narrow distally; labial palpus with three palpomeres, elongate and styliform; ligula wide, short and bifid forming two rounded lobes apically; mentum trapezoidal; pronotum 1.2–1.3 times as wide as long, narrowest at base and widest near middle; hypomeron visible in lateral view; mesoventrite not carinate medially; mesocoxal cavities narrowly separated; isthmus very short; tarsal formula 4-5-5; elytra short, shorter than pronotum, outer posterior angles deeply sinuate, hind wings rudimentary, present as small membranous appendages; abdomen broad, uniformly converging to broadly rounded apex, tergites III–VI moderately impressed at base. Median lobe of aedeagus simple with short tubus. Species of this genus inhabit the mid to upper littoral zone of fine grain sandy beaches, and are often found in association with Thinopinus and

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Tribe Athetini Casey, 1910

447

Thinusa (Ahn and Ashe 1992). They feed on intertidal invertebrates (Ahn and Ashe 1992). 204. Pontomalota 20.240a–i)

opaca

(LeConte) (Fig.

Phytosus opaca LeConte 1863. Casey 1885, Fenyes 1920, Gouix and Klimaszewski 2007 [synonymies], Ahn and Ashe 1992. Diagnosis. Body length 2.6–4.9 mm, narrowly subparallel, with strong microsculpture; colour variable, head light brown to black, pronotum light brown to dark brown, elytra light brown or creamy white, appendages paler that the body (Fig. 20.240a); head slightly deflexed, about 1.1–1.4 times as wide as long; antenna beadshaped; pronotum about 1.2–1.3 times as wide as long, longer than head and elytra; elytra 0.8–0.9 times as long as pronotum; abdomen subparallel with evenly wide paratergites. MALE. Tergite VIII truncate apically (Fig. 20.240d); sternite VIII rounded laterally and truncate medially (Fig. 20.240e); tubus of median lobe of aedeagus long, arcuate ventrally and slightly sinuate medially, apex narrowly triangular and slightly pointed ventrad in lateral view (Fig. 20.240b), in dorsal view bulbus rounded and tubus subparallel basally and triangular apically (Fig. 20.240c). FEMALE. Tergite VIII truncate apically (Fig. 20.240f); sternite VIII arcuate apically (Fig. 20.240g); spermatheca slender, S-shaped, capsule tubular with broad apical invagination, stem long and sinuate (Fig. 20.240h, i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, OR, WA. New locality data: British Columbia, Vancouver Is., 15 km SE of Tofino, Long Beach, 49 040 0300 N, 125 480 0200 W, 16.X.2011, under wood near high tide line on sea beach, A. Davies (CNC) 2 males, 2 females, 7 sex undetermined. COLLECTION AND HABITAT DATA. Habitat: mid to upper littoral zone of fine grain sandy beaches, under wood near high tide line on

sea beach. Collecting period: I, II, V–X. Collecting method: sifting debris in littoral zone of fine grain sandy beaches, collecting from under wood on a beach.

Genus Psammostiba Yosii and Sawada, 1976 (Figs. 20.241a–i and 20.242a–h) (Casey 1910; Yosii and Savada 1976; Gusarov 2003d) Diagnosis (adopted from Gusarov 2003d, see his Figs. 58, 64, 65, 67 and 68). Body length 3.0–5.0 mm, body parallel-sided; anterior margin of labrum concave; antennomeres V–X slightly elongate or subquadrate; ligula long and split in apical half; labial palpus with setae a, b, g and d present; pronotum slightly transverse, 1.1 times as wide as long, with microsetae directed anteriorly along the midline; in lateral portions of the disc microsetae directed laterally; pronotal macrosetae short; pronotal hypomera fully visible in lateral view; medial macroseta of mesotibia inconspicuous and thin, as long as tibial width; tarsal formula 4-5-5; metatarsal segment 1 as long as segment 2; no empodial setae; tarsal claws of different length (best observed in dorsal view), external claw longer than internal; male tergite and sternite VIII with numerous prominent and erected macrosetae; male tergite VIII with serrate margin; flagellum of internal sac of median lobe of aedeagus without pointed apex; proximal portion of spermatheca with one coil. Psammostiba can be distinguished from Atheta by the strong isodiametric microsculpture of the entire body, by the concave anterior margin of the labrum, by the basal impression on tergum VI, by lacking the empodial seta, and by the tarsal claws of different length. Key to Nearctic Species of Psammostiba 1. Body length 4.0–4.6 mm, pronotum subquadrate (Fig. 20.241a); elytra at suture about as long as pronotum along midline, maximum width of elytra one-fifth wider than maximum width of pronotum (Fig. 20.241a); posterior

448

margin of male tergite VIII with small crenulation (Fig. 20.241d); tubus of median lobe of aedeagus moderately produced ventrad in lateral view, apical part short (Fig. 20.241b); spermatheca with large loop of posterior coil (Fig. 20.241h)........... ..................Psammostiba comparabilis (Mäklin) – Body length 3.0–3.1 mm, pronotum transverse (length/width 0.5–0.6 mm) (Fig. 20.242a); elytra at suture longer than pronotum along midline, maximum width of elytra one-third wider than maximum width of pronotum (Fig. 20.242a); posterior margin of male tergite VIII with distinct crenulation (Fig. 20.242d); tubus of median lobe of aedeagus strongly produced ventrad in lateral view, apical part long (Fig. 20.242b); spermatheca with small loop of posterior coil (Fig. 20.242h)...... Psammostiba kenaii Gusarov

205. Psammostiba comparabilis (Mäklin) (Fig. 20.241a–h) Homalota comparabilis Mäklin, 1853. Fenyes, 1920, Seevers 1978, Lohse and Smetana 1985, Gusarov 2003d. Diagnosis (based on description by Gusarov 2003d). Body length 4.0–4.6 mm, colour black with brown legs and light brown tarsi (Fig. 20.241a); head in both sexes with weak medial impression, surface dull, with strong and dense isodiametric microsculpture, punctation fine, distance between punctures equals their diameter; head subquadrate, about as wide as maximum width of pronotum, temple length to eye length ratio 0.9–1.1; antennomeres IV–X elongate, antennomere XI shorter than two last ones combined; pronotum slightly transverse, 1.2 times as wide as head, width to length ratio 1.2, surface dull, with strong and dense isodiametric microsculpture, punctation finer than on head, poorly visible on dull background, distance between punctures equal to ½–1 times their diameter; elytra much wider (1.10–1.30 mm) and longer (0.99–1.17 mm; measured from humeral angle) than pronotum (elytral length to pronotal length ratio 1.5), 1.1 times as wide as long;

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punctation as on pronotum; abdominal terga dull, with strong and dense isodiametric microsculpture, with fine punctation, punctation on terga VI–VII almost as dense as on terga III– V, distance between punctures equals 1–3 times their diameter. Apical margin of tergum 7 with white palisade fringe. MALE. Tergite VIII with finely crenulate posterior margin and with prominent apical macrosetae (Fig. 20.241d); sternite VIII parabolic, rounded at apex and with prominent apical macrosetae (Fig. 20.241e); tubus of median lobe of aedeagus narrow apically and pointing ventrally in lateral view, apical part of tubus short and apex rounded (Fig. 20.241b); in dorsal bulbus oval and tubus short, broad and rounded apically (Fig. 20.241c). FEMALE. Tergite VIII truncate apically with prominent macrosetae in apico-lateral corners of disc (Fig. 20.241f); sternite VIII parabolic, rounded at apex and with prominent apical macrosetae (Fig. 20.241g); spermatheca S-shaped, narrow, capsule tubular with moderate apical invagination, stem sinuate and with large loop of posterior coil (Fig. 20.241h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA. New locality data: British Columbia, Queen Charlotte Is., Tow Hill Park, 13.VII.1983, 83-4, sifting kelp on beach, J.M. Campbell (CNC) 1 male, 3 females, 22 sex undetermined; Charlotte Is., Louis Is., Skedans, 6.VIII.1983, sifting beach wrack and on beach at night, J.M. Campbell (CNC) 2 males, 1 female. COLLECTION AND HABITAT DATA (Gusarov 2003d). Habitat: littoral habitats, found in decomposing seaweed on sandy seashores; some specimens were taken on beach at night. In BC, collected from rocks in intertidal zone, and from kelp and wrack. Collecting period: VII, VIII, X. Collecting method: sifting decomposing seaweed.

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449

Comments. This species was previously recorded only from Victoria, BC (Gusarov 2003d). 206. Psammostiba 20.242a–h)

kenaii

Gusarov (Fig.

Psammostiba kenaii Gusarov 2003d. Holotype (male): Alaska, Haines, 3.VII.1968, Campbell and Smetana; 1310, Holotypus, male, Psammostiba kenaii, V.I. Gusarov 2003; Holotype CNC No. 23420 (CNC). Paratypes: labelled as the holotype (CNC) 1 male, 2 females [one female dissected]. Examined. Diagnosis (based on description by Gusarov 2003d). Body length 3.0–3.1 mm, colour black, elytra dark brown to black, legs brown, tarsi yellowish brown (Fig. 20.242a); head in both sexes with weak medial impression, surface dull, with strong and dense isodiametric microsculpture, punctation fine, distance between punctures equals their diameter; head subquadrate, about as wide as maximum width of pronotum, temple length to eye length ratio 0.9–1.0; antennomeres IV–VII elongate, VIII–X elongate or subquadrate, antennomere XI shorter than two preceding ones combined; pronotum slightly transverse, 1.2 times as wide as head, width to length ratio 1.2, surface dull, with strong and dense isodiametric microsculpture, punctation finer than on head, poorly visible on dull background, distance between punctures equal to ½–1 times their diameter; elytra much wider (0.7–0.8 mm) and longer (0.7 mm; measured from humeral angle) than pronotum (elytral length to pronotal length ratio 1.5), 1.1 times as wide as long; punctation as on pronotum; abdominal terga dull, with strong and dense isodiametric microsculpture, with fine punctation, punctation on terga VI–VII almost as dense as on terga III– V, distance between punctures equals 1–3 times their diameter. Apical margin of tergum VII with white palisade fringe. MALE. Tergite VIII with distinctly crenulate posterior margin and with apical macrosetae prominent (Fig. 20.242d); sternite VIII parabolic, produced apically, apex

rounded; apical macrosetae prominent (Fig. 20.242e); tubus of median lobe of aedeagus evenly narrow apically and pointing ventrally in lateral view, apical part of tubus long (Fig. 20.242b); in dorsal view median lobe with oval bulbus and tubus short, broad and rounded apically (Fig. 20.242c). FEMALE. Tergite VIII broadly arcuate apically with prominent macrosetae in apico-lateral corners of disc (Fig. 20.242f); sternite VIII parabolic, rounded apically, apical macrosetae prominent (Fig. 20.242g); spermatheca S-shaped, narrow, capsule tubular with moderate apical invagination, stem sinuate and with small posterior loop (Fig. 20.242h). DISTRIBUTION. Origin: Nearctic. CANADA: BC [Victoria]. USA: AK, CA. New locality data: British Columbia, Queen Charlotte Is., Tow Hill Park, 13.VII.1983, 83-4, sifting kelp on beach, J.M. Campbell (CNC) 2 males, 7 females; Charlotte Is., Skidegate, 83-17, 17.VII.1983, ex seaweed, J.M. Campbell (CNC) 2 males, 1 female; Vancouver Is., French Beach Provincial Park, 48 230 N, 123 560 W, 6. VII.2010, in beach wrack, A. Davies (CNC) 1 sex undetermined. COLLECTION AND HABITAT DATA (Gusarov 2003d). Habitat: littoral habitats, found in decomposing seaweed on sandy seashores. Collecting period: VII, X. Collecting method: sifting decomposing seaweed.

Genus Schistoglossa Kraatz, 1856 (Figs. 20.243–20.246) (Klimaszewski et al. 2009b, 2018b) Diagnosis. Body subparallel, length 1.8–3.5 mm; colour brown; head slightly narrower than pronotum or nearly as broad; eyes small, about one quarter to one third length of genae in dorsal view, infraorbital carina short, feeble or virtually obsolete; antennae

450

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Tribe Athetini Casey, 1910

(Fig. 20.246h)....................................................... .......Schistoglossa hampshirensis Klimaszewski – Elytra at suture at most as long as pronotum along midline; antennomeres VII–X subquadrate or elongate; other characters not as above ......... 2

approximately reaching posterior margin of elytra, antennomeres VII–X slightly transverse or slightly elongate, ligula completely divided into two separate lobes, mandibles appearing bifid apically but in fact left mandible is bifid and the right mandible bears two apical and one subapical teeth, apical teeth often overlap and appear as one in dorsal view creating an illusion of a bifid mandible, the teeth of left mandible are in the horizontal plane and the two apical teeth of the right mandible are in the vertical plane and can be locked together; maxillary palpi with four palpomeres, terminal article needle-shaped; pronotum transverse to nearly subquadrate, broadest in middle or in apical one-third, hind angles rounded, hypomera clearly visible in lateral view, integument smooth and moderately glossy, microsculpture weak, punctation fine, pubescence on disc directed entirely or partially posteriad along midline, lateroposteriad elsewhere; mesocoxae contiguous; male tergite VIII with two large lateral teeth and apical crenulation in Schistoglossa s. str., tergite VIII truncate apically and without lateral teeth and apical crenulation in the subgenus Boreomorpha; median lobe of aedeagus bearing large bulbus and triangular tubus in dorsal view, crista apicalis small, internal sac with reversed V-shaped structure in dorsal view in Schistoglossa s. str., with semicircular apical structures in the subgenus Boreomorpha; spermatheca with capsule tubular (Schistoglossa s. str.), or approximately spherical (Boreomorpha).

3. Maximum width of pronotum distinctly less than width of elytra at shoulders (Fig. 20.244a); elytral ratio of maximum length to maximum width 0.7 (Fig. 20.244a); apical margin of tergite VIII with two larger lateral teeth and lacking regular crenulation (Fig. 20.244d); spermatheca approximately L-shaped (Fig. 20.244h)............... ..............Schistoglossa charlottae Klimaszewski – Maximum width of pronotum about the same as width of elytra at shoulders (Fig. 20.245a); elytral ratio of maximum length to maximum width 0.8 (Fig. 20.245a); apical margin of male tergite VIII with two larger lateral teeth and regular crenulation (Fig. 20.245d); spermatheca approximately S-shaped (Fig. 20.245h)............... ............. Schistoglossa campbellii Klimaszewski

Comments. These are cold-loving species occurring in arctic and subarctic habitats, and in the temperate regions they are usually confined to bogs, fens, and marshes.

(Subgenus Boreomorpha Klimaszewski and Webster, in Klimaszewski et al. 2009b) (Fig. 20.243a–h) (Klimaszewski et al. 2009b, 2018b)

Key to Species of Schistoglossa 1. Elytra at suture slightly shorter than pronotum along midline (Fig. 20.246a); male tergite VIII emarginate apically and finely crenulate (Fig. 20.246d); antennomeres VII–X distinctly transverse (Fig. 20.246a); apex of median lobe of aedeagus slightly produced ventrally (Fig. 20.246b); spermatheca S-shaped

2. Antennomers VII–X elongate (Fig. 20.243a); eyes strongly protruding (Fig. 20.243a); body length 2.8–3.0 mm, elytra broadening posteriorly (Fig. 20.243a); median lobe of aedeagus and spermatheca as illustrated (Fig. 20.243b, c, h) ........... .............. Schistoglossa carexiana Klimaszewski – Antennomeres VII–X subquadrate; eyes not protruding, streamlined with contour of head; genital structures not as above ........................... 3

Diagnosis. Antennomeres V–X slightly elongate; eyes moderate-sized, about half as long as genae in dorsal view; infraorbital carina weak, present only basally or obsolete; male tergite VIII broadly arcuate apically, apical margin entire and without lateral teeth and crenulation; median lobe of aedeagus with bulbus and tubus evenly coalescent in dorsal view, internal sac without V-shaped structure typical for Schistoglossa

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Tribe Athetini Casey, 1910

s. str. but with two crescent-shaped subapical structures clearly visible in lateral view; spermatheca club-shaped and with approximately spherical capsule. This group contains species that bear some transitional characters between the genera Schistoglossa and Boreophilia but are placed in the former on the grounds of mandibular structures (2 + 3 teeth), consistent with the other Schistoglossa species and the generic type species S. viduata (Erichson). 207. Schistoglossa (Boreomorpha) carexiana Klimaszewski (Fig. 20.243a–h) Schistoglossa carexiana Klimaszewski, in Klimaszewski et al. 2009b, 2015a. Diagnosis. Body length 2.8–3.0 mm, width 0.4–0.5 mm, approximately uniformly dark brown with slightly paler tip of abdomen, legs, antennae, and mouthparts (Fig. 20.243a); integument with weak microsculpture, sculpticells irregularly hexagonal, pubescence short, except slightly longer on head and abdomen, yellowishbrown in artificial light, moderately dense; head slightly elongate, rounded posteriorly, carinate basally, tempora approximately as long as twice the maximum diameter of eye as seen from above; antennae slim, all antennomeres elongate; pronotum slightly transverse, broadly rounded laterally and posteriorly, broadest in apical half, pubescence directed anteriorly on apical third of the midline and laterally forming arched lines elsewhere; elytra strongly transverse, and broadening posteriorly, hind margin nearly straight, pubescence directed obliquely posteriad; abdomen parallel-sided, three basal tergites deeply impressed basally. MALE. Tergite VIII arcuate apically, apical margin without or with small emargination (Fig. 20.243d); sternite VIII parabolic apically, antecostal suture sinuate (Fig. 20.243e); in dorsal view bulbus of median lobe of aedeagus broad and flattened, narrowed apically, tubus short, triangular (Fig. 20.243c), in lateral view venter of tubus sinuate, apex projected ventrally (Fig. 20.243b), apical sclerites

451

of internal sac lunar-shaped, and basal sclerites narrowly elongate in lateral view (Fig. 20.243b). FEMALE. Tergite VIII arcuate apically and truncate medially (Fig. 20.243f); sternite VIII slightly parabolic apically (Fig. 20.243g); spermatheca club-shaped with capsule spherical, bearing internal ribbings and without apparent apical invagination, stem moderately narrow and approximately straight (Fig. 20.243h). DISTRIBUTION. Origin: Nearctic. CANADA: BC, SK. USA: not recorded COLLECTION AND HABITAT DATA. Habitat: in British Columbia, adults were captured from Sphagnum and Carex at the edge of a marsh. In Saskatchewan, one female was captured on a sandy beach. Collecting period: VI– VIII. Collecting method: by treading vegetation at the edge of marsh and sifting debris in littoral zone of fine grain sandy beaches. (Subgenus Schistoglossa Kraatz, 1856) (Figs. 20.244a–h and 20.246a–h) (Klimaszewski et al. 2009b, 2018b) Diagnosis. Antennomeres V–X slightly transverse; eyes small, about one-third as long as genae in dorsal view, infraorbital carinae feeble, present only basally or obsolete; male tergite VIII truncate or emarginate apically, apical margin weakly to coarsely crenulate and bearing two large lateral teeth; median lobe of aedeagus with bulbus and tubus distinctly separated in dorsal view (not evenly coalescent), internal sac with V-shaped structure; spermatheca L- or S-shaped and with tubular capsule. 208. Schistoglossa (Schistoglossa) charlottae Klimaszewski (Fig. 20.244a–h) Schistoglossa (Schistoglossa) charlottae Klimaszewski, in Klimaszewski et al. 2009b. Diagnosis. Body length 2.0–2.2 mm, width 0.4 mm, approximately uniformly dark brown with slightly paler tip of abdomen, and legs

452

(Fig. 20.244a); integument with weak microsculpture, sculpticells irregularly hexagonal, pubescence short, except slightly longer on head and abdomen, yellowish-brown in artificial light, moderately dense; head slightly elongate, rounded posteriorly, carinae not apparent, tempora as long as twice the maximal diameter of eye seen from above; antennae slim, antennomeres V–X subquadrate to slightly transverse; pronotum strongly transverse, broadly rounded laterally and posteriorly, broadest in basal third, pubescence directed anteriorly on apical third of the midline and straight posteriorly and obliquely laterally elsewhere; elytra moderately transverse, and subparallel, hind margin slightly sinuate, pubescence directed straight posteriad; abdomen parallel-sided, three basal tergites deeply impressed basally. MALE. Tergite VIII transverse and broadly emarginated apically, apical margin with two large and two minute lateral teeth (Fig. 20.244d); sternite VIII elongate and parabolic apically, antecostal suture sinuate (Fig. 20.244e); median lobe of aedeagus with broadly oval bulbus and narrowly triangular tubus in dorsal view (Fig. 20.244c); venter of tubus arcuate in lateral view, apex produced ventrad (Fig. 20.244b), apical sclerites of internal sac V-shaped in dorsal view (Fig. 20.244c). FEMALE. Tergite VIII truncate apically (Fig. 20.244f); sternite VIII triangularly produced apically and pointed medially (Fig. 20.244g); spermatheca L-shaped with capsule tubular and without apparent apical invagination, stem moderately narrow and approximately straight except for the twisted end (Fig. 20.244h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: in marshes with Carex and Sphagnum. Collecting period: VIII. Collecting method: treading Sphagnum and in Carex marsh. 209. Schistoglossa (Schistoglossa) campbelli Klimaszewski (Fig. 20.245a–h)

20

Tribe Athetini Casey, 1910

Schistoglossa (Schistoglossa) campbelli Klimaszewski, in Klimaszewski et al. 2009b. Diagnosis. Body length 2.0–2.4 mm, width 0.4 mm, approximately uniformly dark brown with slightly paler tip of abdomen and legs (Fig. 20.245a); integument with weak microsculpture, sculpticells irregularly hexagonal, pubescence short, except slightly longer on head and abdomen, yellowish-brown in artificial light, moderately dense; head slightly elongate, rounded posteriorly, weakly carinate basally, tempora as long as two and a half times the maximal diameter of eye as seen from above; antennae slim, antennomeres V–X slightly transverse; pronotum moderately transverse, broadly rounded laterally and posteriorly, broadest in basal half, pubescence directed anteriorly on apical fourth of the midline and posteriorly and obliquely laterally elsewhere; elytra moderately transverse, and slightly broadened posteriorly, hind margin sinuate, pubescence directed straight posteriad; abdomen parallel sided, three basal tergites deeply impressed basally. MALE. Tergite VIII transverse and broadly emarginate apically, apical margin feebly crenulate and with two large lateral teeth (Fig. 20.245d); sternite VIII elongate and parabolic posteriorly, slightly pointing medially, antecostal suture straight (Fig. 20.245e); venter of tubus of median lobe of aedeagus arcuate with apex produced ventrally in lateral view (Fig. 20.245b), in dorsal view bulbus oval in outline and tubus subparallel basally and triangular apically (Fig. 20.245c), apical sclerites of internal sac V-shaped in dorsal view (Fig. 20.245c). FEMALE. Tergite VIII truncate and slightly emarginated medially (Fig. 20.245f); sternite VIII parabolic apically and pointed medially (Fig. 20.245g); spermatheca S-shaped with capsule spherical apically and tubular basally, and with small apical invagination, stem moderately narrow and strongly sinuate (Fig. 20.245h). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: edge of marsh with Carex and

20

Tribe Athetini Casey, 1910

453

Sphagnum. Collecting period: VIII. Collecting method: treading Sphagnum and in Carex marsh.

sinuate and with (Fig. 20.246h).

210. Schistoglossa (Schistoglossa) hampshirensis Klimaszewski (Fig. 20.246a–h)

DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC [new provincial record], QC. USA: NH.

Schistoglossa (Schistoglossa) hampshirensis Klimaszewski, in Klimaszewski et al. 2009b, 2015a, 2018b. Diagnosis. Body length 1.8–2.0 mm, width 0.3 mm; colour virtually uniformly dark brown with tarsi, antennal bases and mouthparts paler (Fig. 20.246a); integument with fine microsculpture, sculpticells irregularly hexagonal, pubescence short, except slightly longer on head and abdomen, yellowish-brown in artificial light, moderately dense; head roughly round, slightly elongate, with feeble infraorbital carina, eyes about one third length of genae in dorsal view; antennae slim, antennomeres V–X moderately transverse; pronotum slightly transverse, broadly rounded laterally and posteriorly, broadest in basal half, pubescence on disc directed anteriad on apical third of midline and obliquely laterad elsewhere; elytra short, at suture about as long as pronotum along midline, distinctly transverse, subparallel, hind margin approximately straight, pubescence on disc directed obliquely or straight posteriad; abdomen parallel-sided, three basal tergites deeply impressed basally. MALE. Tergite VIII broadly emarginated apically and bearing irregular crenulation and two larger lateral teeth (Fig. 20.246d); sternite VIII parabolic apically (Fig. 20.246e); median lobe of aedeagus moderately broad, venter of tubus arcuate, apex slightly produced ventrally in lateral view (Fig. 20.246b), apical sclerites of internal sac V-shaped in dorsal view (Fig. 20.246c), in dorsal view bulbus oval, tubus short, subparallel and triangular apically (Fig. 20.246c). FEMALE. Tergite VIII sinuate apically and bearing small median emargination (Fig. 20.246f); sternite VIII triangularly produced apically and pointed medially (Fig. 20.246g); spermatheca with tubular capsule, stem narrow,

small

posterior

coil

We report this species from BC for the first time here based on the following: British Columbia, 55 km S of Watson Lake, 29.VI.2015, 59.6843-129.1627, 693 m, Firekill, grass, soil sifting, coll. B. Godin and D, Horwood (LFC) 1 female; Bcol., N of Terrace, Kalum Lake Rd., pond at Findley Lake Road, 54 36.350 N, 128 39.110 W, 30.VII.2012, 220 m, edge of pond in sphagnum, A. Davies (CNC) 1 female. COLLECTION AND HABITAT DATA. Habitat: in marshes or near wetlands; in Carex marshes in sphagnum hummocks, in sphagnum hummocks with Carex and grasses in open wet black spruce bog, in Carex hummocks in Carex marshes, in grass and sphagnum at the base of alders in a marsh with grasses and alders, and along stream margins among cobblestones under alders or in litter; one swept from Carex in small Carex marsh. In BC one female was captured in soil in firekilled forest with grass. Collecting period: IV–VI. Collecting method: treading sedge hummocks into water, sifting litter and sphagnum, sweeping foliage in Carex marsh. Comments (Klimaszewski et al. 2018b). The New Hampshire specimens were collected at elevations of 5200–5700 feet and the only Quebec specimen was found on Mount Albert at 3700 feet, suggesting this may be an arctic alpine and or subarctic species. A female from NB was tentatively associated with this species by Klimaszewski et al (2009c), but it had a spermatheca not typical of S. hampshirensis (stem not turned up posteriorly) and was found in a marsh habitat along with S. brunswickensis and S. pseudocampbelli Klimaszewski and Webster. It is possible that it is an aberrant specimen of S. brunswickensis. No additional specimens of

454

20

S. hampshirensis have been found in NB. We therefore remove this species from NB until more specimens can be found.

Genus Seeversiella Ashe, 1986 (Fig. 20.247a–h) (Gusarov 2003c; Klimaszewski et al. 2018b) Diagnosis. Body subparallel, highly glossy, length 2.0–3.5 mm; antennomere II slightly longer than III, IV–X slightly elongate or subquadrate; anterior margin of labrum concave, sensilla a of epipharynx long, ligula with narrow base and subdivided apically, labial palpi with setae a, b and g present (for illustrations see Gusarov 2003d); pronotum slightly transverse, 1.2 times as wide as long, pubescence on midline of disc directed anteriad and posteriad basally and inwards towards midline in central area of disc, hypomera fully visible in lateral view; mesolateral macroseta of middle tibia robust and long, twice as long as tibial width, hind tarsus with tarsomere I as long as II, one long empodial seta present, tarsal formula 4-5-5; process of mesoventrite broad; male tergite III with lateroapical angles prolonged in form of spines; male tergite VII with a weak to moderately strong median carina; median lobe of aedeagus with two sclerotized bands (sclerite-like flat structures) in internal sac, flagellum (copulatory piece) with pointed apex and sclerotized suspensoria (for illustrations see Gusarov 2003d); spermatheca short. 211. Seeversiella (Fig. 20.247a–h)

globicollis

(Bernhauer)

Atheta (Microdota) globicollis Bernhauer, 1907. Ashe 1986 [as synonym Seeversiella bispinosa], Gusarov 2003c, Majka and Klimaszewski 2008b, Klimaszewski et al. 2007b, 2011, 2015a, 2018b, Work et al. 2013, Webster et al. 2016. Diagnosis. Body narrowly elongate, subparallel, length 2.2–2.5 mm; colour reddish-brown, or

Tribe Athetini Casey, 1910

dark brown to black, head usually darker, and antennae, legs, and posterior section of elytra reddish-brown in some (Fig. 20.247a); integument moderately glossy, forebody with fine, moderately dense punctation, silky pubescence and meshed microsculpture; head rounded, about as wide as pronotum, eyes moderately large; antennae moderately robust, antennomeres IV–X elongate; pronotum moderately transverse, widest at middle; elytra slightly wider than head and as wide as pronotum, at suture as long as pronotum; abdomen with sides broadly arcuate. MALE. Tergite VIII arcuate apically (Fig. 20.247d); sternite VIII rounded apically (Fig. 20.247e); median lobe of aedeagus with tubus broadly arcuate, almost straight ventrally, and narrowly rounded at apex in lateral view (Fig. 20.247b); internal sac with two elongate sclerite in bulbus (Fig. 20.247c), in dorsal view bulbus large, suboval with angular anterior angles, tubus short and triangular (Fig. 20.247c). FEMALE. Tergite VIII arcuate apically (Fig. 20.247f); sternite VIII rounded apically (Fig. 20.247g); spermatheca with small tubular and rounded apically capsule, stem clubshaped and swollen posteriorly (Fig. 20.247h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, NF, NS, ON, QC, SK. USA: AZ, CO, ID, MI, MN, MT, NH, NM, SD, WI. MEXICO. GUATEMALA. HONDURAS. COLLECTION AND HABITAT DATA. Habitat: old-growth northern hardwood forest, old-growth white spruce and balsam fir forest, fir and riparian forests, mountain forests, oak forest, yellow birch dominated forest, balsam fir forest, and meadow; specimens from moose dung, soil and moss under logs, leaf litter near a body of water, river debris, humus under maple, Populus litter at the edge of spruce-cedar forest, pine stump buttress, litter near the sea. Collecting period: III–X. Collecting method: sifting forest litter, pitfall traps, Lindgren funnel traps.

20

Tribe Athetini Casey, 1910

Genus Strigota Casey, 1910 (Figs. 20.248a–h and 20.249a–h) (Gusarov 2003a; Klimaszewski et al. 2011, 2018b) Diagnosis. Body elongate, subfusoid (somewhat Oxypoda-like); integument of forebody with punctation fine and dense, pubescence dense, pilose, appearing silky particularly on abdomen; antennae thick, antennomeres V–X more or less transverse; pronotum 1.3 times as wide as long; pronotal hypomeron not visible in lateral view; pronotum and elytra subequal in length; isthmus almost as long as process of mesoventrite; in both sexes antecostal suture of abdominal tergite VIII joins basal margin (always separated from basal margin and extended laterad in other athetines); apical margin of male sternite VIII with row of microsetae on dorsal side. Key to Species of Strigota (Klimaszewski et al. 2018b) 1. Elytra at suture shorter than pronotum length at midline (Fig. 20.249a); body uniformly dark brown to black (Fig. 20.249a); apical margin of male tergite VIII with sharp, narrow triangular projection in about medial one-fourth (Fig. 20.249d); tubus of median lobe of aedeagus strongly bent ventrad in basal one-third in lateral view (Fig. 20.249b); spermathecal capsule globose and sharply delimited from neck, stem with broad posterior loop (Fig. 20.249h) ..................... ..Strigota obscurata Klimaszewski and Brunke – Elytra at suture about as long as pronotal length at midline (Fig. 20.248a); body bicoloured in most, dark brown to black with paler, usually reddish-brown, elytra (Fig. 20.248a); apical margin of male tergite VIII broadly triangularly produced, rounded at apex and slightly pointed (Fig. 20.248d); tubus of median lobe of aedeagus slightly bent ventrad in basal one-fourth in lateral view (Fig. 20.248b); spermathecal capsule slightly globose and not sharply delimited from neck, stem with tight posterior loop (Fig. 20.248h)....... Strigota ambigua (Erichson)

455

212. Strigota 20.248a–h)

ambigua

(Erichson) (Fig.

Homalota ambigua Erichson, 1839. Bernhauer 1907, Casey 1910, 1911 [as nine synonyms], Notman 1922 [as synonym Anaduosternum brevipennis], Gusarov 2003a, Majka et al. 2008, Klimaszewski et al. 2011, 2012, 2016d, 2018b, Brunke et al. 2012, Webster et al. 2016. Diagnosis. Body elongate-oval, moderately convex, length 2.4–3.0 mm; colour uniformly light brown or dark brown to almost black, or forebody, legs/tarsi, antennae and elytra light brown and abdomen dark brown (Fig. 20.248a); punctation on forebody fine and dense, pubescence fine and silky in appearance; head narrower than pronotum, ratio of maximum width of head to maximum width of pronotum 0.7; antennae robust, antennomeres I–IV elongate, V–X slightly to distinctly transverse; pronotum transverse, sides evenly arcuate, ratio of maximum width to length 1.3, about as wide as elytra; elytra at suture slightly shorter than pronotum; abdomen gradually narrowed posteriad. MALE. Tergite VIII moderately elongate, sinuate basally and produced medially at apex (not as sharp as in S. obscurata), antecostal suture joining base of tergite above spiracles (Fig. 20.248d); sternite VIII elongate, rounded apically (Fig. 20.248e); median lobe of aedeagus with tubus straight and less produced ventrad than that of S. obscurata (Fig. 20.248b), in dorsal view bulbus narrowly oval and tubus narrowly subparallel basomedially and triangular with apex rounded (Fig. 20.248c). FEMALE. Tergite VIII slightly triangularly produced, pointed apically (Fig. 20.248f); sternite VIII rounded apically with dense row of microsetae (Fig. 20.248g); spermatheca with club-shaped capsule with a small invagination, nec long and tubular, stem sinuate and coiled posteriorly (Fig. 20.248h). The spermatheca of this species is nearly identical to that of S. obscurata except for the shape of the capsule, which is less sharply deflexed and more tubular.

456

20

DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], LB, NB, NF, NS, ON, PE, QC, SK, YT. USA: CA, CO, CT, IA, KS, MA, MO, NC, NJ, NM, NY, NV, TX. We report this species from BC for the first time here based on the following: British Columbia, Fairview, townsite, pitfall, FCL-67, 18.VI-2. VII.2010, 49.174-119.468, 494 m, coll. David Marks (RBCM) 1 female; same data except: 5. VII-9.VIII.2010, trap 30, 31 (RBCM) 1 male, 1 female; Crownland CL38, pitfall trap 31, 32 (RBCM) 3 females; pitfall FCL2-44, 13.VIII-28. IX.2010 (RBCM) 1 female; Kootenay, Porcupine road service road, 12.IX.2011, Berlese, moss, 49.2646-117.1815, 820 m, coll. R.G.B. Bennett, D.C. Copley (RBCM) 1 male [specimen slightly smaller, tentative identification]. COLLECTION AND HABITAT DATA. Habitat: open habitats with well-drained soil including sand dunes, lakeshores, beaches, Dryas covered limestone barrens, soybean fields, old fields, open gaps in spruce forest with sandy soil mostly covered with Cladina lichens, weedy disturbed field, and riverbanks. Adults under stones, cobblestones on moist sand, in organic debris in groundhog burrows, organic debris, flood refuse, and in beach-drift debris. Collecting period: IV–IX. Collecting method: pitfall traps, malaise traps, sifting various kinds of litter. 213. Strigota obscurata and Brunke (Fig. 20.249a–h)

Klimaszewski

Strigota obscurata Klimaszewski and Brunke, in Brunke et al. 2012. Klimaszewski et al. 2016d, 2018b, Webster et al. 2016. Diagnosis. Body narrowly elongate, length 2.2–2.5 mm; colour dark brown to black, with legs and/or tarsi brown, central disc of elytra sometimes with traces of reddish tinge (Fig. 20.249a); integument moderately glossy, with dense, meshed microsculpture, pubescence short, dense, appearing somewhat silky; head

Tribe Athetini Casey, 1910

convex, rounded posteriorly, eyes at least slightly shorter than genae in dorsal view, pubescence on disc directed towards midline; antennae stout, antennomeres I–III distinctly elongate, IV–V subquadrate and VI–X moderately transverse; pronotum slightly transverse, widest in basal third, pubescence on disc directed posteriad at midline, obliquely posteriad elsewhere; elytra transverse, at suture shorter than pronotum at midline, pubescence on disc directed straight posteriad; abdomen subparallel with first three visible tergites deeply impressed basally; hind tarsus with basal article as long as two following articles combined. MALE. Tergite VIII with bisinuate base and acutely produced apex (Fig. 20.249d); sternite VIII elongate with broad distance between base and antecostal suture, apex truncate (Fig. 20.249e); in lateral view median lobe of aedeagus with bulbus moderate-sized, tubus bent obliquely ventrad before middle of ventral margin, internal sac with several short, inconspicuous sclerites (Fig. 20.249b); in dorsal view median lobe of aedeagus with tubus constricted basally (Fig. 20.249c). FEMALE. Tergite VIII with bisinuate base and acutely produced apex, less than in male (Fig. 20.249f); sternite VIII broadly rounded apically (Fig. 20.249g); spermatheca with spherical capsule bearing a small invagination, stem sinuate and coiled posteriorly (Fig. 20.249h). The spermatheca of this species is nearly identical to that of S. ambigua except for the capsule, which is more spherical and sharply deflexed, and the stem is more broadly coiled. The distinctive shape of tergite VIII of Strigota obscurata will easily differentiate it from the remaining congeners. DISTRIBUTION. Origin: Nearctic. CANADA: BC [new provincial record], NB, ON, QC, SK. USA: not recorded. We report this species from BC for the first time here based on the following: British Columbia, Log Lk., 65 km N of Prince George, VIII-IX. 2012, pitfall # 204, 54.2-122.616, 780 m,

20

Tribe Athetini Casey, 1910

OM3C2SX, sub-boreal spruce, J.M. Kranabetter (RBCM) 1 male.

457

coll.

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: open habitats; in flood debris on a river margin, on soil at the base of grass in a residential lawn, in Lindgren funnel trap in jack pine forest, soybean fields in pitfalls (most common rove beetle in this habitat in southern ON), oak savanna/alvar habitats under stones and in pitfall traps, hedgerow in pitfall traps, and in malaise pans in a field, in sub-boreal spruce forest in BC. Collecting period: V–X. Collecting method: malaise traps, pitfall traps, sifting litter, Lindgren funnel trap.

Genus Tarphiota Casey, 1893 (Figs. 20.250a–g and 20.251a–i) (Mäklin 1852; Casey 1893, 1910; Fenyes 1920; Ahn 1996) Diagnosis. Body narrowly elongate, dark brown to black; integument of forebody strongly granulose due to fine-meshed, raised reticulation; head without infraorbital carinae; eyes large; pronotum 1.2–1.3 times as wide as long, microsetae evenly distributed, macrosetae absent; elytra longer than pronotum, slightly sinuate apico-laterally; hind wings present; antennomeres VII–X slightly to strongly transverse; mandibles each with blunt apex; ligula narrow, Y-shaped; mesocoxae separated, mesocoxal cavities unmargined, mesoventral process pointed; anterior and middle tibiae each with a row of spines; species inhabit mid to upper littoral zone of fine-grained sandy beaches.

pronotum (Fig. 20.251a); antennomeres VII–X distinctly transverse (Fig. 20.251a); male and female tergite VIII with shallow broadly V-shaped apical emargination (Fig. 20.251d, f) ........................... Tarphiota geniculata (Mäklin)

214. Tarphiota 20.250a–g)

fucicola

(Mäklin) (Fig.

Tachyusa fucicola Mäklin, 1852. Casey 1893 [as T. pallidipes], Casey 1910 [as T. debilicollis], Fenyes 1920, Ahn 1996. Diagnosis. Body length 2.9–3.1 mm; dark brown to black with brown legs and antennae (Fig. 20.250a); integument of forebody strongly granulose; head about 1.2 times as wide as long; antennae slender, antennomeres VII–X subquadrate; pronotum about 1.3 times as wide as long, sides rounded; elytra 1.3 times as long as pronotum; abdominal tergites with evenly distributed pubescence, several macrosetae present. MALE. Tergite VIII truncate apically (Fig. 20.250c); sternite VIII rounded apically (Fig. 20.250d); in lateral view tubus of median lobe of aedeagus gently arcuate ventrally and with narrow and pointed apex slightly produced ventrad (Fig. 20.250b). FEMALE. Tergite VIII broadly arcuate apically (Fig. 20.250e); sternite VIII rounded apically (Fig. 20.250f); spermatheca with medium-sized globular capsule and short elbowed neck, stem very narrow and long (Fig. 20.250g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA.

Key to Species of Tarphiota 1. Body length 2.9–3.1 mm; maximum width of elytra one-fourth wider that maximum width of pronotum (Fig. 20.250a); antennomeres VII–X subquadrate (Fig. 20.250a); male and female tergite VIII without apical emargination (Fig. 20.250c, e); spermathecal stem narrow (Fig. 20.250g)........ Tarphiota fucicola (Mäklin) – Body length 2.0–2.6 mm; maximum width of elytra one-sixth wider that maximum width of

COLLECTION AND HABITAT DATA. Habitat: under rotting seaweed on sandy beaches. Collecting period: III, V–VIII, XI. Collecting method: sifting seaweed. 215. Tarphiota 20.251a–i)

geniculata

(Mäklin) (Fig.

Homalota geniculata Mäklin, 1852. Casey 1910 [as T. iota, T. insolita, T. litorina,

458

T. seditiosa], Ahn 1996.

20

Fenyes

1920,

Hatch

1957,

Diagnosis. Body length 2.0–2.6 mm; dark brown to black with legs and antennae brown (Fig. 20.251a); integument of forebody strongly granulose; head about 1.2 times as wide as long; antennae slender, antennomeres VII–X distinctly transverse; pronotum about 1.2 times as wide as long, sides rounded; elytra 1.2 times as long as pronotum; abdominal tergites with evenly distributed pubescence, several macrosetae present. MALE. Tergite VIII truncate apically and with broadly V-shaped shallow median emargination (Fig. 20.251d); sternite VIII triangularly produced apically (Fig. 20.251e); tubus of median lobe of aedeagus gently sinuate ventrally and with narrow and rounded apex, slightly produced ventrad in lateral view (Fig. 20.251b), in dorsal view bulbus and tubus broad (Fig. 20.251c). FEMALE. Tergite VIII apically emarginate medially (Fig. 20.251f); sternite VIII rounded apically (Fig. 20.251g); spermatheca with small globular capsule bearing apical invagination, neck arcuate, stem moderately broad and moderately long and sinuate (Fig. 20.251h, i). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA. In BC, it was recorded from Queen Charlotte Is., Massett, and Sidney (Ahn 1996). New locality data: British Columbia, Queen Charlotte Is., 3.5 km SW Tow Hill, 22. VIII.1983, 83-110, under debris and kelp on beach, J.M. Campbell (CNC) 1 male, 1 female, 7 sex undetermined; Queen Charlotte Is., Tlell, 17.VII.1983, 83-16, ex rotting seaweed, J.M. Campbell (CNC) 1 female; Queen Charlotte Is., NW corner Graham Is., Lepas Bay, 17. VII.1983, 83-93, under kelp on beach, J.M. Campbell (CNC) 1 male, 1 female; Vancouver Is., Ucluelet, 17.X.2008, under wood on sea beach, A. Davies (CNC) 1 male, 2 sex undetermined; Vancouver Is., Long Beach, 12 mi S Tofino, 23.V.1968, A. Smetana (CNC) 1 male, 2 females, 14 sex undetermined.

Tribe Athetini Casey, 1910

COLLECTION AND HABITAT DATA. Habitat: under rotting seaweed and wood on sandy sea beaches, running on water swept shore. Collecting period: IV–VIII, X. Collecting method: sifting seaweed.

Genus Thinusa Casey, 1893 (Figs. 20.252–20.254) (Casey 1893; Moore and Legner 1976; Seevers 1978; Ahn 1997; Ashe 2001) Diagnosis. Body narrowly subparallel, length 1.8–3.1 mm, integument granulose with coarse microsculpture; pubescence short and more or less uniformly distributed; head slightly deflexed, with incomplete infraorbital carinae, antennomeres V–X strongly transverse; mandible with blunt apex; ligula narrow, Y-shaped; mentum with long V setae; elytra shorter than pronotum, slightly sinuate apico-laterally, hind wings absent; mesocoxae contiguous; mesoventral process pointed; mesocoxal cavities unmarginated; anterior and middle tibiae with row of spines. Species occur in the intertidal zone of sandy beaches from Alaska to Baja California. Key to Species of Thinusa (Modified from Ahn 1997) 1. Body large, length 2.4–3.1 mm (Fig. 20.252a); male tergite VIII truncate apically, margin straight (Fig. 20.252d); tubus of median lobe of aedeagus with subapical dent in lateral view (Fig. 20.252b, c)........... Thinusa fletcheri Casey – Body small, length 1.8–2.2 mm (Figs. 20.253a and 20.254a); male tergite VIII truncate apically, margin shallowly emarginated medially (Fig. 20.253c); tubus of median lobe of aedeagus without subapical dent in lateral view (Fig. 20.253b)........... Thinusa maritima (Casey)

216. Thinusa fletcheri Casey (Fig. 20.252a–h) Thinusa fletcheri Casey, 1906. Casey 1911 [as T. divergens, T. nigra, T. robustula], Fenyes

20

Tribe Athetini Casey, 1910

1920, Hatch 1957, Moore and Legner 1976, Ahn 1997. Lectotype (female): Canada, British Columbia: QCI [Queen Charlotte Islands]; Phytosus fletcheri n. sp. Fvl. [Fauvel]; fletcheri-2, Paratype USNM 38777; Casey bequest 1925; Lectotype Thinusa fletcheri Casey, designated by K.J. Ahn 1995 (USNM). Examined and illustrated. Thinusa robustula Casey, 1911. Synonymized by Ahn 1997. Confirmed same identity as the lectotype of T. fletcheri. Lectotype (female): Metlakatla, B. Columbia; Casey determ. fletcheri 6; Casey bequest 1925; robustula-2, USNM TYPE 38778; Lectotype Thinusa robustula Casey, desig. K.J. Ahn 1995. Examined. Diagnosis. Body narrowly subparallel, length 3.1 mm; abdomen dark brown, head, pronotum and elytra paler-red brown, legs yellow brown, antennomeres I–III yellow brown and IV–XI brown (Fig. 20.252a); head 1.2 times as wide as long, eyes large; mandibles robust, apex hooked and blunt; pronotum broadest in apical third, about one-fifth broader than long; elytra along suture distinctly shorter than pronotum along midline, about one-fourth shorter than length of pronotum along midline; abdomen subparallel, tergites with microsetae uniformly distributed, macrosetae scattered. MALE. Tergite VIII truncate apically (Fig. 20.252d); sternite VIII broadly arcuate apically, distance between base of disc and antecostal suture wide (Fig. 20.252e); in lateral view tubus of median lobe of aedeagus arcuate ventrally, with sharp median dent, subapical part of tubus moderately narrow and gradually produced ventrad, apex moderately broad and rounded (Fig. 20.252b, c). FEMALE (based on the lectotype). Tergite VIII truncate apically (Fig. 20.252f); sternite VIII broadly triangularly produced apically, apex rounded (Fig. 20.252g); spermatheca with club-shaped capsule, apical invagination large and moderately deep, stem wide, S-shaped (Fig. 20.252h).

459

DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA. Confirmed locality data: BC, Queen Charlotte Is., Massett, 20641, Thinusa fletcheri, A. Fenyes coll., fletcheri Casey, Smiths. In., Thinusa det. K.J. Ahn 1997 (CAS) 1 male. COLLECTION AND HABITAT DATA. Habitat: sandy beaches with seaweed. Collecting period: not recorded. Collecting method: sifting seaweed. Comments. Recorded from BC by Ahn (1997) from Queen Charlotte Is., Massett (CAS) 1 male, which we have dissected and confirmed its identity. We have examined the female lectotype of Thinusa fletcheri Casey and it does not match the shape of the spermatheca illustrated by Ahn (1997). We confirm that the male illustrated by Ahn (1997) belongs to this species. We confirm synonymy of T. robustula Casey, with T. fletcheri, by Ahn 1997. The lectotype of T. robustula is slightly smaller, length 2.4 mm, but morphologically conforms with T. fletcheri. 217. Thinusa maritima (Casey) (Figs. 20.253a– d and 20.254a–g) Phytosus maritimus Casey 1885. As Thinusa: Casey 1893, Fenyes 1920, Hatch 1957, Moore and Legner 1976, Ahn 1997. Lectotype (male): Cal. [ifornia]; Thinusa maritima Csy.; TYPE USNM 38775; Casey bequest 1925; Lectotype, Thinusa maritima (Casey), desig. K.J. Ahn 1995 (USNM). Examined and illustrated. Diagnosis. Body narrowly subparallel, length 1.8–2.2 mm; colour variable, from forebody reddish brown and abdomen dark brown to body uniformly brown to black (Figs. 20.253a and 20.254a); head 1.3 times as wide as long; mandibles robust, apex hooked and sharply pointed; pronotum about one-fourth broader than long; elytra along suture distinctly shorter

460

than pronotum, about one-fourth shorter than length of pronotum along midline; abdominal subparallel, tergites with microsetae uniformly distributed, macrosetae scattered. MALE (based on male lectotype). Tergite VIII arcuately emarginated apically (Figs. 20.253c and 20.254c); sternite VIII rounded apically (Figs. 20.253d and 20.254d); tubus of median lobe of aedeagus arcuate without sharp dent, subapical part narrowly elongate and apex narrowly rounded in lateral view (Figs. 20.253b and 20.254b). FEMALE. Tergite VIII truncate apically, antecostal suture strongly sinuate (Fig. 20.254e); sternite VIII slightly produced apically but not distinctly pointed medially (Fig. 20.254f); spermatheca with capsule tubular and broadest in the middle, apical invagination deep, stem narrow, moderately long, S-shaped (Fig. 20.254g). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: AK, CA, WA. MEXICO. Baja California. New locality data: British Columbia, Vancouver Is., Ucluelet, 17.X.2008, under wood on sea beach, A. Davies (CNC) 1 male, 2 females, 7 sex undetermined; Vancouver Is., 15 km SE Tofino, Long Beach, 49 040 0300 N,  0 00 125 48 02 W, 16.X.2011, under wood near high tide line on sea beach, A. Davies (CNC) 2 males; Vancouver Is., Long Beach, 12 mi S Tofino, 23.V.1968, A. Smetana (CNC) 2 males. Confirmed locality data: BC, Queen Charlotte Is., Massett, 20640, Rev. Keene coll., A. Fenyes coll., Thinusa fletcheri Casey, Thinusa maritima (Casey) det. K.J. Ahn 1995 (CAS) 1 male. COLLECTION AND HABITAT DATA. Habitat: under decaying seaweed on sandy beaches, sandspit under damp driftwood, on wave swept shore, surf saturation zone on beach, wrack and beach debris. Collecting period: IV–X. Collecting method: sifting seaweed on beaches, under wood on sea beach.

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Comments. Recorded by Ahn (1997) from Queen Charlotte Is., Massett, Rev. Keen (CAS), 1 male, which we have dissected and confirmed its identity. The female lectotype of this species matches the description of T. maritima by Ahn (1997), and specimens from Ucluelet, Vancouver Island, listed above (CNC). It is the smallest species of this genus.

Genus Trichiusa Casey, 1893 (Figs. 20.255a–d and 20.256a–h) (Casey 1893, 1906; Brunke et al. 2012; Schülke and Smetana 2015; Webster et al. 2016; Klimaszewski et al. 2018b) Diagnosis. Body compact, broad, on average 3.0 mm long, often with long, dense and shaggy pubescence, on pronotum directed anteriad along midline of disc and often erected; body form distinctive with head and pronotum distinctly narrower than elytra, elytra at least 1.2 times as wide as pronotum; mesocoxae widely separated; abdomen robust, oval in outline, broadest at segment V and at least as broad as elytra at widest point; median lobe of aedeagus in dorsal view with bulbus of moderate-size and tubus elongate, and apically triangular, in lateral view tubus long and arcuate ventrally (Figs. 20.255b and 20.256b); spermatheca with tubular or spherical capsule bearing moderate to large apical invagination, stem thin, sinuate and looped or twisted posteriad. Key to Species of Trichiusa 1. Body uniformly dark brown; pubescence on pronotum straight and strongly erect (Fig. 20.256a)................. Trichiusa pilosa Casey – Body bicolored, head, pronotum, apical part of abdomen dark brown and posterior part of elytra, basal abdomen and legs red brown; pubescence on pronotum not strongly erect (Fig. 20.255a) Trichiusa columbica Casey

218. Trichiusa 20.255a–d)

columbica

Casey (Fig.

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Tribe Athetini Casey, 1910

Trichiusa columbica Casey, 1911. Lectotype (male): British Columbia, Metlakatla, Keen; Type USNM 39425; Casey bequest 1925; Trichiusa columbica Casey, V.I. Gusarov des. 2000, lectotype (USNM). Lectotype, present designation. Diagnosis (based on male lectotype). Body broad, ovoid in shape, length 2.0 mm; colour dark brown, except paler, reddish brown posterior part of elytra, base of abdomen, legs and antennomeres I–III (Fig. 20.255a); sparsely punctate and pubescent; setae long, not strongly erect; integument moderately glossy; head narrower than pronotum; pronotum distinctly transverse, rounded laterally and basally, slightly narrower than elytra; elytra broad and at suture about as long as pronotum; abdomen arcuate laterally and broadest at middle of its length. MALE. Tergite VIII strongly transverse and broadly arcuate apically, setae moderately long (Fig. 20.255c); sternite VIII parabolic apically (Fig. 20.255d); median lobe of aedeagus with tubus gently arcuate ventrally, narrowly triangular and narrowly rounded apically in lateral view (Fig. 20.255b); internal sac structures weakly defined (Fig. 20.255b). FEMALE. Undescribed. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: not recorded. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded. Comments. This species is known only from the type locality—Metlakatla, BC. Casey described T. geniculata from a single male from Metlakatla. Gusarov added his lectotype designation label and the identification label as Trichiusa geniculata (Mäklin). However, Homalota geniculata Mäklin belongs to the genus Thinusa as correctly pointed out by Lohse and Smetana (1985). We formally designate Casey’s type of T. columbica as the lectotype. Trichiusa columbica is generally similar to T. hirsuta

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Casey (not recorded from BC but from eastern Canada). However, T. hirsuta has the pronotum more strongly converging apically, male tergite VIII has longer apical setae, and the tubus of the median lobe of the aedeagus is sharply pointed. Due to these differences, we have considered T. columbica as a distinct species. 219. Trichiusa pilosa Casey (Fig. 20.256a–h) Trichiusa pilosa Casey, 1893. Casey 1906 and auct. before 2015 [as synonym T. postica], Klimaszewski et al. 2015a, 2018b, Webster et al. 2016. Diagnosis. Body broad, oval, length 1.5–1.8 mm; colour dark brown with reddish tinge, base of abdomen slightly paler, or uniformly black, appendages usually lighter than rest of body (Fig. 20.256a); sparsely punctate and pubescent; setae straight and erect, particularly on pronotum; integument highly glossy; head slightly narrower than pronotum; pronotum moderately transverse, rounded laterally and basally, distinctly narrower than elytra; elytra broad and at suture about as long as pronotum; abdomen arcuate laterally and broadest at middle of its length. MALE. Tergite VIII strongly transverse and truncate apically (Fig. 20.256d); sternite VIII parabolic and slightly produced apically, apex rounded (Fig. 20.256e); in lateral view tubus of median lobe of aedeagus slightly sinuate and moderately narrow apically with apex rounded (Fig. 20.256b), in dorsal view bulbus narrowly oval and tubus narrowly triangularly shaped, apex rounded (Fig. 20.256c); internal sac structures weakly defined (Fig. 20.256b, c). FEMALE. Tergite VIII broad and truncate apically (Fig. 20.256f); sternite VIII broadly parabolic apically (Fig. 20.256g); spermatheca with broad and spherical capsule with minute apical invagination, neck elbowed, stem narrow, sinuate and tightly coiled posteriorly (Fig. 20.256h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, NB, NS, ON. USA: ID, IN, KS, OH, RI.

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COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018b). Habitat: NB specimens from among cobblestones on lakeshore and from a Lindgren funnel trap in canopy of Populus tremuloides in a hardwood forest; one individual from bison dung in BC and another from a window trap in a 10-year-old regenerating white spruce stand in AB. Collecting period: VI– IX. Collecting method: sifting leaf litter, Lindgren funnel traps, window trap. Athetini with Subtribe affiliation uncertain (At present, phylogenetic position of Paragoniusa within the Athetini could not be firmly established, because of the insufficient phylogenetic knowledge of the tribe. Although Paragoniusa and Goniusa are externally similar in sub-limuloid body shape, this character state is commonly observed in other myrmecophilous staphylinids, and it is possible that the state is convergent)

Genus Paragoniusa Maruyama and Klimaszewski, 2004 (Fig. 20.257a–g) (Maruyama and Klimaszewski 2004) Diagnosis. Body broad, sub-limuloid, uniformly reddish brown; head strongly transverse, temples considerably produced laterad; surface moderately convex, finely sculptured, pubescence short; antennomeres III–X transverse and XI without coeloconic sensilla; labrum transverse, anterior margin straight; mandibles asymmetrical; ligula bifid apically; mentum with anterior margin sinuate; pronotum transverse, 1.8–1.9 times wider than long, pubescence directed posteriad medially and posterolaterad laterally, macrosetae absent; pronotal hypomera invisible in lateral view; metaventrite process very short and slightly produced; abdominal tergite VIII with short and inconspicuous macrosetae; tarsal formula 4-5-5;

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Tribe Athetini Casey, 1910

basal tarsomere of hind leg shorter than second, one empodial seta present; proximal part of spermathecal short and not coiled. Paragoniusa is similar to Goniusa in general appearance but may be distinguished by its larger body, transverse head with produced temples, transverse antennomeres III–X; elongate palpomere III of labial palpi, laterally invisible pronotal hypomera, very short mesoventral process, absent macrosetae of mesotibia, inconspicuous and not suberect macrosetae on abdominal sternites III–VII, and by the proximal part of the spermatheca short and not coiled. 220. Paragoniusa myrmicae Maruyama and Klimaszewski (Fig. 20.257a–g) Paragoniusa myrmicae Maruyama and Klimaszewski, 2004 (Maruyama and Klimaszewski 2006: 126–AB, BC records; Klimaszewski et al. 2011: 172–LB records; Webster et al. 2016: 139–NB records). Diagnosis. Body length 4.5–5.1 mm; reddish brown, head, scutellum, basal areas of elytra and abdominal segments V and VI brownish black, and mouthparts and antennae paler (Fig. 20.257a); head strongly transverse, widest at temples, slightly depressed above, surface without short setae in mesal area; antennae about as long as head and pronotum combined; antennomere I slightly longer than II; antennomere II almost as long as III; antennomeres IV–X slightly transverse; pronotum strongly transverse, widest just after middle, about 1.5 times as wide as head, with lateral areas flattened; elytra wide, almost as wide as pronotum, widest just behind lateral corners. MALE. Tergite VIII transverse, apical margin arcuate, convex, lateral corners slightly produced (Fig. 20.257d); median lobe of aedeagus oval in outline in dorsal view (Fig. 20.257c); in lateral view tubus with basal dent basally and evenly narrow apically in lateral view (Fig. 20.257b).

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Fig. 20.155 (a–h) Images of Adota maritima (Mannerheim): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in ventral view; (d)

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male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.156 (a–g) Images of Aloconota cambrica (Wollaston): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

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Tribe Athetini Casey, 1910

female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

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Fig. 20.157 (a–h) Images of Aloconota carmanah Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

FEMALE. Tergite VIII arcuate apically, slightly emarginated medially (Fig. 20.257e); sternite VIII

rounded apically and slightly emarginated medially (Fig. 20.257f); spermatheca with proximal

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Fig. 20.158 (a–e) Images of Amischa analis (Gravenhorst): (a) habitus in dorsal view (apical part of abdomen removed); (b) female tergite VIII; (c) female

sternite VIII; (d) spermatheca (Canada), (e) (Europe). Male unknown in North America. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

portion of capsule slightly curved, narrowed apicad, apical portion spherical, 3/5 as long as

proximal portion, its inner wall densely wrinkled,

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Tribe Athetini Casey, 1910

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Fig. 20.159 (a–h) Images of Atheta (Alaobia) ventricosa Bernhauer: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

stem about straight and swelled posteriorly (Fig. 20.257g).

DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, NB. USA: not recorded.

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Fig. 20.160 (a–h) Images of Atheta (Atheta) graminicola (Gravenhorst): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

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Tribe Athetini Casey, 1910

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.161 (a–i) Images of Atheta (Atheta) ringi Klimaszewski: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male

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sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

470

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Tribe Athetini Casey, 1910

Fig. 20.162 (a–i) Images of Atheta (Bessobia) cryptica (Lohse): (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male

tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

COLLECTION AND HABITAT DATA. Habitat: ant nest in rotten log,

myrmecophilous species associated with Myrmica ants. Collecting period: V. Collecting

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Tribe Athetini Casey, 1910

Fig. 20.163 (a–h) Images of Atheta (Chaetida) longicornis (Gravenhorst): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

method: sifting leaf litter, Lindgren funnel traps, window trap.

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.164 (a–h) Images of Atheta (Datomicra) celata (Erichson): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

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Tribe Athetini Casey, 1910

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.165 (a–h) Images of Atheta (Datomicra) dadopora C. G. Thomson: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

473

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.166 (a–h) Images of Atheta (Dimetrota) alesi Klimaszewski and Brunke: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

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Tribe Athetini Casey, 1910

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.167 a-h. Images of Atheta (Dimetrota) remulsa Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; d, male tergite VIII; e, male sternite VIII; (f)

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female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.168 (a–h) Images of Atheta (Dimetrota) stercoris Fenyes: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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Tribe Athetini Casey, 1910

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.169 (a–h) Images of Atheta (Dimetrota) winchesteri Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

477

sternite VIII; f, female tergite VIII; g, female sternite VIII; (h) spermatheca. (b–e) holotype, (a, f–h) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.170 (a–h) Images of Atheta (Dimetrota) godini Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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Tribe Athetini Casey, 1910

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) female pygidium. (b–e) holotype, (a, f–h) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.171 (a–h) Images of Atheta (Dimetrota) metlakatlana Bernhauer: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.172 (a–h) Images of Atheta (Dimetrota) pseudometlakatlana Klimaszewski and Godin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe

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Tribe Athetini Casey, 1910

of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.173 (a–h) Images of Atheta (Dimetrota) wheelerae Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

481

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–d) holotype, (c, f–h) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.174 (a–h) Images of Atheta (Dimetrota) districta Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

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Tribe Athetini Casey, 1910

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.175 (a–h) Images of Atheta (Dimetrota) fanatica Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

483

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

484

Fig. 20.176 (a–h) Images of Atheta (Dimetrota) munsteri Bernhauer: (a) male habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

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Tribe Athetini Casey, 1910

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–e) lectotype; (f–h) paralectotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.177 (a–h) Images of Atheta (Dimetrota) hampshirensis Bernhauer: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

485

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.178 (a–h) Images of Atheta (Dimetrota) picipennis (Mannerheim) [¼A. cursor (Mäklin)]: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of

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Tribe Athetini Casey, 1910

aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.179 (a–h) Images of Atheta (Dimetrota) brumalis Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.180 (a–h) Images of Atheta (Dimetrota) capsularis Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

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Tribe Athetini Casey, 1910

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.181 (a–h) Images of Atheta (Dimetrota) prudhoensis (Lohse): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.182 (a–e) Images of male holotype of Atheta (Dimetrota) rurigena Casey with original labels: (a) abdomen and hind legs (forebody missing); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

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Tribe Athetini Casey, 1910

dorsal view; (d) male tergite VIII; (e) male sternite VIII. Scale bar for abdomen ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.183 (a–i) Images of Atheta (Dimetrota) strigosula Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

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(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (h) after Klimaszewski et al. (2005). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 20.184 (a–h) Images of Atheta (Dimetrota) terranovae Klimaszewski and Langor: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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Tribe Athetini Casey, 1910

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.185 (a–j) Images of Atheta (Hydrosmectomorpha) meduxnekeagensis Webster and Klimaszewski: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view (b—BC, c—NB); (d, e) male tergite VIII (d—NB, e—BC); (f) male sternite VIII; (g, h) female

493

tergite VIII (g—NB, h—BC); (i) female sternite VIII; (j) spermatheca. (a–i) after Klimaszewski et al. (2018). Scale line for habitus ¼ 1 mm, remaining scale lines ¼ 0.2 mm. Arrows indicate important diagnostic features

494

Fig. 20.186 (a–h) Images of Atheta (Lamiota) keeni Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

20

Tribe Athetini Casey, 1910

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.187 (a–e) Images of Atheta (Lamiota) relicata (Casey) (lectotype): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in

495

lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

496

Fig. 20.188 (a–h) Images of Atheta (Metadimetrota) cheersae Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

20

Tribe Athetini Casey, 1910

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.189 (a–g) Images of Atheta (Microdota) copleyi Klimaszewski: (a) habitus of male in dorsal view (holotype) (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

497

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. (a–d) holotype, (e–g) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

498

Fig. 20.190 (a–h) Images of Atheta (Microdota) holmbergi Bernhauer: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

20

Tribe Athetini Casey, 1910

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a, b, d–e) holotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.191 (a–h) Images of Atheta (Microdota) lucifera Bernhauer: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

499

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

500

Fig. 20.192 (a–h) Images of Atheta (Microdota) platonoffi Brundin: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

20

Tribe Athetini Casey, 1910

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.193 a-h. Images of Atheta (Microdota) ripariides Newton: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

501

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

502

Fig. 20.194 (a–e) Images of Atheta (Microdota) surgens (Casey) (lectotype): (a) habitus of male in dorsal view (missing some legs and apical part of abdomen); (b) median lobe of aedeagus in lateral view; (c) median lobe

20

Tribe Athetini Casey, 1910

of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.195 a-h. Images of Atheta (Pancota) nescia Casey: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

503

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c) lectotype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

504

Fig. 20.196 (a–h) Images of Atheta (Pseudota) concessa Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite

20

Tribe Athetini Casey, 1910

VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.197 (a–h) Images of Atheta (Pseudota) pseudoklagesi Klimaszewski and Webster: (a) habitus in dorsal view (missing abdominal apex); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

505

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

506

Fig. 20.198 (a–j) Images of Atheta (Tetropla) frosti Bernhauer: (a) habitus in dorsal view (missing abdominal apex); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite

20

Tribe Athetini Casey, 1910

VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i, j) spermatheca. (c, d, i) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.199 (a–h) Images of Atheta (subgenus uncertain) brunswickensis Klimaszewski: (a) habitus in dorsal view (missing abdominal apex); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

507

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

508

Fig. 20.200 (a–i) Images of Atheta (subgenus uncertain) irrupta (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view;

20

Tribe Athetini Casey, 1910

(e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (c) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.201 (a–h) Images of Atheta (subgenus uncertain) pseudoschistoglossa Klimaszewski and Webster: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

509

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

510

Fig. 20.202 (a–i) Images of Palearctic Atheta (subg. undetermined) basicornis (Mulsant and Rey) (apical part of abdomen removed) [listed erroneously in Atheta s. str.], not confirmed in BC: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe

20

Tribe Athetini Casey, 1910

of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (c, h) after Pal (1970). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.203 (a–g) Images of Palearctic Atheta (Dimetrota) subrugosa (Märkel and Kiessenwetter), not confirmed in BC: (a) habitus in dorsal view; (b, c) median lobe of aedeagus in lateral view; (d) median lobe of

511

aedeagus in dorsal view; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. (c, d) after Brundin (1953). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

512

Fig. 20.204 (a–f) Images of Boreophilia nomensis (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c, d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male sternite VIII.

20

Tribe Athetini Casey, 1910

(a, d, e, f) after Klimaszewski et al. (2019); (b, c) after Lohse et al. (1995). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.205 (a–h) Images of Boreostiba pseudolaticollis Klimaszewski and Godin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

513

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; h, spermatheca. (a–h) paratypes. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

514

Fig. 20.206 (a–h) Images of Boreostiba parvipennis (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

20

Tribe Athetini Casey, 1910

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.207 (a–h) Images of Clusiota impressicollis (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

515

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

516

Fig. 20.208 (a–g) Images of Dalotia coriaria (Kraatz): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

20

Tribe Athetini Casey, 1910

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.209 (a–g) Images of Dinaraea angustula (Gyllenhal): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

517

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

518

Fig. 20.210 (a–d) Images of Dinaraea inexpectata Klimaszewski (holotype): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of

20

Tribe Athetini Casey, 1910

aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.211 (a–g) Images of Dinaraea pacei Klimaszewski and Langor: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

519

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

520

Fig. 20.212 (a–g) Images of Dinaraea subdepressa (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

20

Tribe Athetini Casey, 1910

female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.213 (a–h) Images of Dochmonota rudiventris (Eppelsheim): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

521

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

522

Fig. 20.214 (a–h) Images of Earota dentata (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

20

Tribe Athetini Casey, 1910

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (c) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.215 (a–h) Images of Geostiba horwoodae Klimaszewski and Godin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

523

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–e) holotype, (f–h) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

524

Fig. 20.216 (a–g) Images of Goniusa caseyi Gusarov: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view;

20

Tribe Athetini Casey, 1910

(c) male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.217 (a–d) Images of Liogluta atriventris (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

525

male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

526

Fig. 20.218 (a–f) Images of Liogluta nitens (Mäklin): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

20

Tribe Athetini Casey, 1910

(e) female tergite VIII; (f) female sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.219 (a–g) Images of Liogluta quadricollis (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

527

male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

528

Fig. 20.220 (a–g) Images of Liogluta scudderi Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

20

Tribe Athetini Casey, 1910

female tergite VIII; (f) female sternite VIII; (g) spermatheca. (a–d) holotype, (e–g) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.221 (a–g) Images of Liogluta terminalis (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c)

529

male tergite VIII; (d) male sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

530

Fig. 20.222 (a–h) Images of Liogluta trapezicollis Lohse: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d) male tergite VIII; (e) male sternite VIII; (f) female

20

Tribe Athetini Casey, 1910

tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.223 (a–g) Images of Liogluta wickhami (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

531

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

532

Fig. 20.224 (a–i) Images of Lypoglossa angularis (Mäklin): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in ventral view; (d) apical part of median lobe of aedeagus in

20

Tribe Athetini Casey, 1910

ventral view; (e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (b–d, i) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.225 (a–i) Images of Lypoglossa franclemonti Hoebeke: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in ventral view; (d) median lobe of aedeagus in dorsal view;

533

(e) male tergite VIII; (f) male sternite VIII; (g) female tergite VIII; (h) female sternite VIII; (i) spermatheca. (b, c) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

534

Fig. 20.226 (a–h) Images of Mocyta breviuscula (Mäklin): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

20

Tribe Athetini Casey, 1910

female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.227 (a–j) Images of Mocyta fungi (Gravenhorst): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) female

535

tergite VIII; (e) female sternite VIII; (f–j) spermatheca. (b, c) based on European specimen (after Brundin 1940), males do not occur in North America. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

536

Fig. 20.228 (a–g) Images of Mocyta discreta (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

20

Tribe Athetini Casey, 1910

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.229 (a–g) Images of Nehemitropia lividipennis (Mannerheim): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

537

female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

538

Fig. 20.230 (a–g) Images of Ousipalia pacifica Casey: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

20

Tribe Athetini Casey, 1910

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.231 (a–g) Images of Paraleptonia pacei Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b, c) median lobe of aedeagus in lateral view; (d, e) median lobe of aedeagus in dorsal

539

view; (f) male tergite VIII; (g) male sternite VIII. (c, e) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

540

Fig. 20.232 (a–h) Images of Pelioptera (Tropimenelytron) robusta (Gusarov): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

20

Tribe Athetini Casey, 1910

ventral view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b–e) after Gusarov (2002a, b, c). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.233 (a–g) Images of Philhygra angusticauda (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

541

female tergite VIII; (f) female sternite VIII; (g) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

542

Fig. 20.234 (a–h) Images of Philhygra botanicarum (Muona): (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

20

Tribe Athetini Casey, 1910

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.235 (a–h) Images of Philhygra charlottae Klimaszewski: (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view;

543

(d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) female pygidium. (a–e) holotype; (f–h) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

544

Fig. 20.236 (a–g) Images of Philhygra clemens (Casey): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e) female

20

Tribe Athetini Casey, 1910

tergite VIII; (f) female sternite VIII; (g) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.237 (a–i) Images of Philhygra laevicollis (Mäklin) [BC specimens]: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

545

female tergite VIII; (g) female sternite VIII; (h) female pygidium; (i) membraneous structures of female pygidium. (a–e) holotype; (c, i) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

546

Fig. 20.238 (a–g) Images of Philhygra terrestris Klimaszewski and Godin: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male

20

Tribe Athetini Casey, 1910

sternite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.239 (a–g) Images of Philhygra terrivaga Klimaszewski, sp. n.: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

547

(e) female tergite VIII; (f) female sternite VIII; (g) female pygidium. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

548

Fig. 20.240 (a–i) Images of Pontomalota opaca (LeConte): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

20

Tribe Athetini Casey, 1910

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (i) after Ahn and Ashe (1992). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.241 (a–h) Images of Psammostiba comparabilis (Mäklin): (a) habitus of male in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in ventral view; (d)

549

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

550

Fig. 20.242 (a–h) Images of Psammostiba kenaii Gusarov: (a) habitus of male in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

20

Tribe Athetini Casey, 1910

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.243 (a–h) Images of Schistoglossa (Boreomorpha) carexiana Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

551

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

552

Fig. 20.244 (a–h) Images of Schistoglossa (Schistoglossa) charlottae Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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Tribe Athetini Casey, 1910

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.245 (a–h) Images of Schistoglossa (Schistoglossa) campbelli Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

553

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

554

Fig. 20.246 (a–h) Images of Schistoglossa (Schistoglossa) hampshirensis Klimaszewski: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

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Tribe Athetini Casey, 1910

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.247 (a–h) Images of Seeversiella globicollis (Bernhauer): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

555

tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, d–h) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

556

Fig. 20.248 (a–h) Images of Strigota ambigua (Erichson): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d)

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Tribe Athetini Casey, 1910

male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, f–h) after Gusarov (2003a). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.249 (a–h) Images of Strigota obscurata Klimaszewski and Brunke: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

557

dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

558

Fig. 20.250 (a–g) Images of Tarphiota fucicola (Mäklin): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (e)

20

Tribe Athetini Casey, 1910

female tergite VIII; (f) female sternite VIII; (g) spermatheca. (g) after Ahn (1997). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.251 (a–i) Images of Tarphiota geniculata (Mäklin): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f)

559

female tergite VIII; (g) female sternite VIII; (h, i) spermatheca. (h) after Ahn (1997). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

560

Fig. 20.252 (a–h) Images of Thinusa fletcheri Casey: (a) habitus in dorsal view (female); (b, c) median lobe of aedeagus in lateral view; (d) male tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite

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Tribe Athetini Casey, 1910

VIII; (h) spermatheca. (a, f–h) lectotype (USNM); (c) after Ahn (1997). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.253 (a–d) Images of Thinusa maritima (Casey) [male lectotype]: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d)

561

male sternite VIII. Specimen from California with photographs of the labels. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

562

Fig. 20.254 (a–g) Images of Thinusa maritima (Casey) [specimens from BC]: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII; (f) female tergite VIII; (g)

20

Tribe Athetini Casey, 1910

female sternite VIII; (h) spermatheca. Specimens from British Columbia. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

20

Tribe Athetini Casey, 1910

Fig. 20.255 (a–d) Images of Trichiusa columbica Casey (lectotype): (a) habitus in dorsal view (middle tarsi missing apical tarsomeres, hind legs reconstructed; apical part of abdomen removed); (b) median lobe of aedeagus in

563

lateral view; (c) male tergite VIII; (d) male sternite VIII. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

564

Fig. 20.256 (a–h) Images of Trichiusa pilosa Casey: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e)

20

Tribe Athetini Casey, 1910

male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Tribe Athetini Casey, 1910

Fig. 20.257 (a–g) Images of Paragoniusa myrmicae Maruyama and Klimaszewski: (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in

565

dorsal view; (d) male tergite VIII; (e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. (b–d) after Maruyama and Klimaszewski (2004). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

566

References Ahn K-J (1996) Revision of the intertidal aleocharine genus Tarphiota (Coleoptera: Staphylinidae). Entomol News 107:177–185 Ahn K-J (1997) Revision and systematic position of the intertidal genus Thinusa Casey (Coleoptera: Staphylinidae: Aleocharinae). Entomol Scand 28:75–81 Ahn KJ, Ashe JS (1992) Revision of the intertidal aleocharine genus Pontomalota Casey (Coleoptera: Staphylinidae) with a discussion of its phylogenetic relationships. Entomol Scand 23:347–359 Ashe JS (1986) Seeversiella bispinosa, a new genus and species of athetine Aleocharinae (Coleoptera: Staphylinidae) from North America. J NY Entomol Soc 94(4):500–511 Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (Eds.) American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Assing V (2001) A revision of the Turkish species of Geostiba Thomson 1858 and Tropimenelytron Pace 1983 (Coleoptera: Staphylinidae, Aleocharinae). Linzer Biol Beitr 33(1):137–185 Benick G (1973) Die von Prof. Dr. Z. Kaszab in der Mongolei gesammelten Arten der Subfamilie Athetae (Col. Staphyl.). Nouv Rev Entomol 3(4):211–217 Benick G, Lohse GA (1974) Tribus 14 (Callicerini) [pp 72–220]. In: Freude H, Harde KW, Lohse A (eds) Die Käfer Mitteleuropas. Band 5: Staphylinidae II (Hypocyphtinae und Aleocharinae) Pselaphidae. Goecke & Evers. Krefeld, 381 pp Bernhauer M (1902) Die Staphyliniden der paläarktischen Fauna. 1. Tribus: Aleocharini (part II). Verh K-Kg Zool-Bot Ges Wien 52:87–284 Bernhauer M (1906) Neue Aleocharinen aus Nordamerika. (II. Teil). Dtsch Entomol Z 1906(2):337–348 Bernhauer M (1907) Neue Aleocharini aus Nordamerika. (Col.) (3. Stück.). Dtsch Entomol Z 1907(4):381–405 Bernhauer M (1909) Neue Aleocharini aus Nordamerika. (Col.) (4. Stück.). Dtsch Entomol Z 1909(4):515–528 Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013) Checklist of beetles (Coleoptera) of Canada and Alaska, 2nd edn. Pensoft, Series Faunistica No. 109. Sofia-Moscow, 402 pp Brundin L (1948) Microdota-Studien. (Col. Staphylinidae). Entomol Tidsskr 69(1–2):8–66 Brunke A, Klimaszewski J, Dorval J-A, Bourdon C, Paiero SM, Marshall SA (2012) New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:119–206 Casey TL (1885) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1(4):283–336

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Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7[1894]:281–606 [see reference in Chapter 7 for explanation of date] Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Transactions of the Academy of Sciences of St. Louis 16(6):125–434 Casey TL (1910) New species of the staphylinid tribe Myrmedoniini [pp 1–183]. In: Memoirs on the Coleoptera. I. New Era Printing, Lancaster, PA, 205 pp Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp. 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing, Lancaster, PA, 259 pp Clark ME (1949) An annotated list of the Coleoptera taken at or near Terrace, British Columbia. Part 2. Proc Entomol Soc Br Columbia 45([1948]):21–24 Criddle N (1922) The entomological record, 1921. Annu Rep Entomol Soc Ont 52(1921):57–70 Crotch GR (1870) The genera of Coleoptera studied chronologically (1802–1821). Trans Entomol Soc London 213–241 Dollin PE, Majka CG, Duinker PN (2008) Saproxylic beetle (Coleoptera) communities and forest management practices in coniferous stands in southwestern Nova Scotia, Canada. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 2:291–336 Elven H, Bachmann L, Gusarov VI (2010) Phylogeny of the tribe Athetini (Coleoptera: Staphylinidae) inferred from mitochondrial and nuclear sequence data. Mol Phylogenet Evol 57(1):84–100 Elven H, Bachmann L, Gusarov VI (2012) Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta). Zool Scr 41(6):617–636 Eppelsheim E (1886) Neue Staphylinen vom Amur. Dtsch Entomol Z 30(1):33–46 Erichson WF (1837) Die Käfer der Mark Brandenburg. Erster Band, Erste Abtheilung. F. H. Morin, Berlin, viii + 384 pp Erichson WF (1839) Genera et species Staphylinorum insectorum coleopterorum familiae. Erster Band. F. H. Morin Berlin, VIII + 400 pp Fauvel A (1889) Liste des coléoptères communs à l’Europe et a l’Amérique du Nord. D’après le catalogue de M. J. Hamilton. Avec remarques et additions. Rev Entomol 8(3):92–174 Fenyes A (1918) Coleoptera. Family Staphylinidae. Subfamily Aleocharinae [pp 1–110]. In: Wytsman P (ed) Genera Insectorum, vol XXVII, Fascicule 173A. M. Nijhoff, Den Haag, 453 pp + 7 pls Fenyes A (1920) Coleoptera. Fam. Staphylinidae. Subfam. Aleocharinae [pp 111–414]. In: Wytsman P (ed) Genera Insectorum, vol XXVII, Fascicule 173B. L. Desmet-Verteneuil, Bruxelles, 453 pp + 7 pls Frank JH (1981) Atheta coriaria (Kraatz) (Aleocharinae) and Sunius confluentus (Say) (Paederinae) in Florida (Coleoptera, Staphylinidae). Coleopt Bull 34[1980] (4):338

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Gouix N, Klimaszewski J (2007) Catalogue of aleocharine rove beetles of Canada and Alaska (Coleoptera, Staphylinidae, Aleocharinae). Pensoft, Sofia-Moscow, 165 pp Gozis M (1886) Recherches de l’espèce typique de quelque anciens genres: Rectifications synonymiques et notes diverse. Herbin, Montluçon, 36 pp Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Gravenhorst JLC (1806) Monographia Coleopterorum Micropterorum. Henricus Dieterich, Göttingen, xvi + 236 + [12] pp Gusarov VI (2002a) A revision of Nearctic species of the genus Earota Mulsant & Rey, 1873 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 92:1–16 Gusarov VI (2002b) A revision of Nearctic species of the genus Geostiba Thomson, 1858 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 81:1–88 Gusarov VI (2002c) A revision of Nearctic species of the genus Tropimenelytron Pace, 1983 (Coleoptera: Staphylinidae: Aleocharinae), a new genus for North America. Zootaxa 114:1–24 Gusarov VI (2003a) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353:1–134 Gusarov VI (2003b) A catalogue of the athetine species of America north of Mexico (Coleoptera: Staphylinidae: Aleocharinae: Athetini). (I.S.: Internet Site). https:// web.archive.org/web/20100613213828/http://www. nhm.ku.edu/ksem/peet/catalogs/cataweb.htm Gusarov VI (2003c) A revision of the genus Seeversiella Ashe, 1986 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 142:1–102 Gusarov VI (2003d) A revision of Nearctic species of the genera Adota Casey, 1910 and Psammostiba Yosii & Sawada, 1976 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 185:1–35 Gusarov VI (2004) A revision of the genus Lypoglossa Fenyes, 1918 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 747:1–36 Gyllenhal L (1810) Insecta Svecica. Classis I: Coleoptera sive Eleuterata. Tomi I pars II. F.J. Leverentz, Scaris. xix + [1] + 660 pp Hamilton J (1889) Catalogue of the Coleoptera common to North America, northern Asia and Europe, with the distribution and bibliography. Trans Am Entomol Soc 16(3–5):88–162 Hatch MH (1957) The beetles of the Pacific Northwest. Part II: Staphyliniformia. University of Washington Publications in Biology, 16, IX + 384 pp Hoebeke ER (1992) Taxonomy and distribution of the athetine genus Lypoglossa Fenyes (Coleoptera: Staphylinidae: Aleocharinae) in North America, with description of a new species. J NY Entomol Soc 100 (2):381–398

567 Klimaszewski J, Peck SB (1986) A review of the cavernicolous Staphylinidae (Coleoptera) of eastern North America: Part I. Aleocharinae. Quaestiones Entomol 22(2):51–113 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belge Entomol 40:3–126 Klimaszewski J, Sweeney J, Price J, Pelletier G (2005) Rove beetles (Coleoptera: Staphylinidae) in red spruce stands, eastern Canada: diversity, abundance, and descriptions of new species. Can Entomol 137(1):1–48 Klimaszewski J, Assing V, Majka CG, Pelletier G, Webster RP, Langor D (2007a) Records of adventive aleocharine beetles (Coleoptera: Staphylinidae: Aleocharinae) found in Canada. Can Entomol 139 (1):54–79 Klimaszewski J, Langor D, Savard K, Pelletier G, Chandler DS, Sweeney J (2007b) Rove beetles (Coleoptera: Staphylinidae) in yellow birch-dominated stands of southeastern Quebec, Canada: diversity, abundance, and description of a new species. Can Entomol 139 (6):793–833 Klimaszewski J, Godin B, Pelletier G, Savard K (2008a) Six new species and records of aleocharine beetles from the Yukon and Alaska (Coleoptera: Staphylinidae: Aleocharinae). Can Entomol 140 (3):265–291 Klimaszewski J, Langor DW, Work TT, Hammond JHE, Savard K (2008b) Smaller and more numerous harvesting gaps emulate natural forest disturbances: a biodiversity test case using rove beetles (Coleoptera, Staphylinidae). Divers Distrib 14(6):969–982 Klimaszewski J, Lynch D, Majka CG, Renkema J, Savard K, Hlaváč P (2009a) Pella glooscapi, a new rove beetle, and new records of aleocharines from Nova Scotia, Canada (Coleoptera, Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:35–44 Klimaszewski J, Webster RP, Savard K (2009b) First record of the genus Schistoglossa Kraatz from Canada with descriptions of seven new species (Coleoptera, Staphylinidae, Aleocharinae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:45–79 Klimaszewski J, Langor D, Majka CG, Bouchard P, Bousquet Y, LeSage L, Smetana A, Sylvestre P, Pelletier G, Davies A, DesRochers P, Goulet H, Webster R, Sweeney J (2010) Review of adventive species of Coleoptera (Insecta) recorded from eastern Canada. Pensoft Series Faunistica No. 94. Pensoft SofiaMoscow, 272 pp Klimaszewski J, Langor D, Pelletier G, Bourdon C, Perdereau L (2011) Aleocharine beetles (Coleoptera, Staphylinidae) of the province of Newfoundland and Labrador, Canada. Pensoft Series Faunistica No. 98. Pensoft, Sofia-Moscow, 313 pp

568 Klimaszewski J, Godin B, Bourdon C (2012) Further contributions to the aleocharine fauna of the Yukon Territory, Canada (Coleoptera, Staphylinidae). In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:207–237 Klimaszewski J, Brunke A, Assing V, Langor DW, Newton AF, Bourdon C, Pelletier G, Webster RP, Herman L, Perdereau L, Davies A, Smetana A, Chandler DS, Majka C, Scudder GGE (2013a) Synopsis of adventive species of Coleoptera (Insecta) recorded from Canada. Part 2: Staphylinidae. Pensoft Series Faunistica No. 104. Pensoft, Sofia-Moscow, 360 pp Klimaszewski J, Webster RP, Langor DW, Bourdon C, Jacobs J (2013b) Review of Canadian species of the genus Dinaraea Thomson, with descriptions of six new species (Coleoptera, Staphylinidae, Aleocharinae, Athetini). ZooKeys 327:65–101 Klimaszewski J, Godin B, Langor D, Bourdon C, Lee S-I, Horwood D (2015a) New distribution records for Canadian Aleocharinae (Coleoptera, Staphylinidae), and new synonymies for Trichiusa. ZooKeys 498:51–91 Klimaszewski J, Webster RP, Bourdon C, Pelletier G, Godin B, Langor DW (2015b) Review of Canadian species of the genus Mocyta Mulsant & Rey (Coleoptera, Staphylinidae, Aleocharinae), with the description of a new species and a new synonymy. ZooKeys 487:111–139 Klimaszewski J, Webster RP, Sikes D, Bourdon C, Labrecque M (2015c) A review of Canadian and Alaskan species of the genera Clusiota Casey and Atheta Thomson, subgenus Microdota Mulsant & Rey (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 524:103–136 Klimaszewski J, Webster RP, Langor D, Sikes D, Bourdon C, Godin B, Ernst C (2016a) A review of Canadian and Alaskan species of the genus Liogluta Thomson, and descriptions of three new species (Coleoptera, Staphylinidae, Aleocharinae). In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:217–256 Klimaszewski J, Langor DW, Hammond HEJ, Bourdon C (2016b) A new species of Anomognathus and new Canadian and provincial records of aleocharine rove beetles from Alberta, Canada (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 581:141–164 Klimaszewski J, Langor DW, Bourdon C, Gilbert A, Labrecque M (2016c) Two new species and new provincial records of aleocharine rove beetles from Newfoundland and Labrador, Canada (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 593:49–89 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016d) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112

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Klimaszewski J, Webster RP, Davies A, Bourdon C (2018a) Description of Hydrosmectomorpha Klimaszewski and Webster, a new subgenus of Atheta C.G. Thomson, with three new Canadian species (Coleoptera: Staphylinidae: Aleocharinae). Insecta Mundi 0648:1–12 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018b) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham. 902 pp Klimaszewski J, Sikes DS, Brunke A, Bourdon C (2019) Species review of the genus Boreophilia Benick from North America (Coleoptera: Staphylinidae: Aleocharinae: Athetini): systematics, habitat and distribution. ZooKeys 848:57–102 Kraatz G (1856) Staphylinii. Naturgeschichte der Insecten Deutschlands. Erste Abtheilung Coleoptera. Zweiter Band. Erste, zweite Lieferung. Bogen 1–24. Nicolai, Berlin, viii + 376 pp Kraatz G (1857). Genera Aleocharinorum illustrata. Linnaea Entomologica 11:1–43, 2 pls LeConte JL (1863) New species of North American Coleoptera. Smithson Misc Collections 6:1–56 Lee S-G, Ahn K-J (2015) A taxonomic review of Korean species of the Atheta Thomson subgenus Microdota Mulsant & Rey, with descriptions of two new species (Coleoptera, Staphylinidae, Aleocharinae). ZooKeys 502:61–97 Lee S-G, Ahn K-J (2017) A taxonomic review of the Korean species of the genus Datomicra Mulsant and Rey of the genus Atheta Thomson, with description of a new species (Coleoptera, Staphylinidae, Aleocharinae). Zootaxa 4262:508–522 Lohse GA (1971) Über gattungsfremde Arten und Artemkreise innerhalb der “Grossgattung” Atheta Thomson. Verh Ver Nat Heimatforschung Hanburg 38:67–83 Lohse GA, Smetana A (1985) Revision of the types of species of Oxypodini and Athetini (sensu Seevers) described by Mannerheim and Mäklin from North America (Coleoptera: Staphylinidae). Coleopt Bull 39 (3):281–300 Lohse GA, Smetana A (1988) Four new species of Geostiba Thomson from the Appalachian Mountains of North Carolina, with a key to North American species and synonymic notes (Coleoptera: Staphylinidae: Aleocharinae). Coleopt Bull 42:269–278 Lohse GA, Klimaszewski J, Smetana A (1990) Revision of arctic Aleocharinae of North America (Coleoptera: Staphylinidae). Coleopt Bull 44(2):121–202 Majka CG, Klimaszewski J (2008a) Introduced Staphylinidae (Coleoptera) in the Maritime Provinces of Canada. Can Entomol 140(1):48–72 Majka CG, Klimaszewski J (2008b) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity,

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biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:85–114 Majka CG, Klimaszewski J (2008c) Adventive Staphylinidae (Coleoptera) of the Maritime Provinces of Canada: further contributions. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:151–174 Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–39 Majka CG, Sikes D (2009) Thomas L. Casey and Rhode Island’s precinctive beetles: taxonomic lessons and the utility of distributional checklists. ZooKeys 22:267–283 Majka CG, Klimaszewski J, Lauff RF (2006) New Coleoptera records from owl nests in Nova Scotia, Canada. Zootaxa 1194:33–47 Majka CG, Klimaszewski J, Lauff RF (2008) The coastal rove beetles (Coleoptera, Staphylinidae) of Atlantic Canada: a survey and new records. ZooKeys 2:115–150 Mäklin FW (1852) [New species and notes]. In: Mannerheim CG: Zweiter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Imp Nat Moscou 25(2):283–372 Mäklin FW (1853) [New species and notes]. In: Mannerheim CG: Dritter Nachtrag zur Kaefer-Fauna der Nord-Amerikanischen Laender des Russischen Reiches. Bull Soc Imp Nat Moscou 26(3):95–273 Mannerheim CG (1830) Précis d’un nouvel arrangement de la famille des brachélytres de l’ordre des insectes coléoptères. St. Petersburg, 87 pp [separate, re-issued in Mémoires Présentées à l’Académie Impériale des Sciences de St.-Pétersbourg 1:415–501, in 1831] Mannerheim CG (1843) Beitrag zur Kaefer-fauna der Aleutischen Inseln, der Insel Sitkha und Neu-Californiens. Bull Soc Imp Nat Moscou 16 (2):175–314 Märkel F, Kiessenwetter EAH (1848) Bericht über eine entomologische Excursion in die Käarthner Alpen im Jahre 1847. Entomol Ztg (Stettin) 9:321–329 Maruyama M, Klimaszewski J (2004) A new genus and species of the myrmecophilous Athetini, Paragoniusa myrmicae (Coleoptera: Staphylinidae: Aleocharinae) from Canada. Entomol Rev Jpn 59(2):241–248 Maruyama M, Klimaszewski J (2006) Notes on myrmecophilous aleocharines (Insecta, Coleoptera, Staphylinidae) from Canada, with a description of a new species of Myrmoecia. Bull Natl Sci Mus, Tokyo (A) 32(3):125–131 McLean JA, Klimaszewski J, Li A, Savard K (2009) Survey of rove beetles (Coleoptera, Staphylinidae) from Stanley Park, Vancouver, British Columbia, Canada, with new records and description of a new species. Part 1. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:5–17

569 Michaud J-P, Majka CG, Privé J-P, Moreau G (2010) Natural and anthropogenic changes in the insect fauna associated with carcasses in the North American Maritime lowlands. Forensic Sci Int 202(1–3):64–70 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Moore I, Legner EF (1976) Intertidal rove beetles (Coleoptera: Staphylinidae). pp 521–551. In: Cheng (ed) Marine insects. Amsterdam Mulsant E, Rey C (1852) Description de quelques coléoptères nouveaux ou peu connus de la tribu des brachélytres. Opusc Entomol 1:15–46, 11 pl Mulsant E, Rey C (1873a) Description de divers coléoptères brévipennes nouveaux ou peu connus. Opusc Entomol 15:147–189 Mulsant E, Rey C (1873b) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens (Suite) [Tableau... des Aléochariens + Septième Branche, Myrmédoniaires, part 1]. Deyrolle, Paris, [2] + 695 pp, pls 1–5 Muona J (1979) Staphylinidae. In: Silfverberg H, Enumeratio Coleopterorum Fennoscandiae et Daniae. Helsingfors Entomologiska Bytesförening, Helsinki, pp 14–28 Muona J (1983) Two new Palearctic Atheta species (Coleoptera, Staphylinidae). Ann Entomol Fenn 49 (2):57–58 Muona J (1984) Review of Palearctic Aleocharinae also occurring in North America (Coleoptera: Staphylinidae). Entomol Scand 15(2):227–231 Muona J (1990) The Fennoscandian and Danish species of the genus Amischa Thomson (Coleoptera, Staphylinidae). Entomol Tidsskr 111:17–24 Notman H (1922) Some new genera and species of Coleoptera collected at Westfield, Chautauqua Co, NY. J NY Entomol Soc 29 [1921](3–4):145–160 Pace R (1983) Il genere Tropimenelytron Scheerpeltz (Coleoptera, Staphylinidae) (XXXIX contributo alla conoscenza delle Aleocharinae). Nouv Rev Entomol 13(2):185–190 Pace R (2006) Aleocharinae del Nepal al Naturkundemuseum di Erfurt (Insecta: Coleoptera: Staphylinidae) [pp 343–408] In: Hartmann M, Weipert J (eds) Biodiversität und Naturausstattung im Himalaya II. Erfurt, Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V., Erfurt Palm T (1970) Svensk Insektfauna. 9. Skalbaggar. Coleoptera. Hafte 6. Kortvingar: Fam. Staphylinidae underfam. Aleocharinae (Atheta), (Nr. 52). Entomologiske Föreningen i Stockholm, Stockholm: 296 pp, 21 pls Pochubay EA, Grieshop MJ (2012) Intraguild predation of Neoseiulus cucumeris by Stratiolaelaps miles and Atheta coriaria in greenhouse open rearing systems. Biol Control 63(2):195–200 Ratnasingham S, Hebert PDN (2013) A DNA-based registry for all animal species: The Bar–code Index

570 Number (BIN) System. PLoS One 8:e66213. https:// doi.org/10.1371/journal.pone.0066213 Renkema JM, Telfer Z, Gariepy T, Hallett R (2015) Dalotia coriaria as predator of Drosophila suzukii: functional responses, reduced fruit infestation and molecular diagnostics. Biol Control 89:1–10 Schülke M, Smetana A (2015) Family Staphylinidae Latreille, 1802. In: Löbl I, Löbl D (eds) Catalogue of Palearctic Coleoptera, vol 2. Revised and updated edition. Hydrophiloidea – Staphylinoidea. Brill, Leiden, pp 304–1134 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana Zool 71:vi + 289 pp Smetana A (2004) Family Staphylinidae Latreille, 1802 New nomenclatorial and taxonomic acts, and comments [pp 29–35], subfamilies OmaliinaeDasycerinae [pp 237–272], subfamilies Phloeocharinae-Apateticinae [pp 329–495], subfamilies Piestinae-Staphylininae [pp 504–698]. In: Löbl I, Smetana A (eds) Catalogue of Palearctic Coleoptera, vol 2. Hydrophiloidea – Histeroidea – Staphylinoidea. Apollo Books, Stenstrup, 942 pp Stefani FOP, Klimaszewski J, Morency M-J, Bourdon C, Labrie P, Blais M, Venier L, Séguin A (2016) Fungal community composition in the gut of rove beetles (Coleoptera: Staphylinidae) from the Canadian boreal forest reveals possible endosymbiotic interactions for dietary needs. Fungal Ecol 23:164–171 Strand A, Vik A (1964) Die Genitalorgane der nordischen Arten der Gattung Atheta Thoms. (Col., Staphylinidae). Norsk Entomol Tidsskr 12 (5–8):327–335, pls 1–21 Thomson CG (1858) Försök till uppställning af Sveriges Staphyliner. Öfversigt af Kongliga VetenskapsAkademiens Förhandlingar 15(1):27–40 Thomson CG (1859) Skandinaviens Coleoptera, synoptiskt bearbetade. I. Tom Berlingska Boktryckeriet, Lund, [3] + 290 pp

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Thomson CG (1867) Skandinaviens Coleoptera, synoptiskt bearbetade. Supplementum. Tom IX. Lundbergska Boktryckeriet, Lund, 407 pp Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:171–248 Webster RP, Klimaszewski J, Sweeney J D, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and Ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186: 83–118 Webster RP, Klimaszewski J, Bourdon C Sweeney JD, Hughes CC, Labrecque M (2016) Further contribution to the Aleocharinae (Coleoptera, Staphilinidae) fauna of New Brunswick and Canada including description of 27 new species. In: Webster RP, Bouchard P, Klimaszewski J (eds) The Coleoptera of New Brunswick and Canada: providing baseline biodiversity and natural history data. ZooKeys [Special Issue] 573:85–216 Wollaston TV (1855) Descriptions of two Coleopterous insects new to British fauna. Zoologists 13:205–207 Work TT, Klimaszewski J, Thiffault E, Bourdon C, Paré D, Bousquet Y, Venier L, Titus B (2013) Initial responses of rove and ground beetles (Coleoptera, Staphylinidae, Carabidae) to removal of logging residues following clearcut harvesting in the boreal forest of Quebec, Canada. ZooKeys 258:31–52 Yosii R, Savada K (1976) Studies on the genus Atheta Thomson and its allies (Coleoptera, Staphylinidae). II: Diagnostic characters of genera and subgenera with description of representative species. Contr Biol Lab Kyoto Univ 25:11–140

Tribe Falagriini Mulsant and Rey, 1873

21

Tribe Falagriini Mulsant and Rey, 1873 Key to Genera of Falagriini (Figs. 21.258, 21.260–21.262) (Hoebeke 1985; Ahn and Ashe 1995; Ashe 2001; Klimaszewski et al. 2018) Diagnosis. Body narrow and elongate, length 2.0–4.0 mm; head with well defined narrow neck, which is less than half width of head; ligula bifid, laciniae spinose at apex and ciliate along inner margin, galeae finely and densely ciliate on membraneous apex; pronotum widest subapically, narrowed basally to about threefourths of width of pronotum, disc with a slightly impressed to very deep median longitudinal sulcus; legs long and slender, tarsal formula 4-5-5 (Canadian species); mesocoxal cavities moderately separated, proventrite elongate behind level of procoxal insertions, often extending laterally to pronotal hypomera; mesospiracular plates or peritremes present around spiracles on mesoventrite, peritreme size ranging from very small and narrowly surrounding spiracles, to very large and contiguous along midline; abdomen broad and slightly constricted at base, tergite IV with distinctive medial gland opening, III–V each transversely and deeply impressed at base, impressions smooth or coarsely sculptured; paramere of aedeagus with velum of paramerite and condylite clearly separated into two lobes.

1. Elytra at suture shorter than pronotum along midline, and about as wide as pronotum (Fig. 21.258a)........................... Bryobiota Casey – Elytra at suture longer than pronotum along midline, and distinctly wider than pronotum .....2 2. Scutellum bicarinate medially (Fig. 21.262a) ............................... Falagria Leach – Scutellum not bicarinate medially.................3 3. Abdominal tergite VII with apical margin densely fimbriate (Fig. 21.261a); elytron at base with small, distinct impression near humeral angle (Fig. 21.261a); scutellum without median longitudinal carina.....................Cordalia Jacobs – Abdominal tergite VII with apical margin simple, not densely fimbriate (Fig. 22.263a); elytron at base without distinct impression near humeral angle (Fig. 22.263a); scutellum with complete or incomplete median longitudinal carina............................... ................................... Myrmecocephalus MacLeay

Genus Bryobiota Casey, 1893 [Revised Concept] (Figs. 21.258a–d and 21.259a–h) (Casey 1893; Ahn and Ashe 1995; Ashe 2001)

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_21

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Diagnosis [based on female lectotype of Bryobiota bicolor (Casey), Fig. 21.258a–d]. Body length 2.2 mm, slender, narrow, moderately flattened, and parallelsided; colour yellow-reddish brown, with dark brown abdomen; body sculpture of head, pronotum, elytra, mostly hexagonal; integument moderately glossy, on head, pronotum, and elytra faintly granulose (more so on elytra) as a result of a fine-meshed raised reticulation, more or less uniformly pubescent with moderately long microsetae; head with neck but often not visible because contracted under pronotum, postocular region long subparallel with slightly angular, rounded genae; eyes small, each about three times shorter than length of postocular region in dorsal view; three basal antennomeres elongate, III slightly shorter than II, III–IV subquadrate, and V–X progressively from slightly to strongly transverse; maxillary palpus with four palpomeres, penultimate palpomere wide, and last one small and needle-shaped; labial palpus with three palpomeres; ligula elongate and bifid apically; pronotum slightly longitudinally impressed medially, truncate basally and rounded apically, broadest in apical third, pubescence on median line of disc directed apically and laterad on sides forming arcuate lines, hypomera delimited from disc by marginal carina; elytra shorter than pronotum, hind wings absent; abdomen slightly swollen, sides arcuate, broadest at middle length, four basal tergites with deep impressions, apical margins of tergites straight medially and slightly arcuate laterally; spermatheca S-shaped, capsule tubular with wide apical invagination, stem about as broad as capsule and strongly sinuate; species restricted to the intertidal zone of sandy beaches. Comments. The genus Bryobiota is here redefined based on the morphology of the lectotype of B. bicolor Casey, the type species of the genus. 221. Bryobiota bicolor (Casey) (Figs. 21.258a–d and 21.259a–h) Phytosus bicolor Casey, 1885. Casey 1893 [as Bryobita], Sawada 1972, Ahn and Ashe 1995.

21 Tribe Falagriini Mulsant and Rey, 1873

Lectotype (female): Cal.[ifornia]; Bryobiota bicolor Csy; TYPE USNM 38774; Casey bequest 1925; Lectotype Bryobiota bicolor (Casey), Desig. K. J. Ahn and J. S. Ashe 1992. Examined (Fig. 21.258a–d). Myrmecopora (Xenusa Mulsant and Rey) californica Scheerpeltz, 1964. Synonymized by Ahn and Ashe (1995). Not verified. Diagnosis [based on lectotype]. Body length 2.2 mm, slender, colour reddish-yellow with dark brown abdomen and slightly darker antennae than forebody (Fig. 21.258a); forebody finely punctate, moderately glossy; head about 1.1 times as wide as long, convex, genae rounded and slightly angular; antennomere III slightly shorter than II, VI–X distinctly transverse; pronotum subquadrate, rounded apically and straight basally, broadest in apical half, convex; elytra short, at suture distinctly shorter than length of pronotum along midline; abdomen slightly swollen, arcuate laterally, with several distinct macrosetae present on each segment, the anterior margin of abdominal terga III–V straight. Association of male with this species, presented by Ahn and Ashe (1995), needs confirmation. MALE. Median lobe of aedeagus was illustrated by Ahn and Ashe (1995) (Fig. 21.258e). FEMALE. Tergite VIII truncate apically (Fig. 21.258b); sternite VIII truncate apically and slightly produced medially, apical margin slightly arcuate medially (Fig. 21.258c); spermatheca S-shaped, capsule broadly tubular with wide and shallow invagination, stem sinuate, broadly tubular, slightly swollen posteriorly (Fig. 21.258d). MALE. Median lobe of aedeagus (Fig. 21.258e), after Ahn and Ashe (1995). We did not find males in BC material and were not able to confirm male association for this species. DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CA. New locality data: British Columbia, Kye Bay, 6. V.32, Gregson; sea-weed; det. 4906 F.E. Blaisdall, Bryobiota bicolor (Casey) 4906; Bryobiota bicolor (Casey) det. K-J. Ahn 1994 (UBC) 2 females [same as female lectotype of B. bicolor].

21

Tribe Falagriini Mulsant and Rey, 1873

COLLECTION AND HABITAT DATA (Ahn and Ashe 1995). Habitat: under decaying seaweed on sandy beaches, rock crevices in splash zones. Collecting period: I, II, VIII. Collecting method: sifting seaweed. Comments. Bryobiota bicolor (Casey 1885) (Fig. 21.258a–e) was recorded in Canada from Key Bay in BC as a different species from B. giulianii (Moore 1978) [originally Salinamexus Moore] (Fig. 21.259a–h), recorded from California (Ahn and Ashe 1995). Ahn and Ashe (1995) considered the two species as belonging to the genus Bryobiota. The median lobe of the aedeagus, and the wrongly affiliated spermathecae for the two species were illustrated by Ahn and Ashe (1995). These authors designated a lectotype for B. bicolor but did not dissect it. We have dissected the lectotype of Bryobiota bicolor (Casey), and discovered that it is a female (Fig. 21.258a–d), with the same shape of the spermatheca as illustrated for B. giulianii by these authors. When studying additional specimens, we have found that the illustrations of the spemathecae provided by Ahn and Ashe (1995) were wrongly assigned to species, and should be switched. They are illustrated and assigned to the proper species in this book. “Salinamexus” giulianii Moore was incorrectly transferred to Bryobiota and is here considered as belonging to a new and distinct genus. We propose a new name, Ashella Klimaszewski new genus, in honor of James S. Ashe, with Ashella giulianii (Moore) as the generic type (CAS) (Fig. 21.259a–h). The genus may be distinguished from Bryobiota and other aleocharine genera by the very narrow, linear and flattened body; forebody with coarse punctation; head angular posteriorly with long genae, gena almost three times longer than diameter of an eye in dorsal view; eyes moderately sized; antennomere III subquadrate; first two visible abdominal tergites with V-shaped basal emargination; and the tubus of the median lobe of the aedeagus narrow and with apical hook in lateral view. All these characters are illustrated in Fig. 21.259a–h.

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We suspect that this genus is related to species with subquadrate, not elongate, antennomere III, of the Tribe Liparocephalini.

Genus Cordalia Jacobs, 1925 (Fig. 21.260a–h) (Hoebeke 1985) Diagnosis. Body of distinctive shape (Fig. 21.260a); head about as broad as pronotum, arcuately truncate at base, posterior angles broadly rounded, vertex impressed at middle in males; eyes small, shorter than genae in dorsal view; mandibles moderate-sized, slightly curved apically, right mandible with small obtuse tooth at middle of inner margin, finely crenulate between tooth and apex, left mandible simple, untoothed, maxillae with galeae shorter than laciniae, maxillary palpi with palpomere III much longer and broader than II, and IV less than half length of III; ligula bifid to middle, lobes rounded at apex, labial palpi with palpomere II much shorter and narrower than I, III longer than II, feebly conical and truncate at apex; antenna moderately long, antennomeres I–II subequal in length, IV slightly transverse, V–X gradually more transverse towards apex; pronotum cordiform, highly convex, much narrower than elytra at greatest width, sides strongly converging basad, disc finely or obsoletely sulcate along median line, sulcus without antebasal impression; hypomera not delimited from disc; mesospiracular peritremes within prosternal membrane large, quadrate, contiguous along median line; process of mesoventrite very short, not quite extending to anterior fourth of mesocoxae, broad and truncate apically; process of metaventrite longer than that of mesoventrite, flat and truncate apically, isthmus short; mesocoxae broadly separated, acetabula margined behind by fine ridge; scutellum flat, moderately punctate and pubescent; elytra at suture longer than pronotum, each elytron with distinct depression at base near humerus; abdomen broad, at base slightly narrower than elytra, tergites III–V transversely impressed at base, impressions coarsely punctate.

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222. Cordalia obscura (Gravenhorst) { (Fig. 21.260a–h) Aleochara obscura Gravenhorst, 1802. Muona 1984, Hoebeke 1985, Humble et al. 2000, Majka and Klimaszewski 2008, Klimaszewski et al. 2010, 2018, Webster et al. 2012.

21 Tribe Falagriini Mulsant and Rey, 1873

decaying, moldy corncobs and cornhusks (common) near mixed forest and some from QC and NH from compost in mixed forests. Collecting period: V–X. Collecting method: sifting organic debris, bacon-baited pitfall trap.

Genus Falagria Leach, 1819 Diagnosis. Body ant-shaped, stout, convex, length 2.0–2.8 mm; colour dark rufo-piceous, with elytra usually lighter (Fig. 21.260a); head small and arcuate-truncate at base, eyes shorter than genae in dorsal view; antennae extended to middle of elytra, antennomeres IV–X quadrate to slightly transverse; pronotum cordiform, highly convex dorsally, sides strongly converging to base, disc sulcate along median line; elytra short, wider than rest of the body; abdomen broad at base and arcuate laterally. MALE. Tergite VIII with apical margin broadly emarginate at middle and densely fimbriate (Fig. 21.260d); sternite VIII truncate apically (Fig. 21.260e); median lobe of aedeagus with suboval bulbus, and short and straight tubus in lateral view, its apex slightly pointing ventrally (Fig. 21.260b); in dorsal view bulbus large, oval, tubus short and narrowly triangular at apex (Fig. 21.260c). FEMALE. Tergite VIII with apical margin broadly emarginate at middle and densely fimbriate (Fig. 21.260f); sternite VIII truncate apically (Fig. 21.260g); spermatheca with capsule clubshaped, flattened apically and bearing long indentation, stem short, broad and swollen posteriorly (Fig. 21.260h). DISTRIBUTION. Origin: Palearctic, adventive in North America. CANADA: BC, NB, NS, ON, QC. USA: CA, CO, CT, IL, MA, MD, MI, NH, NJ, NY, OH, PA, WA. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: forested and anthropogenic habitats; in North America, reported from various kinds of organic debris such as grass clippings, decaying vegetation and compost, rotting bracket fungi, garden soil, and from bacon bait trap. NB specimens from

(Fig. 21.261a–h) (Hoebeke 1985) Diagnosis. Head moderate in size, about as wide as pronotum at maximum width, arcuate-truncate at base, hind angles rounded, neck about one-fourth to one-sixth as wide as head across eyes; eyes small, slightly prominent; mandibles robust, right mandible with small tooth at middle of inner margin, left mandible simple, galeae pubescent on membranous apex, laciniae curved towards hooked apex, with long spines, maxillary palpi with palpomere III longer than II and incrassate towards apex, IV narrow, subuliform, about a one-third length of III, ligula small, narrow, distinctly split to about middle, lobes rounded apically, labial palpi with palpomere II dilated towards truncate apex; antennae moderately long, widened towards apex, antennomeres I–III elongate, III as long as or shorter than II, IV– X transverse; pronotum cordiform, broadest at apical one-third, sides strongly converging basad, disc with deep sulcus along median line, with fine, transverse and impressed line before base; prosternum moderately short before procoxae, posterior margin with obtuse-angular process; hypomera delimited from pronotal disc by marginal ridge; mesospiracular peritremes large, quadrate, and contiguous along median line; process of mesoventrite extending to about half length of mesocoxae, obtusely rounded or narrowly truncate apically; process of metaventrite obtuse, separated from that of mesoventrite by short isthmus; mesocoxae moderately separated; scutellum strongly bicarinate; elytra convex, slightly bisinuate basally at outer angles, integument without microsculpture; abdomen almost as broad as elytra at base, dilated at middle, first three visible tergites transversely impressed basally, impressions coarsely punctate,

21

Tribe Falagriini Mulsant and Rey, 1873

apical margin of tergite VIII with ctenidium of minute spinules. 223. Falagria dissecta Erichson (Fig. 21.261a–h) Falagria dissecta Erichson, 1839. Melsheimer 1844 [as two synonyms], Casey 1906, 1911 [as six synonyms], Fenyes 1912, Hoebeke 1985, Klimaszewski et al. 2018. Diagnosis. Body length 1.9–2.6 mm; colour pale piceo-testaceous to black or piceous-black, with mouthparts, basal antennomeres, elytra in some, and legs paler (Fig. 21.261a). MALE. Tergite VIII with apical margin arcuate, with comb of very minute denticles (Fig. 21.261d); sternite VIII truncate apically (Fig. 21.261e); median lobe of aedeagus with tubus moderately long, narrow subapically and slightly pointed ventrally at apex in lateral view (Fig. 21.261b), in dorsal view bulbus oval, tubus moderately long and broad, triangular apically (Fig. 21.261c). FEMALE. Tergite VIII with apical margin arcuate, with comb of minute denticles (Fig. 21.261f); sternite VIII broadly arcuate apically, truncate medially (Fig. 21.261g); spermatheca curved, with capsule large, subspherical, stem short and broad, not swollen posteriorly (Fig. 21.261h). Externally, this species appears very similar to F. sulcata, from which it differs in having a deep pronotal sulcus interrupted near the middle by a small depression, the abdomen without long, erect setae ventrally on the margins and apically, and in the shapes of the median lobe of the aedeagus and the spermatheca. DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NS, ON, QC, SK. USA: AL, AZ, CA, CO, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, ME, MA, MD, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NY, OH, OR, PA, RI, SC, SD, TX, UT, VA, VT, WA, WI, WV, WY. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: occurs in decaying organic debris in various habitats. Adults from decaying vegetation, compost,

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animal carcasses, under animal dung, fungi, debris on a beach, nests or nest lining of various rodents such as beaver, marmot, and the voles Microtus pennsylvanicus (Ord.) and M. ochrogaster (Wagner). Specimens also captured in malt, black light, and pitfall traps; NB specimens from compost (common), in sea wrack on sandy sea beach, on bare soil among grass on residential lawn, and from debris in entrance of fox den in a pasture. Collecting period: IV–XI. Collecting method: sifting organic litter and debris, malt trap, blacklight, and pitfall traps.

Genus Myrmecocephalus MacLeay, 1871 (Fig. 21.262a–h) (Hoebeke 1985) Diagnosis. Head rounded or arcuate-truncate at base, at least as wide as maximum width of pronotum, neck very narrow; eyes large and prominent; mandibles moderately sized, curved towards acute apex, right mandible with obtuse tooth at middle of inner margin, left mandible simple, galeae shorter than laciniae and pubescent at membranous apex, laciniae hooked at apex and with 4–5 spines between apex and middle of inner margin, maxillary palpi slender, palpomere III longer than II, IV long and subulate, ligula bifid almost to base, lobes rounded apically, labial palpi with palpomere I long and broad, II much shorter and narrower than I, III longer than II, subconical, truncate at apex; antennae moderately long, antennomeres I–III elongate and subequal in length, XI almost as long as IX and X combined; pronotum with hypomera long and narrow, delimited from pronotal disc by marginal ridge; scutellum densely punctate, finely carinate along median line, in some specimens carina only distinct at base; prosternum moderately large before front coxae, posterior margin finely cusped at middle; mesospiracular peritremes large, quadrate, contiguous along median line; process of mesoventrite projecting to apical one-third or half of mesocoxae, subparallel, rounded and free at apex and separated from process of metaventrite by a short and depressed isthmus;

576

mesocoxae moderately separated; elytra elongate, much broader than pronotum, posterior margins sinuate at outer angles; abdomen subparallel, first three visible tergites transversely impressed at base, impressions punctate or impunctate; apical margin of tergite VIII simple, without ctenidium. 224. Myrmecocephalus arizonicus (Casey) (Fig. 21.262a–h) Lorinota arizonica Casey, 1906. Moore and Legner 1975. Diagnosis. Body length 3.2–3.6 mm; colour rufocastaneous, head and apical four abdominal tergites dark brown, legs/tarsi and antennae yellowish brown (Fig. 21.262a); head elongate, orbicular, broadly and evenly rounded at base, surface between punctation glossy; eyes shorter than genae in dorsal view; antennae long, reaching apical third of elytra, with antennomeres I–VII distinctly elongate, VIII–X subquadrate to slightly transverse; pronotum narrow and elongate, narrower than head, broadest in apical third, sides strongly converging to base, sinuate, hind margins distinctly narrowly deflexed, disc sulcate along median line except for extreme apex, surface sparsely and minutely punctate and pubescent, surface between punctures smooth

21 Tribe Falagriini Mulsant and Rey, 1873

and glossy; scutellum finely carinate, carina in most confined to base; elytra with surface sparsely and minutely punctate, surface between punctures smooth and glossy; abdomen arcuate laterally, narrow at base, tergites III–V deeply, transversely impressed at base, impression coarsely punctate, tergal surface minutely and densely punctate. MALE. Tergite VIII with apical margin truncate and simple (Fig. 21.262d); sternite VIII parabolically rounded from base to apex (Fig. 21.262e); tubus of median lobe of aedeagus short and strongly sinuate ventrally in lateral view (Fig. 21.262b), in dorsal view as illustrated (Fig. 21.262c). FEMALE. Tergite VIII with apical margin truncate and simple (Fig. 21.262f); sternite VIII with apical margin broadly rounded, antecostal suture strongly sinuate and strongly produced medially (Fig. 21.262g); spermatheca S-shaped, stem sinuate, broadly curved posteriorly (Fig. 21.262h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, SK. USA: AZ, CO, NE, NM, SD, UT, WA. COLLECTION AND HABITAT DATA. Habitat: not recorded. Collecting period: not recorded. Collecting method: not recorded.

21

Tribe Falagriini Mulsant and Rey, 1873

Fig. 21.258 (a–e) Images of Bryobiota bicolor (Casey) [lectotype with images of the original labels]: (a) habitus in dorsal view; (b) female tergite VIII; (c) female sternite

577

VIII; (d) spermatheca; (e) median lobe of aedeagus (after Ahn and Ashe 1995). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

578

Fig. 21.259 (a–h) Images of Ashella giulanii (Moore) [new combination]: (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

21 Tribe Falagriini Mulsant and Rey, 1873

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. (a–e) holotype; (f–h) non-type specimen. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

21

Tribe Falagriini Mulsant and Rey, 1873

Fig. 21.260 (a–h) Images of Cordalia obscura (Gravenhorst): (a) habitus in dorsal view (apical part of abdomen removed); (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male

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tergite VIII; (e) male sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

580

Fig. 21.261 (a–h) Images of Falagria dissecta Erichson: (a) habitus in dorsal view with enlarged scutellum; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

21 Tribe Falagriini Mulsant and Rey, 1873

sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

21

Tribe Falagriini Mulsant and Rey, 1873

Fig. 21.262 (a–h) Images of Myrmecocephalus arizonicus (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male

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sternite VIII; (f) female tergite VIII; (g) female sternite VIII; (h) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

582

References Ahn K-J, Ashe JS (1995) Systematic position of the intertidal genus Bryobiota Casey and a revised phylogeny of the falagriine genera of America north of Mexico (Coleoptera: Staphylinidae: Aleocharinae). Ann Entomol Soc Am 88(2):143–154 Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton. 443 pp. [publ. 28 Dec 2000 according to CRC] Casey TL (1885) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1(4):285–336 Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7[1894]:281–606 [see reference in Chapter 7 for explanation of date] Casey TL (1906) Observations on the staphylinid groups Aleocharinae and Xantholinini, chiefly of America. Trans Acad Sci St Louis 16(6):125–134 Casey TL (1911) New American species of Aleocharinae and Myllaeninae [pp 1–245]. In: Memoirs on the Coleoptera. II. The New Era Printing, Lancaster, Pennsylvania, 259 pp Erichson WF (1839) Genera et species Staphylinorum insectorum coleopterorum familiae. Erster Band. F. H. Morin, Berlin, viii + 400 pp Fenyes A (1912) Falagria Mannh. and its relatives. J NY Entomol Soc 20:20–27 Gravenhorst JLC (1802) Coleoptera Microptera Brunsvicensia nec non exoticorum quotquot exstant in collectionibus entomologorum Brunsvicensium in genera familias et species distribuit. Carolus Reichard, Brunsuigae, lxvi + 206 + [1, errata] pp Hoebeke ER (1985) A revision of the rove beetle tribe Falagriini of America north of Mexico (Coleoptera: Staphylinidae: Aleocharinae). J NY Entomol Soc 93 (2):913–1018 Humble LM, Winchester NN, Ring RA (2000) The potentially rare and endangered terrestrial arthropods in British Columbia: revisiting British Columbia’s biodiversity [pp 101–108]. In: Proceedings of a conference on the biology and management of species and habitats at risk, Kamloops, BC, 15–19 Feb, 1999, vol 1. BC Ministry of Environment, Lands and Parks, Victoria BC and University College of the Cariboo, Kamloops, 490 pp Jacobs W (1925) Über den Gattungsnamen Cardiola Mulsant et Rey (Col.). Entomol Z (Frankfurt am Main) 38:82 Klimaszewski J, Langor D, Majka CG, Bouchard P, Bousquet Y, LeSage L, Smetana A, Sylvestre P, Pelletier G, Davies A, DesRochers P, Goulet H,

21 Tribe Falagriini Mulsant and Rey, 1873 Webster R, Sweeney J (2010) Review of adventive species of Coleoptera (Insecta) recorded from eastern Canada. Pensoft Series Faunistica No. 94. Pensoft, Sofia-Moscow, 272 pp Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Leach WE (1819) New genera. In: Samouelle (ed) The entomologist’s useful compendium; or an introduction to the knowledge of British insects, comprising the best means of obtaining and preserving them, and a description of the apparatus generally used; etc. Thomas Boys, London, 496 pp + 12 pls MacLeay W (1871) Notes on a collection of insects from Gayndah. Trans Entomol Soc New South Wales 2 [1869–1873] (2):79–158; (3):159–205 Majka CG, Klimaszewski J (2008) Introduced Staphylinidae (Coleoptera) in the Maritime Provinces of Canada. Can Entomol 140(1):48–72 Melsheimer FE (1844) Descriptions of new species of Coleoptera of the United States. Proc Acad Nat Sci 2 (2–3):26–43 Moore I (1978) Two new species of Salinamexus from western North America (Coleoptera: Staphylinidae). Entomol News 89(2–3):113–115 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Mulsant E, Rey C (1873) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens (Suite) [Tableau... des Aléochariens + Septième Branche, Myrmédoniaires, part 1]. Deyrolle, Paris, [2] + 695 pp, pls 1–5 Muona J (1984) Review of Palearctic Aleocharinae also occurring in North America (Coleoptera: Staphylinidae). Entomol Scand 15(2):227–231 Sawada K (1972) Methodological research in the taxonomy of Aleocharine. Contrib Biol Lab Kyoto Univ 24 (1):31–59 Scheerpeltz O (1964) Eine neue Art der Gattung Myrmecopora Saulcy von der pazifischen Küste Nordamericas (Col., Staph.). Deut Entomol Z 12 (2):49–51 Webster RP, Klimaszewski J, Sweeney J D, DeMerchant I (2012) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, and an addition to the fauna of Quebec, Canada: Aleocharinae. In: Klimaszewski J, Anderson R (eds) Biosystematics and ecology of Canadian Staphylinidae (Coleoptera) II. ZooKeys [Special Issue] 186:83–118

Tribe Lomechusini Fleming, 1821

22

Tribe Lomechusini Fleming (Figs. 22.263–22.266) (Ashe 2001; Maruyama 2006; Elven et al. 2012; Klimaszewski et al. 2018) Diagnosis. Epipharynx with sensillum α fully developed; galeae and laciniae moderately to greatly elongate, length of galea equal to or greater than distance to base of galea from cardo; mesocoxae broadly separated by broad processes of meso- and metaventrite, process of mesoventrite broad and short; tarsal formula 4-55; median lobe of aedeagus with ventral bridge, compressor plate covered by extension from base of bulbus. Comments. Many taxa are associated with ants. Key to Subtribes of Lomechusini 1. Pronotum broad, highly explanate (Fig. 22.263a); abdomen with prominent tufts of gold-yellow trichomes on lateral parts of sternites (Fig. 22.263a)................. Lomechusina Fleming – Pronotum moderately broad, convex and not explanate (Figs. 22.264a, 22.265a and 22.266a); abdomen without trichomes ................................. .......................... Myrmedoniina C.G. Thomson

Subtribe Lomechusina Fleming, 1821 Diagnosis. Pronotum broad and explanate; prominent tufts of gold-yellow trichomes on lateral parts of abdomen. Only one genus, Xenodusa Wasmann, occurs in North America.

Genus Xenodusa Wasmann, 1894 (Fig. 22.263a–g) (Hoebeke 1976; Klimaszewski et al. 2018) Diagnosis. Body robust, depressed, broadly subparallel, ferrugineous; length 4.0–6.5 mm; head small, half as wide as base of pronotum, with frontal impression between antennal fossae, occiput with raised band along dorsum of foramen magnum; mouthparts prognathous, labrum separate from clypeus, mandibles expanded basally, strongly curved into apical blunt point, galeae and laciniae densely pubescent, maxillary palpi with four palpomeres, apical one short and acicular; antennomere I enlarged, and incrassate, II shorter than III, remaining ones elongate; eyes equal in length to genae in dorsal view; pronotum broadly

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7_22

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transverse, highly explanate and reflexed laterally, anterior margin broadly emarginate, front angles broadly rounded, hind angles acute; hypomera broad in lateral view; prosternum of moderate size in front of procoxae with lateral projections almost reaching inflexed pronotal margins; abdomen with prominent golden-yellow tufts of trichomes laterally on segments III–V; legs long and slender. Adults and larvae are usually found in association with ants of the genera Camponotus and Formica (Hoebeke 1976). 225. Xenodusa 22.263a–g)

reflexa

Tribe Lomechusini Fleming, 1821

sinuate laterally (Fig. 22.263e); sternite VIII broadly V-shaped with apex rounded (Fig. 22.263f); spermatheca with spherical capsule with large and deep apical invagination, stem narrow, long and irregularly, loosely coiled posteriorly (Fig. 22.263g). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, MB, NB, NS, ON, QC, SK. USA: AZ, CA, CO, CT, IL, IA, MA, MD, MI, MN, MT, ND, NE, NH, NM, NV, NY, OR, RI, SC, UT, VT, WA, WI, WY.

(Walker) (Fig.

Atemeles reflexus Walker, 1866. Casey 1885 [as synonym Lomechusa montana], Wasmann 1894 [as synonym Atemeles cavus var. hirsutus], Hoebeke 1976, Majka and Klimaszewski 2008, Webster et al. 2009, Klimaszewski et al. 2018. Diagnosis. Body robust, depressed, subparallel, length 5.3–6.4 mm; colour uniformly dark ferruginous (Fig. 22.263a); forebody minutely granulose, with sparse, fine pubescence; abdomen polished, with few scattered setae dorsally and long, erect setae ventrally along sternal margins; head small, partially covered by pronotum, front with broad impression; antennae long and slender, antennomere II slightly elongate, remaining ones elongate; pronotum twice as wide as head, front margin broadly emarginate at middle, front angles broadly rounded, sides reflexed, hind angles acute; elytra as wide as base of pronotum, just over half as long as wide, at suture longer than pronotum; abdomen as wide as elytra, lateral trichomes ferruginous on segments III–V, dorsum polished, with few short setae, venter convex with scattered erect setae along sternite margins and/or setae on disc of each sternite; legs long and slender. MALE. Tergite VIII with apical margin broadly emarginate, lateral edges of emargination rounded (Fig. 22.263c); sternite VIII with sides obliquely narrowing to broadly arcuate apex (Fig. 22.263d); tubus of median lobe of aedeagus in lateral view with valves slender, long, usually contiguous, apices of gonostyli stout (Fig. 22.263b). FEMALE: tergite VIII with apical margin very shallowly emarginate, irregularly

COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: reported from ant nests of Camponotus noveboracensis (Fitch), C. laevigatus (F. Smith), C. herculeanus modoc Wheeler, and Formica subpolita Mayr. One specimen from NB captured in a Lindgren funnel trap in an old mixed forest. Several QC specimens were found in December hibernating under rock with colony of ants at the border of agriculture field. Collecting period: IV–XI. Collecting method: hand-collecting in ant nests or near ant nests, Lindgren funnel trap. Subtribe Myrmedoniina C.G. Thomson, 1867 (Maruyama 2006; Klimaszewski et al. 2018) Diagnosis. This group differs from other subtribes of Lomechusiini in having the galeae elongate, the ligula bifid, and the first three visible abdominal segments without trichomes (Maruyama 2006); otherwise the group is not well defined. Key to Genera of Myrmedoniina 1. Second and third visible abdominal tergites with large median tuberosities (best observed in lateral view) (Figs. 22.264a, b and 22.265a, b); antennomeres IV–X appearing somewhat beadshaped (Figs. 22.264a and 22.265a); forebody with short pubescence appressed to integument (Figs. 22.264a and 22.265a); genitalia as illustrated (Figs. 22.264e and 22.265c, d) ........... .............................Myrmoecia Mulsant and Rey – Second and third visible abdominal tergites without median tuberosities (Fig. 22.266a);

22

Tribe Lomechusini Fleming, 1821

585

antennomeres IV–X variable in shape but never bead-shaped (Fig. 22.266a); forebody almost without pubescence or with pubescence not appressed to integument; genitalia differently shaped, as illustrated (Figs. 22.266b, c, h)........... .................................................... Zyras Stephens

Genus Myrmoecia Mulsant and Rey, 1873 (Figs. 22.264a–e and 22.265a–g) (Klimaszewski et al. 2005, 2018; Maruyama and Klimaszewski 2006) Diagnosis. Forebody with short pubescence appressed to integument, appearing somewhat silky; pronotum trapezoidal, about one-fourth broader than long, disc moderately deeply impressed, impression broader near apex than base; second and third visible tergites of abdomen with large apico-medial tuberosities (better visible in lateral view); antennomeres IV–X appearing somewhat bead-shaped; mesoventrite with isthmus very short. 226. Myrmoecia lugubris 22.264a–e and 22.265a–g)

(Casey) (Figs.

Nototaphra lugubris Casey, 1893. Moore and Legner 1975, Seevers 1978, Klimaszewski et al. 2005, Maruyama and Klimaszewski 2006. Holotype (female): USA: La Veta, 4,7, Col. [orado]; Type USNM 39408; Notot. lugubris; Casey bequest 1925 (USNM). Examined. Diagnosis. Body length 2.4 mm; moderately broad, elongate, broadest at elytral base (Fig. 22.264a); head, pronotum, baso-lateral part of elytra and median part of abdomen dark brown to almost black, most of elytra, tibiae, tarsi, antennae and maxillary palps reddish-brown; forebody with distinct meshed microsculpture; pubescence short and moderately dense; pronotum trapezoidal in shape, broadest in apical third, broadly impressed medially, distinctly narrower than elytra, width/length ratio 1.2; elytra at suture slightly longer than pronotum along

midline, distinctly broader than pronotum, elytra/pronotum length ratio 1.4; basal metatarsomere less than three fourths as long as the two following tarsomeres combined. MALE. Unknown, see comments below. FEMALE. Tergite VIII rounded apically, antecostal suture approximately straight (Fig. 22.264c); sternite VIII parabolic apically, antecostal suture slightly sinuate, apex rounded (Fig. 22.264d); spermatheca with short, tubular and slightly broadening apically capsule bearing broadly conical invagination, stem sinuate anteriorly and posteriorly (Fig. 22.264e). DISTRIBUTION. Origin: Nearctic. CANADA: BC. USA: CO. COLLECTION AND HABITAT DATA. Habitat: not reported. Collecting period: not reported. Collecting method: not reported. Comments. Casey misidentified the sex of the holotype as a male because it is a female. Myrmoecia canadensis Maruyama and Klimaszewski (2006), described from Alberta (Fig. 22.265a–g), is very similar to M. lugubris in general appearance and colour but may be distinguished from it by the tubercle on the 5th abdominal tergite larger than that on the 4th tergite (smaller than that on the 4th in M. lugubris), and the spermatheca with the basal part forming a ring (irregularly sinuate in M. lugubris) (Maruyama and Klimaszewski 2006). These differences have to be confirmed for a larger series of specimens because the comparison between the two species was made between male and female of M. canadensis and only female of M. lugubris. For now, the shape of the spermathecal stem is the only reliable difference between the two species. It is possible that M. canadensis is conspecific with M. lugubris. Future examination of both sexes of M. lugubris from BC should clarify the status of the two species.

Genus Zyras Stephens, 1835 (Fig. 22.266a–h) (Hlaváč and Jászay 2009; Klimaszewski et al. 2005, 2018)

586

Diagnosis. Body robust, 4.0–6.0 mm, integument highly glossy, often with bright contrasting colours, moderately convex, punctation usually large and coarse; pubescence fine and slightly or not adhering to the integument, without silky appearance; head moderately convex; antennomere I as long as II and III combined, II smaller than III, IV slightly transverse, V–X moderately to distinctly transverse and more so toward the apex, terminal antennomere about as long as two preceding ones combined; pronotum broadest in apical one-third, usually about one-fourth broader than long and about one-fourth narrower than elytra, impressed basally, usually with well-defined medial antebasal fovea, punctation moderately large to large, pubescence on disc directed obliquely posteriad along arcuate lines on each side; second and third visible abdominal tergites without tuberosities; median lobe of aedeagus with enlarged bulbus and narrowly conical tubus; spermatheca very small with capsule narrowly elongate, without apparent invagination, stem short, seminal canal with multiple coils forming a bundle. Comments (Klimaszewski et al. 2018). The genus Zyras is treated here in a narrow sense following Klimaszewski et al. (2005). Seevers (1978) predicted the generic subdivision of Nearctic Zyras (sensu lato) would include transfer of many species to the genus Pella. Seevers (1978) considered Nearctic Zyras in a broad sense, with the exception of the genera Apalonia and Myrmoecia, and provided a key to twelve Nearctic species. Kistner (1971) treated at least some of the subgenera recognized by Lohse (1974) as distinct genera (e.g., Pella). 227. Zyras obliquus (Casey) (Figs. 22.266a–h) Myrmedonia obliqua Casey, 1893. Seevers 1978 [as synonym of Z. haworthi (Stephens, 1832)], Klimaszewski and Peck 1986, Klimaszewski and Winchester 2002 [as Z. pseudohaworthi], Klimaszewski et al. 2005, 2009, 2016, 2018, Webster et al. 2009, Majka and Klimaszewski 2010.

22

Tribe Lomechusini Fleming, 1821

Diagnosis. Body narrowly elongate, length 4.0–6.5 mm; head pronotum and abdomen, except for its reddish-brown margins, dark brown to black, elytra bicoloured reddish-brown with base and lateral parts of disc dark brown to black, appendages reddish-brown (Fig. 22.266a); integument highly glossy, without distinct meshed microsculpture; pubescence moderately long and sparse; antennomere I about as long as the two following antennomeres combined, II smaller than III, IV quadrate, V–X moderately transverse and slightly more so toward the apex, and the terminal one about as long as the two preceding antennomeres combined; pronotum approximately trapezoidal in shape, with small basal impression, distinctly narrower than elytra; elytra about twice as broad as length of suture; hind tarsi with basal article slightly longer than two following articles combined. MALE. Tergite VIII with two apical rounded teeth that are close together (Fig. 22.266d); sternite VIII elongate, truncate apically (Fig. 22.266e); median lobe of aedeagus with tubus straight ventrally and slightly sinuate apically with narrow and rounded apex slightly pointed ventrad in lateral view (Fig. 22.266b); in dorsal view bulbus enlarged and tubus conical (Fig. 22.266c), internal sac with two narrowly elongate subparallel structures (Figs. 22.266b, c). FEMALE. Tergite VIII truncate apically (Fig. 22.266f); sternite VIII broadly rounded apically (Fig. 22.266g); capsule of spermatheca, L-shaped, small, narrow, with subapical constriction, stem short, seminal canal very long with multiple coils forming a bundle (Fig. 22.266h). DISTRIBUTION. Origin: Nearctic. CANADA: AB, BC, LB, MB, NB, NF, NS, ON, QC, SK. USA: MI, MO, NH, NY, OR. COLLECTION AND HABITAT DATA (Klimaszewski et al. 2018). Habitat: adults from various traps in 3–5-year-old post-harvest Douglas-fir forest, transition zone of Sitka spruce forest, mixed Populus/Salix/Alnus forest, and Pinus banksiana forest, under red oak in red oak forest. Microhabitat unknown. Collecting period: VI–VIII. Collecting method: pitfall traps, window traps, and pan-traps.

22

Tribe Lomechusini Fleming, 1821

Fig. 22.263 (a–g) Images of Xenodusa reflexa (Walker): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) male tergite VIII; (d) male sternite VIII;

587

(e) female tergite VIII; (f) female sternite VIII; (g) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

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Fig. 22.264 (a–e) Images of Myrmoecia lugubris (Casey) (female holotype): (a) habitus in dorsal view (apical part of abdomen removed); (b) dorsal tubercules

22

Tribe Lomechusini Fleming, 1821

of abdomen in lateral view; (c) female tergite VIII; (d) female sternite VIII; (e) spermatheca. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

22

Tribe Lomechusini Fleming, 1821

Fig. 22.265 (a–g) Images of Myrmoecia canadensis Maruyama and Klimaszewski: (a) habitus in dorsal view (male) (apical part of abdomen removed); (b) dorsal tubercules of abdomen in lateral view; (c) median lobe of

589

aedeagus in lateral view; (d) median lobe of aedeagus in dorsal view; (e) male tergite VIII; (f) male sternite VIII; (g) spermatheca. (a–f) holotype; (g) paratype. Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

590

Fig. 22.266 (a–h) Images of Zyras obliquus (Casey): (a) habitus in dorsal view; (b) median lobe of aedeagus in lateral view; (c) median lobe of aedeagus in dorsal view; (d) male tergite VIII; (e) male sternite VIII; (f) female

22

Tribe Lomechusini Fleming, 1821

tergite VIII; (g) female sternite VIII; (h) spermatheca. (b, c, h) after Klimaszewski and Winchester (2002). Scale bar for habitus ¼ 1 mm, remaining scale bars ¼ 0.2 mm

22

Tribe Lomechusini Fleming, 1821

References Ashe JS (2001) Key to the tribes and genera of Nearctic Aleocharinae. 8. pp 299–374. In: Arnett RH Jr, Thomas CM (eds) American beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC, Boca Raton, 443 pp [publ. 28 Dec 2000 according to CRC] Casey TL (1885) New genera and species of Californian Coleoptera. Bull Calif Acad Sci 1(4):283–336 Casey TL (1893) Coleopterological notices. V. Ann NY Acad Sci 7 [1894]:281–606 [see reference in Chapter 7 for explanation of date] Elven H, Bachmann L, Gusarov VI (2012) Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta). Zool Scr 41(6):617–636 Fleming J (1821) Insecta [pp 41–56]. In: Supplement to the fourth, fifth and sixth editions of the Encyclopaedia Britannica, with preliminary dissertations on the history of sciences, 5th vol. A. Constable and Company, Edinburgh, 584 pp Hlaváč P, Jászay T (2009) A revision of the genus Zyras (Zyras) Stephens, 1835 (Coleoptera, Staphylinidae, Aleocharinae). I. Current classification status and the redefinition of the genus. ZooKeys 29:49–71 Hoebeke ER (1976) A revision of the genus Xenodusa (Staphylinidae, Aleocharinae) for North America. Sociobiology 2(2):109–143 Kistner DK (1971) Studies of Japanese myrmecophiles. Part I. The genera Pella and Falagria (Coleoptera, Staphylinidae). Entomological Essays to Commemorate the Retirement of Professor K. Yasumatsu, 1971:141–165. Hokuryukan Publishing, Tokyo Klimaszewski J, Peck SB (1986) A review of the cavernicolous Staphylinidae (Coleoptera) of eastern North America: Part I. Aleocharinae. Quaestiones Entomol 22(2):51–113 Klimaszewski J, Winchester NN (2002) Aleocharine rove beetles (Coleoptera Staphylinidae) of the ancient Sitka spruce forest on Vancouver Island, British Columbia, Canada. Mém Soc R Belge Entomol 40:3–126 Klimaszewski J, Pelletier G, Maruyama M, Hlaváč P (2005) Canadian species of the Zyras group of genera and review of types from America north of Mexico (Coleoptera, Staphylinidae, Aleocharinae). Rev Suisse Zool 112(3):703–733 Klimaszewski J, Lynch D, Majka CG, Renkema J, Savard K, Hlaváč P (2009) Pella glooscapi, a new rove beetle, and new records of aleocharines from Nova Scotia, Canada (Coleoptera, Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:35–44 Klimaszewski J, Larson DJ, Labrecque M, Bourdon C (2016) Twelve new species and fifty-three new provincial distribution records of Aleocharinae rove beetles of Saskatchewan, Canada (Coleoptera, Staphylinidae). ZooKeys 610:45–112

591 Klimaszewski J, Webster RP, Langor DW, Brunke A, Davies A, Bourdon C, Labrecque M, Newton AF, Dorval J-A, Frank JH (2018) Aleocharine rove beetles of eastern Canada (Coleoptera, Staphylinidae, Aleocharinae): a glimpse of megadiversity. Springer, Cham, 902 pp Lohse GA (1974) Hypocyphtinae, Aleocharinae [in part, pp 7–72, 221–292]. In: Freude H, Harde KW, Lohse A (eds) Die Käfer Mitteleuropas. Band 5: Staphylinidae II (Hypocyphtinae und Aleocharinae), Pselaphidae. Goecke & Evers, Krefeld, 381 pp Majka CG, Klimaszewski J (2008) New records of Canadian Aleocharinae (Coleoptera: Staphylinidae). In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys [Special Issue] 2:85–114 Majka CG, Klimaszewski J (2010) Contributions to the knowledge of the Aleocharinae (Coleoptera, Staphylinidae) in the Maritime Provinces of Canada. ZooKeys 46:15–19 Maruyama M (2006) Revision of the Palearctic species of the myrmecophilous genus Pella (Coleoptera, Staphylinidae, Aleocharinae). Natl Sci Mus Monogr (Tokyo) 32:iii + 207 pp Maruyama M, Klimaszewski J (2006) Notes on myrmecophilous aleocharines (Insecta, Coleoptera, Staphylinidae) from Canada, with a description of a new species of Myrmoecia. Bull Natl Sci Mus Tokyo (A) 32(3):125–131 Moore I, Legner EF (1975) A catalogue of the Staphylinidae of America north of Mexico (Coleoptera). University of California, Division of Agricultural Sciences, Special Publication No. 3015:1–514 Mulsant E, Rey C (1873) Histoire naturelle des Coléoptères de France. Brévipennes. Aléochariens (Suite) [Tableau... des Aléochariens + Septième Branche, Myrmédoniaires, part 1]. Deyrolle, Paris, [2] + 695 pp, pls 1–5 Seevers CH (1978) A generic and tribal revision of the North American Aleocharinae (Coleoptera: Staphylinidae) [with additions and annotations by Lee H. Herman]. Fieldiana: Zoology 71:vi + 289 pp Stephens JF (1832) Illustrations of British entomology, or a synopsis of indigenous insects, containing their generic and specific distinctions, with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable. Mandibulata, vol 5. Baldwin & Cradock, London, pp 1–240, pls 24–26 Stephens JF (1835) Illustrations of British entomology, or a synopsis of indigenous insects, containing their generic and specific distinctions, with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable. Mandibulata, vol 5. Baldwin and Cradock, London, pp 369–477 Thomson CG (1867) Skandinaviens Coleoptera, synoptiskt bearbetade. Supplementum. Tom IX. Lundbergska Boktryckeriet, Lund, 407 pp

592 Walker F (1866) List of Coleoptera [and descriptions]. [Appendix pp 309–334]. In: Lord JK (ed) The naturalist in Vancouver Island and British Columbia, vol II. R. Bentley, London, vii + [2] + 375 pp Wasmann E (1894) Kritisches Verzeichniss der myrmekophilen und termitophilen Arthropoden. Mit Angabe der Lebensweise und mit Beschreibung neuer Arten. F. L. Dames, Berlin, xiii + 231 pp

22

Tribe Lomechusini Fleming, 1821

Webster RP, Klimaszewski J, Pelletier G, Savard K (2009) New Staphylinidae (Coleoptera) records with new collection data from New Brunswick, Canada. I. Aleocharinae. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys [Special Issue] 22:171–248

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.1 List of aleocharine species recorded from British Columbia with their distribution records in Canada and the United States Taxonomic name Tribe Aleocharini Fleming Aleochara acerba (Casey) Aleochara assiniboin Klimaszewski Aleochara bilineata Gyllenhal{ Aleochara bimaculata Gravenhorst

Aleochara carmanah Klimaszewski Aleochara castaneipennis Mannerheim Aleochara curtidens Klimaszewski Aleochara curtula (Goeze){ Aleochara fumata Gravenhorst{

Canadian distribution

United States distribution

BC BC, MB, NT, ON, SK, YT AB, BC, MB, NF, NS, NB, ON, PE, QC, SK AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK

ID, WA

BC AB, BC, LB, NB, NF, NS, NT, ON, QC, YT BC BC, NB, NF, NS, ON, PE, QC AB, BC, MB, NB, NF, NS, ON, QC, PE, YT

Aleochara gracilicornis Bernhauer

BC, MB, NB, NF, NS, NT, ON, QC, SK

Aleochara lacertina Sharp

AB, BC, MB, NB, NF, NS, ON, QC, SK

Aleochara lanuginosa Gravenhorst{

AB, BC, MB, NB, NF, NS, ON, QC BC, SK BC, NB, NF, NS, QC AB, BC, NB, NF, NS, ON, QC, SK BC BC BC, NB, ON, SK AB, BC, MB, QC, SK BC

Aleochara laramiensis (Casey) Aleochara litoralis (Mäklin) Aleochara morion Gravenhorst{ Aleochara pacifica (Casey) Aleochara quadrata Sharp Aleochara rubricalis (Casey) Aleochara suffusa (Casey)
Aleochara sulcicollis Mannerheim


ID, IL, MA, ME, NY, OR, RI, WA, WI AL, AZ, CA, CO, CT, DC, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY AK, AZ, CO, WA, WY CA IL, MA, ME, NH, NJ, NY, OH, PA, RI, VT AL, AR, AZ, CA, GA, MA, MD, ME, MN, MO, NC, NH, NJ, NM, NY, OK, OR, PA, TN, VT, WA, WI, WV, WY AR, AZ, CO, FL, IL, IN, KS, LA, MA, MD, ME, MI, MN, MO, MT, NH, NJ, NM, NY, OH, PA, RI, SD, UT, WI AZ, CA, CO, FL, MA, ME, MN, MT, ND, NH, NM, NV, NY, OK, OR, PA, RI, TN, TX, UT, WA, WI CA, CO, ID, MA, MN, NC, NH, NJ, NY, OR, PA, RI, WA, WI CO, WY AK, CA, FL, MA, NJ, NY, RI, VA CA, CT, NV CA, WA AZ, CA, OR, WA AK, AZ, CO, ID, NM, WA, WY AK, CA, OR, WA (continued)

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.1 (continued) Taxonomic name Aleochara tahoensis Casey Aleochara verna Say

Aleochara villosa Mannerheim{ Tribe Athetini Casey Adota maritima (Mannerheim) Aloconota cambrica (Wollastone){ Aloconota carmanah Klimaszewski Amischa analis (Gravenhorst){

Atheta alesi Klimaszewski and Brunke Atheta brumalis Casey Atheta brunswickensis Klimaszewski Atheta capsularis Klimaszewski Atheta celata (Erichson)
Atheta cheersae Klimaszewski Atheta concessa Casey Atheta copleyi Klimaszewski, sp. n. Atheta cryptica (Lohse) Atheta dadopora C.G. Thomson


Atheta districta Casey Atheta fanatica Casey Atheta formalis (Casey) Atheta frosti Bernhauer Atheta godini Klimaszewski, sp. n. Atheta graminicola (Gravenhorst)


Atheta hampshirensis Bernhauer Atheta holmbergi Bernhauer Atheta irrupta (Casey) Atheta keeni Casey Atheta longicornis (Gravenhorst){ Atheta lucifera Bernhauer Atheta meduxnekeagensis Webster and Klimaszewski Atheta metlakatlana Bernhauer Atheta munsteri Bernhauer


Canadian distribution AB, BC, MB, NB, NF, NS, NT, ON, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT AB, BC, NB, ON, QC, SK BC BC BC BC [NPR], LB, MB, NB, NF, NS, ON, PE, QC, SK ON, BC [NPR] BC BC [NPR], NB, NS, ON, QC, YT BC [NPR], LB, NB, NF, ON, QC, YT BC, NB, NF, NS, SK BC BC BC BC, NF, QC, YT AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, QC, SK AB, BC, LB, NB, NS, QC, SK, YT BC BC, LB, MB, NB, NS, ON, QC, SK BC AB, BC, LB, MB, NB, NF, NT, ON, QC, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, QC BC BC BC BC, NB, NF, NS, QC, SK BC [NPR] NB, BC BC BC [NPR], MB, NT, YT

United States distribution AR, CA, CO, MT, NH, NM, NV, OR, SD, WA, WI AK, AZ, CA, CO, DC, FL, GA, IA,ID, IL, IN, KS, MA, ME, MI, MN, MO, MT, NH, NC, NJ, NM, NY, NV, OK, OR, PA, SC, SD, TN, TX, UT, VT, WA, WI, WY AK, CA, OK, OR, WA

AK, CA CA CA, CO, IN, MA, ME, MI, NH, NY, PA, VT, WA

CA

AK, MA, WV

AK, NY, PA, RI

NV

MA, NC, NH, NY, PA, RI, VT, WV

AK, OR

AK, CA, NC, NH, NY, PA, RI, WA [as A. moesta] AK AK, OR CA, MN, OR CA, WA

OR AK (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

595

Table A.1 (continued) Taxonomic name Atheta nescia (Casey) Atheta picipennis (Mannerheim) Atheta platonoffi Brundin
Atheta prudhoensis (Lohse) Atheta pseudoklagesi Klimaszewski and Webster Atheta pseudometlakatlana Klimaszewski and Godin Atheta pseudoschistoglossa Klimaszewski and Webster Atheta relicata (Casey) Atheta remulsa Casey

Atheta ringi Klimaszewski Atheta ripariides Newton Atheta rurigena Casey Atheta stercoris Fenyes Atheta strigosula Casey Atheta surgens (Casey) Atheta terranovae Klimaszewski and Langor Atheta ventricosa Bernhauer Atheta wheelerae Klimaszewski, sp. n. Atheta winchesteri Klimaszewski, sp. n. Boreophilia nomensis (Casey) Boreostiba pseudolaticollis Klimaszewski and Godin, sp. n. Boreostiba parvipennis (Bernhauer) Clusiota impressicollis (Bernhauer) Dalotia coriaria (Kraatz){ Dinaraea angustula (Gyllenhal){

Dinaraea inexpectata Klimaszewski, sp. n. Dinaraea pacei Klimaszewski and Langor Dinaraea subdepressa (Bernhauer) Dochmonota rudiventris (Eppelsheim) Earota dentata (Bernhauer)

Canadian distribution BC BC AB, BC, LB, NB, NF, NS, ON, SK, YT BC, LB, NB, NF, NS, ON, QC, SK, YT BC [NPR], AB, QC, NB, NF, SK, YT BC [NPR], YT BC, NB, SK BC AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT BC BC [NPR], YT BC BC BC, LB, MB, NB, NF, ON, QC, SK, YT BC BC [NPR], LB, NB, NF, ON, QC, SK, YT AB, BC, LB, NB, NF, NS, ON, QC, SK, YT BC BC BC BC

United States distribution AK AK AK, NY, VT

AK

AK, NY

AK

AK, NY

AK [NSR] AK, DC, IN, NC, NJ, NY, PA, VT

AK AK [NSR]

BC [NPR], AB, LB, NF, NT, QC, YT BC, NB, NF, ON AB, BC, NB, NS, ON, QC AB, BC [NPR], LB, NB, NF, NS, ON, PE, QC, SK, YT BC

AK, NH

BC, LB, NB, NF, QC, SK, YT BC [NPR], NB, ON [NPR] BC [NPR], NB, NF, NT, QC, YT AB, BC, MB, NB, NF, NS, ON, QC, SK, YT

AK

CA, WA CA, FL, LA, MA, NJ, NY CA, NY, PA

NH ID, MA AK, AL, AZ, CA, CO, DC, IA, ID, IL, IN, MA, ME, MD, MT, NC, NH, NJ, NM, NV, NY, OH, OR, PA, VA, WA (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.1 (continued) Taxonomic name Geostiba horwoodae Klimaszewski & Godin, sp. n. Goniusa caseyi Gusarov Liogluta atriventris (Casey) Liogluta nitens (Mäklin) Liogluta quadricollis (Casey) Liogluta scudderi Klimaszewski, sp. n. Liogluta terminalis (Casey) Liogluta trapezicollis Lohse Liogluta wickhami (Casey) Lypoglossa angularis (Mäklin, 1853) Lypoglossa franclemonti Hoebeke

Mocyta breviuscula (Mäklin)

Mocyta discreta (Casey) Mocyta fungi (Gravenhorst){

Nehemitropia lividipennis (Mannerheim) { Ousipalia pacifica Casey Paragoniusa myrmicae Maruyama and Klimaszewski Paraleptonia pacei Klimaszewski Pelioptera robusta (Gusarov) Philhygra angusticauda (Bernhauer) Philhygra botanicarum (Muona)
Philhygra charlottae Klimaszewski, sp. n. Philhygra clemens (Casey) Philhygra laevicollis (Mäklin) Philhygra terrestris Klimaszewski and Godin Philhygra terrivaga Klimaszewski, sp. n. Pontomalota opaca (LeConte) Psammostiba comparabilis (Mäklin) Psammostiba kenaii Gusarov Schistoglossa campbelli Klimaszewski Schistoglossa carexiana Klimaszewski Schistoglossa charlottae Klimaszewski Schistoglossa hampshirensis Klimaszewski

Canadian distribution BC

United States distribution

AB, BC, MB BC AB, BC, YT BC BC AB, BC, LB, NB, NF, NS, ON, QC, YT BC [NPR], YT BC AB, BC, NF, QC, YT BC [NPR], AB, MB, NB, NF, NS, NT, ON, QC, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT BC [NPR], , MB, ON, QC, SK AB, BC, LB, MB, NB, NF, NS, NU, ON, PE, QC, SK, YT BC [NPR], NB, NF, NS, ON, PE, QC, SK BC, SK [NPR] AB, BC, LB, NB

DC, MA, NH, NJ, NY, TX

BC BC BC, NB BC, LB, NB, NF, NS, ON, SK, YT BC

AK, OR, WA

MT, NH AK AK, AZ, CA, CO, ME, NH, NM, OR, WA ME, NH, NY, VT

AK, CA, NV, OR

IA, MN, WA [NSR] AK, CA, ME, MA, MN, NH, NY, OR, RI, WA

CA, LA, MA, MI, MN, NE, NJ, NM, NY, PA, TX, VT CA, WA [NSR]

OR AK, NH

BC, MB, NB, NS, ON, QC, YT BC, NB, NS, ON BC [NPR], NB, SK, YT

OH, WI AK, WA AK [NSR]

BC

OR

BC BC BC AB, BC BC, SK BC AB, BC [NPR], QC

AK, CA, OR, WA AK, CA AK, CA

NH (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

597

Table A.1 (continued) Taxonomic name Seeversiella globicollis (Bernhauer) Strigota ambigua (Erichson)

Strigota obscurata Klimaszewski and Brunke Tarphiota fucicola (Mäklin) Tarphiota geniculata (Mäklin) Thinusa fletcheri Casey Thinusa maritima (Casey) Trichiusa columbica Casey Trichiusa pilosa Casey Tribe Autaliini C.G. Thomson Autalia puncticollis Sharp{ Autalia rivularis (Gravenhorst){ Autalia truncatula Casey Tribe Diglottini Jakobson Bryothinusa catalinae Casey Tribe Falagriini Mulsant and Rey Bryobiota bicolor (Casey) Cordalia obscura (Gravenhorst){

Canadian distribution AB, BC, MB, NB, NF, NS, ON, QC, SK BC [NPR], LB, NB, NF, NS, ON, PE, QC, SK, YT BC [NPR], NB, ON, QC, SK BC BC BC BC BC AB, BC, NB, NS, ON

United States distribution AZ, CO, ID, MI, MN, MT, NH, NM, SD, WI

BC AB, BC, LB, NB, NF, NS, ON, QC, SK BC [NPR] BC [NPR]

CA, WA CA, ME, MI, MN, NJ, NY, OR, PA, VT

BC BC, NB, NS, ON, QC

CA CA, CO, CT, IL, MA, MD, MI, NH, NJ, NY, OH, PA, WA AL, AZ, CA, CO, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, ME, MA, MD, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NY, OH, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY AZ, CO, NE, NM, SD, UT, WA

Falagria dissecta Erichson

AB, BC, MB, NB, NS, ON, QC, SK

Myrmecocephalus arizonicus (Casey) Tribe Gymnusini Heer Gymnusa atra Casey


AB, BC, SK

Gymnusa grandiceps Casey Gymnusa pseudovariegata Klimaszewski Tribe Homalotini Heer Encephalus americanus Seevers Gyrophaena affinis Mannerheim{

Gyrophaena californica (Casey) Gyrophaena keeni Casey Gyrophaena nana (Paykull)
Gyrophaena uteana Casey

CA, CO, CT, IA, KS, MA, MO, NC, NJ, NM, NY, NV, TX

AK, CA AK, CA AK, CA AK, CA, OR, WA ID, IN, KS, OH, RI

AK, CA CA

AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT BC [NPR], LB, MB, NB, NF, NS, ON, QC AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT

AK, IL, IA, MA, ME, MI, MN, NH, NY

AB, BC BC, MB, NB, NF, NS, ON, QC, SK

IL, MT, NM AZ, DC, IL, IN, IA, KY, MA, ME, MI, MN, MO, NC, NH, NJ, NM, NY, OH, PA, TN, WA, WI, WV CA, CO, NM FL, MA, ME, MT, NH, NY, OR, TN, WA, WY

BC AB, BC, LB, MB, NB, NF, NS, ON, QC, YT AB, BC, MB, NB, NF, ON, YT AB, BC, NB, ON, QC, SK

New England states, IL, MA, MD, ME, MI, NH, NY, RI AK, CA, MA, OR, WA

AK, CA, MA, ME, MI, MT, WI, WY CA, CO, UT (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.1 (continued) Taxonomic name Leptusa gatineauensis Klimaszewski and Pelletier Leptusomorpha claudiae Klimaszewski sp. n., Leptusomorpha Klimaszewski and Hoebeke, gen. n. Silusa californica (Bernhauer)

Silusa opaca Fenyes Silusa vesperis Casey Stictalia brevicornis Casey Stictalia californica (Casey) Stictalia carlottae Casey Stictalia kranabetteri Klimaszewski and Godin, sp. n. Tribe Hypocyphtini Laporte Cypha crotchii (Horn) Holobus vancouveri Klimaszewski Oligota parva Kraatz{ Tribe Liparocephalini Fenyes Amblopusa brevipes Casey Diaulota densissima Casey Liparocephalus brevipennis Mäklin Liparocephalus cordicollis LeConte Paramblopusa borealis (Casey) Tribe Lomechusini Fleming Myrmoecia lugubris (Casey) Xenodusa reflexa (Walker)

Zyras obliquus (Casey) Tribe Myllaenini Ganglbauer Myllaena audax Casey Myllaena decreta Casey Myllaena fenyesi Bernhauer Myllaena insomnis Casey

Tribe Oxypodini C.G. Thomson Alfocalea montana Klimaszewski Betocalea pacifica Klimaszewski Blepharhymenus illectus (Casey) Crataraea suturalis (Mannerheim){ Calodera bennetti Klimaszewski, sp. n. Devia prospera (Erichson)
Gnathusa eva Fenyes Gnathusa tenuicornis Fenyes Megocalea lemieuxi Klimaszewski

Canadian distribution AB, BC, NB, NF, NS, ON, QC, SK BC

United States distribution

AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT BC BC BC BC BC BC

AK, CA, MN

AB, BC, SK BC BC [NPR], NB, PE

ID, OR, WA

BC BC BC BC BC

AK, CA AK, CA, OR, WA AK AK, CA, OR, WA AK, CA, OR, WA

BC AB, BC, MB, NB, NS, ON, QC, SK

CO, WA AZ, CA, CO, CT, IL, IA, MA, MD, MI, MN, MT, ND, NE, NH, NM, NV, NY, OR, RI, SC, UT, VT, WA, WI, WY MI, MO, NH, NY, OR

AB, BC, LB, MB, NB, NF, NS, ON, QC, SK BC, LB, NB, NF, NT, ON, QC AB, BC BC AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK, YT AB, BC BC BC [NPR] BC, LB, NB, NS, ON, QC, SK BC AB, BC, LB, MB, NB, NT, ON, QC, SK, YT AB, BC, SK, YT AB, BC, NB, YT BC

CA CA, OR, WA CA CA, AK

CA, MA, MO, NV, TX

AK, CA, GA, IL, LA, MA, NH, NJ, NY, OR, RI, UT, WA AZ, CA, NM, OR, UT, WA AZ, CA, NV, OR, UT, WA AK, ID, MA, MI, MN, MT, NY, WI

OR CA, IA, IL, IN, MA, MO, PA, RI, SC, VA, VT AK [NSR] AK, CA, CO, MA, MI, MN, MT, NH, NM, OR, SD, UT, WA, WI, WY CA AK, CA, OR (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

599

Table A.1 (continued) Taxonomic name Meotica pallens (Redtenbacher){ Metocalea lindgreni Klimaszewski Mniusa yukonensis (Klimaszewski and Godin) Neoisoglossa agnita (Casey) Neothetalia canadiana Klimaszewski Neothetalia columbiana (Klimaszewski) Neothetalia nimia (Casey) Neothetalia pallitarsis (Kirby) Neothetalia robergei Klimaszewski, sp. n. Neothetalia smetanai Klimaszewski Ocalea vancouveri Casey Ocyusa canadensis Lohse Oxypoda canadensis Klimaszewski

Oxypoda convergens Casey

Oxypoda famula Casey Oxypoda frigida Bernhauer Oxypoda gatosensis Bernhauer Oxypoda glenorae Casey Oxypoda impressa Casey Oxypoda irrasa Mäklin Oxypoda lacustris Casey

Oxypoda longicarinata Klimaszewski Oxypoda manitobae Casey Oxypoda nimbata Casey Oxypoda opaca (Gravenhorst){ Oxypoda orbicollis Casey

Oxypoda regressa Casey Oxypoda smithi Klimaszewski Oxypoda stanleyi Klimaszewski & McLean Oxypoda sylvia Casey

Oxypoda vancouveri Klimaszewski Parocalea pseudobaicalica Lohse Phloeopora arctica Lohse

Canadian distribution BC, NB, NS, ON, YT [NPR] BC BC, NB, NS, QC, YT

United States distribution NJ, RI

BC [NPR], AB BC, LB, NB, QC, YT BC BC AB, BC BC

AZ AK AK

BC BC BC [NPR], NB, NF, ON, SK, YT AB, BC [NPR], LB, MB, NF, NT, ON, QC, YT. AB, BC [NPR], LB, MB, NB, NF, NS, ON, QC BC AB, BC, LB, NB, NF, NS, NT, ON, QC, YT BC BC BC AB, BC [NPR], SK, YT AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK, YT BC BC, MB, SK BC BC, LB, NB, NF, NS, ON, QC BC [NPR], AB, LB, NB, NS, ON, QC, SK, YT BC BC BC AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT BC BC [NPR], NT, QC, YT AB [NPR], BC [NPR], NT, ON, YT

AK

AK AK, NH

IA, MO, NY

AK, NH CA

AK, OR AK, WA

CO CA NC, NY, PA, SC, TN, VT WI

AK, ME, NH

AK, NH

(continued)

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Table A.1 (continued) Taxonomic name Phloeopora canadensis Klimaszewski and Langor Phloeopora oregona Casey Tribe Placusini Mulsant & Rey Placusa incompleta Sjöberg{ Placusa petulans Casey Placusa pseudosuecica Klimaszewski Placusa tachyporoides (Waltl){ Placusa tacomae Casey Placusa vaga Casey Tribe Tachyusini C.G. Thomson Brachyusa americana (Fenyes) [unsettled status] Brachyusa helenae (Casey) Gnypeta baranowskii Klimaszewski, sp. n. Gnypeta brevicornis Casey Gnypeta caerulea (C.R. Sahlberg)


Canadian distribution BC [NPR], NF, NB

United States distribution

BC [NPR], NB

OR

AB, BC, NB, NF, NS, ON, QC, SK BC AB, BC, ON, QC, SK AB, BC, NB, NS, ON, QC, SK AB, BC, NB, NF, NS, NT, ON, QC, SK, YT AB, BC, NB, NS, NT, ON, QC, SK, YT

WA

CA, MA AZ, CO, MA, ME, MN, NY, WA CA

BC BC [NPR], LB, NB, NF, NT, ON, SK, YT BC

Tachyusa cavicollis LeConte

BC AB, BC, LB, MB, NF, NS, NB, NT, ON, PE, QC, SK, YT BC AB, BC, ON AB, BC AB, BC AB, BC, MB,NB, NF, NS, NT, ON, QC BC

Tachyusa obsoleta Casey

BC, NB, SK

Gnypeta crebrepunctata (Casey) Gnypeta helenae Casey Gnypeta lohsei Klimaszewski Paradilacra densissima (Bernhauer) Tachyusa americanoides Paśnik

CA, WA

AK, MT

AK

CA, OR AZ, CA, CO, MT, NV, NM, ND, OR AK, WA CA, MT, NV, ND, OR, UT IL, MA, MT, NH, NY, PA, VT AR, FL, IL, IN, KS, LA, MA, ME, MD, NE, NC, NH, NJ, NM, OH, PA, SD, TX, VA, VT, WY AZ, AR, CO, IL, IN, KS, KY, MA, MN, MT, NC, NE, ND, NM, NH, NJ, OH, PA, SD, TX, VT, WA, WY


Holarctic species, {adventive species, {species of uncertain distribution status—adventive in Canada or Holarctic, species without marks are native, NPR new provincial record, NSR new state record. Tribes, genera and species are listed in alphabetical order

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601

Table A.2 List of aleocharine species recorded from Canada with their distribution records in Canada and the United States Taxonomic name Tribe Aleocharini Fleming Aleochara acerba (Casey) Aleochara assiniboin Klimaszewski Aleochara bilineata Gyllenhal{ Aleochara bimaculata Gravenhorst

Aleochara carmanah Klimaszewski Aleochara castaneimarmotae Klimaszewski, Webster and Brunke Aleochara castaneipennis Mannerheim Aleochara caviceps (Casey) Aleochara curtidens Klimaszewski Aleochara curtula (Goeze){

Canadian distribution

United States distribution

BC BC, MB, ON, SK, YT

ID, WA

AB, BC, MB, NF, NS, NB, ON, PE, QC, SK AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK

ID, IL, MA, ME, NY, OR, RI, WA, WI AL, AZ, CA, CO, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY

BC NB, ON

AR, IL, KY, MO, TN

AB, BC, LB, NB, NF, NS, NT, ON, QC, YT NB, ON, QC BC

AK, AR, AZ, CO, WA, WY

BC, NB, NF, NS, ON, PE, QC ON

AZ, IA, ID, IN, MT, NJ, NV, NY, PA CA IL, MA, ME, NH, NJ, NY, OH, PA, RI, VT

Aleochara daviesi Klimaszewski and Brunke Aleochara elisabethae Klimaszewski and Larson Aleochara fumata Gravenhorst{

AB, BC, MB, NB, NF, NS, ON, QC, PE, YT

Aleochara gracilicornis Bernhauer

BC, MB, NB, NF, NS, NT, ON, QC, SK

Aleochara inexspectata Klimaszewski Aleochara lacertina Sharp

NB, NF, NS, ON, QC, SK

AL, AR, AZ, CA, GA, MA, MD, ME, MN, MO, NC, NH, NJ, NM, NY, OK, OR, PA, TN, VT, WA, WI, WV, WY AR, AZ, CO, FL, IL, IN, KS, LA, MA, MD, ME, MI, MN, MO, MT, NH, NJ, NM, NY, OH, PA, RI, SD, UT, WI MI, WI

AB, BC, MB, NB, NF, NS, ON, QC, SK AB, BC, NB, NF, NS, ON, QC BC, SK

AZ, CA, CO, FL, MA, ME, MN, MT, ND, NH, NM, NV, NY, OK, OR, PA, RI, TN, TX, UT, WA, WI CA, CO, ID, MA, MN, NC, NH, NJ, NY, OR, PA, RI, WA, WI CO, WY

ON, QC

AL, AR, AZ, CA, CO, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, MA, ME, MD, MI, MN, MO, NC, NE, NH, NJ, NY, OH, OK, PA, RI, SC, TN, TX, VA, VT, WV, WI AK, CA, FL, MA, NJ, NY, RI, VA AL, AR, AZ, DC, FL, GA, IL, IN, KS, KY, LA, MA, MD, ME, MI, MO, NC, NH, NJ, NY, OH, OK, PA, SC, TN, TX, VA, WV, WI CA, CT, NV

Aleochara lanuginosa Gravenhorst{ Aleochara laramiensis (Casey) Aleochara lata Gravenhorst{

AB, SK

Aleochara litoralis (Mäklin) Aleochara lustrica Say

BC, NB, NF, NS, QC ON

Aleochara morion Gravenhorst{ Aleochara ocularis Klimaszewski

AB, BC, NB, NF, NS, ON, QC, SK MB, NB, ON, QC

IA, IN, KY, MA, MO, PA (continued)

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Table A.2 (continued) Taxonomic name Aleochara pacifica (Casey) Aleochara quadrata Sharp Aleochara rubricalis (Casey) Aleochara rubripes Blatchley Aleochara sculptiventris (Casey) Aleochara sekanai Klimaszewski Aleochara shelleyae Klimaszewski and Langor Aleochara speculicollis Bernhauer Aleochara suffusa (Casey)
Aleochara sulcicollis Mannerheim
Aleochara tahoensis Casey Aleochara thoracica (Casey) Aleochara trisecta (Casey) Aleochara tristis Gravenhorst { Aleochara verna Say

Aleochara villosa Mannerheim{ Amarochara brevios Assing Amarochara duryi (Casey) Amarochara fenyesi Blatchley Amarochara inquilina (Casey) Tribe Athetini Casey Acrotona onthophilides Newton, 2017 Acrotona brachyoptera Klimaszewski and Webster Acrotona horwoodae Klimaszewski and Godin Acrotona manitobensis Klimaszewski and Godin Acrotona pseudopygmaeidesides Newton [2017: 10] Acrotona pseudopygmaeides Newton Acrotona recondita (Erichson) Acrotona sequestralis (Casey) Acrotona smithi (Casey) Acrotona sphagnorum Klimaszewski and Webster Acrotona subpygmaea (Bernhauer)

Canadian distribution BC BC BC, NB, ON, SK MB, NB, ON, QC NB, NF, ON, QC AB, LB, MB, NB, NT, ON, SK, YT NF, ON

United States distribution CA, WA CA, NV, OR, WA. Mexico AZ, CA, OR, WA CA, IN, KY, MA, ME, NH, PA, WA AL, AR, DC, MA, ME, NC, NH, NJ, NY, OH, PA, RI, SC, TN, VT, WI AK

AB, NB, ON, QC, SK

AR, AZ, CA, CO, MI, NV, TX. Mexico

AB, BC, MB, QC, SK BC

AK, AZ, CO, ID, NM, WA, WY AK, CA, OR, WA

AB, BC, MB, NB, NF, NS, NT, ON, SK, YT NB, NF, ON ON, QC NB, NF, ON, QC

AR, CA, CO, MT, NH, NM, NV, OR, SD, WA, WI

AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT

IL, MA, ME, MI, NH, NJ, PA, RI, WI AZ, CA, IA, ID, NY, OR, PA, TN CA, MN, NE, PA, VT

AB, BC, NB, ON, QC, SK

AK, AZ, CA, CO, DC, FL, GA, IA,ID, IL, IN, KS, MA, ME, MI, MN, MO, MT, NH, NC, NJ, NM, NY, NV, OR, PA, SC, SD, TN, TX, UT, VT, WA, WI, WY AK, CA, OK, OR, WA

ON NB, QC ON NB

KS IA, MA, NY, PA GA, IN, KS. IA

YT

AK

MB, NB, ON YT MB SK

NF SK NB, LB MB, NB, ON, QC NB

AR, CA, NH, NV, NY, PA, RI IA NY

NB, NS, ON, SK

IN, MA, NY, RI (continued)

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603

Table A.2 (continued) Taxonomic name Adota maritima (Mannerheim) Alevonota gracilenta (Erichson){ Aloconota cambrica (Wollaston) Aloconota carmanah Klimaszewski Aloconota neocambrica Klimaszewski and Langor Aloconota sulcifrons (Stephens){ Amischa analis (Gravenhorst){ Amischa praelonga Klimaszewski and Godin Anatheta planulicollis (Casey) Anatheta surrufa (Casey) Atheta acadiensis Klimaszewski and Majka Atheta aemula (Erichson) Atheta alesi Klimaszewski and Brunke Atheta alphacrenuliventris Klimaszewski and Webster Atheta altaica Bernhauer
Atheta amicula (Stephens){ Atheta annexa Casey Atheta atramentaria (Gyllenhal){ Atheta avalon Klimaszewski and Langor Atheta borealides Newton Atheta brumalis Casey Atheta brunswickensis Klimaszewski Atheta bubo Klimaszewski and Webster Atheta burwelli (Lohse) Atheta cadeti Klimaszewski and Godin Atheta campbelli (Lohse) Atheta capsularis Klimaszewski Atheta caribou (Lohse) Atheta celata (Erichson)
Atheta chartersensis Klimaszewski and Webster Atheta cheersae Klimaszewski Atheta circulicollis Lohse Atheta concessa Casey

Canadian distribution BC

United States distribution AK, CA

NB, NF, ON BC

CA

BC LB, NB, NF MB, NB, NF, ON, QC

AL, IL, IN, KY, MO, NH, NY, TN, VA, WA, WV

BC [NPR], LB, MB, NB, NF, NS, ON, PE, QC, SK YT

CA, CO, IN, MA, ME, MI, NH, NY, PA, VT, WA WY

MB MB NB, NF, NS, PE, QC

KS KS

NB, ON, QC ON, BC [NPR]

CA, IA, KS, MA, MS, NC, NH, NJ, NY, PA, TX

NB LB, NF, NT, YT NF, NS, ON NB, NF, NS, ON, QC

AK CA, WA AL, FL, GA, IA, IL, IN, KS, KY, LA, MO, MS, NC, NY, OH, TN, VA, WI, WV

LB, NF LB, NF AB, NF, NU, ON BC BC [NPR], MB, NB, NS, ON, QC, YT NB

CA

LB, NB, NF, ON, QC, YT YT LB, NB, NF, ON, YT BC [NPR], MB, LB, NB, NF, ON, QC, YT LB, YT BC, NB, NF, NS, QC, SK NB

AK

AK, MA, WV

BC LB, NB, NF, ON, QC BC (continued)

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Table A.2 (continued) Taxonomic name Atheta copleyi Klimaszewski, sp. n. Atheta cranberriensis Klimaszewski and Webster Atheta crenuliventris Bernhauer Atheta cryptica (Lohse) Atheta curtipenis Klimaszewski and Webster Atheta curvipennis Klimaszewski and Langor Atheta dadopora C.G. Thomson
Atheta districta Casey Atheta fanatica Casey Atheta festinans (Erichson) Atheta finita Moore and Legner Atheta formalis (Casey) Atheta formicaensis Klimaszewski and Webster Atheta frosti Bernhauer Atheta gelida J.R. Sahlberg
Atheta giguereae Klimaszewski and Webster Atheta godini Klimaszewski, sp. n. Atheta graminicola (Gravenhorst)
Atheta hampshirensis Bernhauer Atheta holmbergi Bernhauer Atheta newfounlandica Klimaszewski and Langor Atheta irrupta (Casey) Atheta keeni Casey Atheta klagesi Bernhauer Atheta larsonae Klimaszewski and Larson Atheta lindrothides Newton Atheta longiclava (Casey) Atheta longicornis (Gravenhorst){ Atheta lucifera Bernhauer Atheta macesi Klimaszewski and Webster Atheta makepeacei Klimaszewski and Webster Atheta manitobae Klimaszewski and Godin

Canadian distribution BC

United States distribution

NB LB, MB, NB, NF, ON, QC, SK BC, NF, QC, YT NB

ME

LB, NF AB, BC, LB, MB, NB, NF, NS, ON, PE, QC, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, QC, SK AB, BC, LB, NB, NS, QC, SK, YT ON, QC AB

AK, NY, PA, RI

NV AZ, CT, IA, IN, KS, MA, ME, MI, NH, NY, PA, RI CA

BC NB BC, LB, MB, NB, NS, ON, QC, SK MB, NT, QC, YT NB, NF, NS, ON

MA, NC, NH, NY, PA, RI, VT, WV AK

BC AB, BC, LB, MB, NB, NF, NT, ON, QC, SK, YT AB, BC, LB, MB, NB, NF, NS, ON, QC BC NF BC BC AB, MB, NB, NF, QC SK NF ON BC, MB, NB, NF, NS, QC, SK BC [NPR] NB

AK, OR AK, CA, NC, NH, NY, PA, RI AK

AK, OR ME, PA

RI CA, MN, OR CA, WA

NB MB (continued)

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Table A.2 (continued) Taxonomic name Atheta martini Lohse Atheta mcalpinei Klimaszewski and Webster Atheta meduxnekeagensis Webster and Klimaszewski Atheta metlakatlana Bernhauer Atheta microelytrata Klimaszewski and Godin Atheta modesta (Melsheimer) Atheta munsteri Bernhauer Atheta nearctica (Lohse) Atheta nescia (Casey) Atheta newfoundlandica (Klimaszewski and Langor) Atheta nigra (Kraatz){ Atheta novaescotiae Klimaszewski and Majka Atheta particula (Casey) Atheta pecki Klimaszewski and Langor Atheta pennsylvanica Bernhauer Atheta petitcapensis Klimaszewski and Webster Atheta picipennis (Mannerheim) Atheta platonoffi Brundin
Atheta pratensis (Mäklin) Atheta prudhoensis (Lohse) Atheta pseudocrenuliventris Klimaszewski Atheta pseudodistricta Klimaszewski and Langor Atheta pseudoklagesi Klimaszewski and Webster Atheta pseudometlakatlana Klimaszewski and Godin Atheta pseudomodesta Klimaszewski Atheta pseudopittionii Klimaszewski and Larson Atheta pseudoschistoglossa Klimaszewski and Webster Atheta pseudosubtilis Klimaszewski and Langor Atheta pseudovestita Klimaszewski and Langor Atheta quebecensis Webster and Klimaszewski

Canadian distribution YT NB

United States distribution

NB, BC BC

OR

YT AB, MB, NB, NS, ON, QC BC [NPR], MB, NT, YT LB, NT, YT BC NB, NF, TN SK NF, NB, NS. FRANCE: St. Pierre et Miquelon NB, NS, ON, QC LB

CT, DC, ME, MI, NY, PA, RI, VT, WV AK AK

NY, RI

LB, MB, NB, NF, NS, ON, QC NB

IN, MA, MN, NY, PA, RI, VA, VT

BC

AK, CA, WA

AB, BC, LB, NB, NF, NS, ON, SK, YT YT BC, LB, NB, NF, NS, ON, QC, SK, YT LB, NB, NF, NS, ON, YT

AK AK AK, NY, VT

NF AB, BC [NPR], MB, NB, NF, QC, SK, YT BC [NPR], SK, YT NB, NF, NS, ON, QC SK BC, NB, SK

AK

AB, LB, NB, NF NF QC (continued)

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Table A.2 (continued) Taxonomic name Atheta recondita (Erichson) Atheta regissalmonis (Lohse) Atheta relicta (Casey) Atheta remulsa Casey Atheta richardsoni Klimaszewski and Larson Atheta ringi Klimaszewski Atheta ripariides Newton Atheta rurigena Casey Atheta savardae Klimaszewski and Majka Atheta sculptisoma Klimaszewski and Langor Atheta smetanai (Lohse) Atheta sparreschneideri Munster
Atheta spermathecorum Klimaszewski and Larson Atheta sphagnicola Klimaszewski and Webster Atheta stercoris Fenyes Atheta strigosula Casey Atheta subsinuata (Erichson){ Atheta subtilis (Scriba){ Atheta surgens (Casey) Atheta terranovae Klimaszewski and Langor Atheta trinotata (Kraatz){ Atheta ventricosa Bernhauer Atheta vestita (Gravenhorst){ Atheta vincenti Webster and Klimaszewski Atheta wheelerae Klimaszewski, sp. n. Atheta whitehorsensis Klimaszewski and Godin Atheta winchesteri Klimaszewski, sp. n. Boreophilia caseyi Lohse Boreophilia davidgei Klimaszewski and Godin Boreophilia eremita (Rye)
Boreophilia fusca (C.R. Sahlberg)
Boreophilia hyperborea (Brundin)
Boreophilia islandica (Kraatz)


Canadian distribution SK LB, NF, QC BC AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT SK BC BC [NPR], MB, SK, YT BC NB, NF, NS, ON, QC

United States distribution CA, NH, NV, PA AK AK, NY

AK

NF LB, MB, NT, QC, YT YT

AK AK

SK NB BC BC, LB,MB, NB, NF, ON, QC, SK, YT YT, SK LB, NB, NF, QC BC BC [NPR], LB, MB, NB, NF, ON, QC, SK, YT NF AB, BC, LB, NB, NF, NS, ON, QC, SK, YT NB, NF, NS NB

AK, NY

AK [NSR]

AK, CA, DC, IN, MA, NC, NH, NJ, NY, PA, VT, WA.

BC NB, YT BC MB, NU, NT, YT AB, YT

AK

LB, MB, NB NT

AK AK

NT, NU

AK

AB, NF, NT, NU, YT

AK (continued)

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Table A.2 (continued) Taxonomic name Boreophilia nearctica Lohse Boreophilia neoinsecuta Klimaszewski Boreophilia nomensis (Casey) Boreophilia ovalis Klimaszewski and Langor Boreophilia subplana (J. Sahlberg)
Boreophilia vega (Fenyes)
Boreophilia venti Lohse Boreostiba campbelliana Lohse Boreostiba frigida (J.R. Sahlberg)
Boreostiba lagunae Lohse Boreostiba parvipennis (Bernhauer) Boreostiba pseudolaticollis Klimaszewski and Godin, sp. n. Boreostiba sibirica (Mäklin)
Boreostiba websteri Klimaszewski and Langor Callicerus obscurus Gravenhorst{ Callicerus rigidicornis (Erichson){ Clusiota grandipenis Klimaszewski and Webster Clusiota impressicollis (Bernhauer) Dalotia coriaria (Kraatz){ Dinaraea angustula (Gyllenhal){ Dinaraea backusensis Klimaszewski and Brunke Dinaraea bicornis Klimaszewski and Webster Dinaraea borealis Lohse Dinaraea curtipenis Klimaszewski and Webster Dinaraea inexpectata Klimaszewski, sp. n. Dinaraea longipenis Klimaszewski and Webster Dinaraea pacei Klimaszewski and Langor Dinaraea piceana Klimaszewski and Jacobs

Canadian distribution AB, LB, MB, NF, QC, YT MB, YT

United States distribution AK AK

BC, YT

AK

NF NT, NU

AK, NH

YT YT YT

AK AK

LB, NT, NU, QC, YT

AK

YT AB, BC, LB, NF, NT, QC, YT BC

LB, NT, NU, QC, YT LB, NB

AK, NH AK [NSR]

AK, NH

ON NF, ON NB BC, NB, NF, ON

CA, WA

AB, BC, NB, NS, ON, QC AB, BC [NPR], LB, NB, NF, NS, ON, PE, QC, SK, YT NB, ON

CA, FL, LA, MA, NJ, NY CA, NY, PA

MA

NB, ON NB, ON, QC NB BC NB BC, LB, NB, NF, QC, SK, YT QC

AK

(continued)

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Table A.2 (continued) Taxonomic name Dinaraea planaris (Mäklin) [status uncertain] Dinaraea quadricornis Klimaszewski and Webster Dinaraea subdepressa (Bernhauer) Dinaraea worki Klimaszewski and Jacobs Dochmonota langori Klimaszewski and Larson Dochmonota rudiventris (Eppelsheim){ Dochmonota simulans Klimaszewski and Larson Dochmonota websteri Klimaszewski and Larson Earota dentata (Bernhauer)

Emmelostiba microptera (Lohse) Geostiba appalachigena Gusarov Geostiba circellaris (Gravenhorst){ Geostiba horwoodae Klimaszewski and Godin, sp. n. Goniusa alperti Kistner Goniusa carrorum Maruyama and Klimaszewski Goniusa caseyi Gusarov Hydrosmecta borealis (Klimaszewski and Langor) Hydrosmecta caduca Casey Hydrosmecta canadensis Webster and Klimaszewski Hydrosmecta dulcis Casey Hydrosmecta minutissimoides Webster and Klimaszewski Hydrosmecta pseudodiosica Lohse Liogluta atriventris (Casey) Liogluta castoris Klimaszewski and Webster Liogluta gigantea Klimaszewski and Langor Liogluta granulosa Lohse Liogluta intermedia Klimaszewski and Langor

Canadian distribution YT

United States distribution AK

NB BC [NPR], NB, ON [NPR]

NH

AB, QC SK BC [NPR], NB, NF, NT QC, YT SK

ID, MA

SK AB, BC, MB, NB, NF, NS, ON, QC, SK, YT YT

AK, AL, CA, CO, DC, IA, ID, IL, KS, IN, MA, ME, MD, MT, NC, NH, NJ, NM, NV, NY, OH, OR, PA, VA, WA AK

NB, NS, QC

MD, ME, MA, NH, NJ, NY, PA, RI, VA, WI, WV

NB, NF BC

AB AB

WA

AB, BC, MB NF

DC, MA, NH, NJ, NY, TX

NB NB

NY

NB NB

WI

NB, ON, YT BC NB, NS, ON, QC LB, ON, QC YT LB, NF, NS, ON, QC

AK NH (continued)

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Table A.2 (continued) Taxonomic name Liogluta microgranulosa Klimaszewski and Webster Liogluta nigropolita (Bernhauer) Liogluta nitens (Mäklin) Liogluta pseudocastoris Klimaszewski and Webster Liogluta quadricollis (Casey) Liogluta scudderi Klimaszewski, sp. n. Liogluta terminalis (Casey) Liogluta trapezicollis Lohse Liogluta wickhami (Casey) Lypoglossa angularis (Mäklin) Lypoglossa franclemonti Hoebeke Lypoglossa manitobae Gusarov Meronera venustula (Erichson) Mocyta amblystegii (Brundin)
Mocyta breviuscula (Mäklin) Mocyta discreta (Casey) Mocyta fungi (Gravenhorst){

Mocyta luteola (Erichson) Mocyta sphagnorum Klimaszewski and Webster Nehemitropia lividipennis (Mannerheim){ Ousipalia pacifica Casey Paragoniusa myrmicae Maruyama and Klimaszewski Paraleptonia pacei Klimaszewski Pelioptera americana (Gusarov) Pelioptera robusta (Gusarov) Pelioptera thujae (Klimaszewski and Webster) Philhygra angusticauda (Bernhauer)

Canadian distribution NB

United States distribution

LB, NF, NT, NU, QC, YT

NH

AB, BC, YT NB

AK, OR, WA

BC BC AB, BC, LB, NB, NF, NS, ON, QC, YT BC [NPR], YT BC AB, BC, LB, NB, NF, QC, YT AB, BC [NPR], MB, NB, NF, NS, NT, ON, QC, SK, YT MB NB, ON, QC MB, NT, NU, YT AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT BC [NPR], , MB, ON, QC, SK AB, BC, LB, MB, NB, NF, NS, NU, ON, PE, QC, SK, YT MB, NB, NF, ON, QC

MT, NH AK AK, AZ, CA, CO, MA, ME, MT, NH, NM, OR, WA ME, NH, NY, VT

Widespread in eastern USA, westward to CO, TX and UT AK AK, CA, NV, OR IA, MN, WA [NSR] AK, CA, ME, MA, MN, NY, OR, RI, WA

DC, IA, IN, MA, MI, MN, MO, MS, NC, NY, PA, RI, WI

NB, NF, ON, QC, SK BC [NPR], NB, NF, NS, ON, PE, QC, SK BC, SK [NPR] AB, BC, LB, NB

CA, LA, MA, MI, MN, NE, NJ, NM, NY, PA, TX, VT CA, WA [NSR]

BC QC

CT, IA, IN, MO, NJ, NY, OH, PA, WI

BC NB

OR

BC, NB

AK, NH (continued)

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Table A.2 (continued) Taxonomic name Philhygra atypicalis Klimaszewski and Webster Philhygra botanicarum (Muona)
Philhygra charlottae Klimaszewski, sp. n. Philhygra clemens (Casey) Philhygra criddlei (Casey) Philhygra falcifera Lohse Philhygra hygrotopora (Kraatz){ Philhygra jarmilae Klimaszewski and Langor Philhygra junii Lohse Philhygra laevicollis (Mäklin) Philhygra larsoni Klimaszewski and Langor Philhygra leechi Lohse Philhygra luridipennis (Mannerheim){ Philhygra malleoides Lohse Philhygra manitobae (Casey) Philhygra palustris (Kiessenwetter){ Philhygra proterminalis (Bernhauer) Philhygra pseudoboreostiba Lohse Philhygra pseudolarsoni Klimaszewski and Godin Philhygra pseudopolaris Klimaszewski and Langor Philhygra pseudoterminalis Klimaszewski and Langor Philhygra ripicoloides Lohse Philhygra rostrifera Lohse Philhygra satanas (Bernhauer) Philhygra sinuipennis Klimaszewski and Langor Philhygra subpolaris (Fenyes) Philhygra terrestris Klimaszewski and Godin Philhygra terrivaga Klimaszewski, sp. n. Philhygra varula (Casey) Philhygra wisconsinica (Casey) Pontomalota opaca (LeConte)

Canadian distribution NB

United States distribution

BC, LB, NB, NF, NS, ON, SK, YT BC BC, MB, NB, NS, ON, QC, YT MB MB, SK NB, NF

OH, WI

MB, NB, NF, ON, SK, YT YT BC, NB, NS, ON

AK, WA

NB, NF MB, NT, YT NB, NF, ON

NY?

MB, NF, NT, QC, YT MB MB

AK IA, ME, NH, NY, PA, WI

NB, ON

CO, PA, RI

YT NB, YT MB, NF, NT, QC, YT

AK, CA

NF NF, NT, SK, YT LB, NT, SK AB

AK CA

NB, NF, SK, YT AB, SK BC [NPR], NB, SK, YT

AZ, NM AK [NSR]

BC

OR, WA

LB, MB, NB, NF, QC MB

WI

BC

AK, CA, OR, WA (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

611

Table A.2 (continued) Taxonomic name Psammostiba comparabilis (Mäklin) Psammostiba kenaii Gusarov Schistoglossa blatchleyi (Bernhauer and Scheerpeltz) Schistoglossa brunswickensis Klimaszewski and Webster Schistoglossa campbelli Klimaszewski Schistoglossa carexiana Klimaszewski Schistoglossa charlottae Klimaszewski Schistoglossa hamshirensis Klimaszewski Schistoglossa pelletieri Klimaszewski and Webster Schistoglossa pseudocampbelli Klimaszewski and Webster Schistoglossa sphagnorum Klimaszewski and Webster Seeversiella globicollis (Bernhauer) Stethusa klimschi (Bernhauer) Stethusa spuriella (Casey) Strigota ambigua (Erichson) Strigota obscurata Klimaszewski and Brunke Strophogastra pencillata Fenyes Tarphiota fucicola (Mäklin) Tarphiota geniculata (Mäklin) Thamiaraea brittoni (Casey) Thamiaraea claydeni Klimaszewski and Webster Thamiaraea corverae Klimaszewski and Webster Thinusa fletcheri Casey Thinusa maritima (Casey) Tomoglossa decora (Casey) Trichiusa columbica Casey Trichiusa compacta Casey Trichiusa hirsuta Casey Trichiusa pilosa Casey Trichiusa pseudopostica Klimaszewski and Langor Trichiusa robustula Casey Tribe Autaliini C.G. Thomson Autalia puncticollis Sharp{

Canadian distribution BC

United States distribution AK, CA

BC MB, NB, NT, ON, QC, SK, YT NB, ON, QC

AK, CA? AK, IN

AB, BC BC, SK BC AB, BC [NPR], QC

NH

NB NB

NB AB, BC, MB, NB, NF, NS, ON, QC, SK ON NF, ON, QC BC [NPR], LB, NB, NF, NS, ON, PE, QC, SK, YT BC [NPR], NB, ON, QC, SK AB, MB, NB, NS, ON, QC

AZ, CO, ID, MI, MN, MT, NH, NM, SD, WI IN, LA, MS DE, FL, GA, IN, MO, NJ, NY, OH, PA CA, CO, CT, IA, KS, MA, MO, NC, NJ, NM, NY, NV, TX

BC BC

AK, CA AK, CA

NB, ON, QC NB

CT, KS, MS, NC, NJ, NY, OH, PA, WI

NB BC BC NB BC ON NB, ON AB, BC, NB, NS, ON NF

AK, CA AK, CA, OR, WA AR, MS, PA, VT DC, NY?, OH VA ID, IN, KS, OH, RI

NB, ON

IA

BC

CA, WA (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.2 (continued) Taxonomic name Canadian distribution Autalia rivularis AB, BC, LB, NB, NF, NS, (Gravenhorst){ ON, QC, SK Autalia truncatula Casey BC Tribe Boreocyphini Klimaszewski and Langor Boreocypha websteri LB, NB, PE Klimaszewski and Langor Tribe Diglottini Jacobson Diglotta mersa (Haliday){ NB Bryothinusa catalinae Casey BC [NPR] Tribe Falagriini Mulsant and Rey Aleodorus bilobatus Say ON, QC

United States distribution CA, ME, MI, NH, NJ, NY, OR, PA, VT AK, CA

CA

Aleodorus intricatus (Casey) Aleodorus scutellaris (LeConte) Borboropora quadriceps (LeConte) Bryobiota bicolor (Casey) Cordalia obscura (Gravenhorst){ Falagria caesa Erichson{ Falagria dissecta Erichson

AB, SK ON

CT, DC, GA, IA, IL, IN, KY, MA, MD, MI, MO, PA, NC, NJ, NY, OH, TN, VA, WV AZ, CA, CO, MT, NM IL, LA, MA, MD, MI, MS, NH, NY, TN

ON

DC, IL, LA, MD, NC, NY, PA, SC, TN, VA, WV

BC BC, NB, NS, ON, QC

Myrmecocephalus arizonicus (Casey) Myrmecocephalus cingulatus (LeConte)

AB, BC, SK

CA CA, CO, CT, IL, MA, MD, MI, NH, NJ, NY, OH, PA, WA IL, MA, MD, NJ, NY, UT, VA AL, AZ, CA, CO, CT, DC, FL, GA, IA, IL, IN, KS, KY, LA, ME, MA, MD, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NY, OH, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY AZ, CO, NE, NM, SD, UT, WA

Myrmecocephalus concinnus (Erichson){ Myrmecocephalus gatineauensis Hoebeke Myrmecopora vaga (LeConte) Tribe Gymnusini Heer Deinopsis canadensis Klimaszewski Deinopsis harringtoni Casey

ON, QC

Deinopsis illinoisensis Klimaszewski Deinopsis rhadina Klimaszewski Gymnusa atra Casey
Gymnusa brevicollis (Paykull) { Gymnusa campbelli Klimaszewski Gymnusa grandiceps Casey

AB, NB, ON, QC, SK AB, BC, MB, NB, NS, ON, QC, SK

NS, ON

NB, QC, ON

AL, AR, DC, FL, IA, IL, IN, KS, KY, LA, MD, MI, MO, MS, NC, NH, NJ, NY, OH, PA, TN, VA, WV, WI AL, AZ, CA, FL, IL, IA, KS, LA, MA, MD, NC, NY, OH, TX, VA, WA NH, NJ, NY, PA

NB, NS

« Lake Superior region»

LB, NB, NF, ON LB, MB, NB, NF, NS, ON, QC ON

AR, CA, MA, ME, MI, NH, WI CT, FL, IL, KY, LA, MA, MI, MS, OK, PA, TX

NB, ON, QC

AL, IA, MA

AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT LB, NF, QC

AK, IL, IA, MA, ME, MI, MN, NH, NY. France: St. Pierre et Miquelon MA, ME. France: St. Pierre et Miquelon

MB, NB, NF, NT, ON, QC, SK, YT BC [NPR], LB, MB, NB, NF, NS, ON, QC

AK, MA New England states, IL, MA, MD, ME, MI, NH, NY, RI (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

613

Table A.2 (continued) Taxonomic name Gymnusa konopackii Klimaszewski Gymnusa lindrothi Klimaszewski and Langor Gymnusa pseudovariegata Klimaszewski Gymnusa smetanai Klimaszewski
Tribe Homalotini Heer Agaricochara aldersonae (Klimaszewski and Webster) Agaricochara pulchra Klimaszewski and Larson Agaricomorpha websteri Klimaszewski and Brunke Anomognathus americanus (Casey) Anomognathus athabascensis Klimaszewski, Hammond, Langor Cyphea curtula (Erichson){ Cyphea wallisi Fenyes Encephalus americanus Seevers Eumicrota corruscula (Erichson)

Canadian distribution MB, NT, NU, YT

United States distribution AK

LB, NF, NB AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, YT MB, NF, NT, ON, YT

AK, CA, MA, OR, WA AK

NB SK NB, NS, ON, QC NB

NY

AB

AB, NB, QC MB AB, BC NB, ON, QC

IL, MT, NM

Eumicrota socia (Erichson)

NB, NS, ON, PE, QC

Gyrophaena affinis Mannerheim{ Gyrophaena antennalis Casey Gyrophaena brevicollis Seevers Gyrophaena californica (Casey) Gyrophaena caseyi Seevers Gyrophaena chippewa Seevers Gyrophaena criddlei Casey Gyrophaena dybasi Seevers Gyrophaena egena Casey Gyrophaena flavicornis Melsheimer Gyrophaena fuscicollis Casey Gyrophaena gaudens Casey Gyrophaena gilvicollis Casey Gyrophaena gracilis Seevers Gyrophaena illiana Seevers Gyrophaena insolens Casey Gyrophaena involuta Casey

BC, MB, NB, NF, NS, ON, QC, SK LB, NB, NF, ON NB, ON

AL, CT, DC, FL, GA, IL, IN, IA, KS, KY, LA, MA, MI, MO, NJ, NY, OH, PA, RI, SC, TN, TX, VA, WI, WV AR, DC, FL, IL, IN, KS, KY, LA, MA, ME, MD, MI, MO, NY, NC, OH, PA, SC, TN, TX, VA, WI, WV AZ, DC, IL, IN, IA, KY, MA, ME, MI, MN, MO, NC, NH, NJ, NM, NY, OH, PA, TN, WA, WI, WV MA, NC, NY, TN IN, IL, MO, NC

BC

CA, CO, NM

MB, NB, ON, QC NB, NF

MI, NC, NY, PA MI, NC, WI

LB, MB, NB, ON, SK, YT? NB, ON ON, QC MB, NB, NS, ON, QC NB, ON AB, MB, ON, PE MB, NB, ON NB NB LB, MB, NB, NF, ON, SK LB, NB

IL, IN, MO, NC, WI MA, NC, PA, RI CT, DC, IL, IN, KY, MA, MD, ME, MI, MO, NC, NH, NJ, NY, PA, RI, TN, VA, VT, WI, WV DC, IL, MO, NY, PA, WI IL, IN, MA, MI, PA, WI DC, IL, IN, KS, MI, NC, NY, PA, VA, WV WI IL, IN, WI MI MA, ME, NH, NY, WI (continued)

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Table A.2 (continued) Taxonomic name Gyrophaena keeni Casey Gyrophaena laetula Casey Gyrophaena lobata Casey Gyrophaena meduxnekeagensis Klimaszewski and Webster Gyrophaena michigana Seevers Gyrophaena modesta Casey Gyrophaena nana (Paykull)
Gyrophaena nanoides Seevers Gyrophaena neonana Seevers Gyrophaena pseudocriddlei Klimaszewski and Webster Gyrophaena sculptipennis Casey Gyrophaena stroheckeri Seevers Gyrophaena subnitens Casey Gyrophaena uteana Casey Gyrophaena vitrina Casey

Canadian distribution AB, BC, LB, MB, NB, NF, NS, ON, QC, YT MB, NB, NF NB, SK NB, ON, QC

United States distribution FL, MA, ME, MT, NH, NY, OR, TN, WA, WY DC, IL, IN, KY, MA, NY, PA, TN, VA, WI DC, IL, IN, KY, MI, NY, WI

NB

IL, MI, WI

AB, MB, NB, NF, NS, ON AB, BC, MB, NB, NF, ON, SK, YT MB, NB, NF, ON, QC NB, NF, ON, YT? NB

IL, IN, MI, MN, NH, NY AK, CA, MA, ME, MI, MT, WI, WY DC, IA, IN, KS, MA, NC, TN, VA, WI IN, NC, PA, WI

AB, NB, NS, ON, QC

MA, NH, NY, WI

ON

IN, NC, WI

MB, NB, NS, ON, SK AB, BC, NB, ON, QC, SK MB, NB, ON, PE, QC

IL, KS, ME, MI, MN, MO, NY, WI CA, CO, UT AL, IL, IN, KY, ME, MI, NC, NH, NY, PA, TN, WV, WI IL, WI

Gyrophaena wisconsinica Seevers Homalota plana (Gyllenhal){

AB, NB, QC

Leptusa brevicollis Casey Leptusa canonica Casey Leptusa carolinensis Pace Leptusa cribratula (Casey) Leptusa elegans Blatchley Leptusa gatineauensis Klimaszewski and Pelletier Leptusa jucunda Klimaszewski and Majka Leptusa opaca Casey Leptusa pseudopaca Klimaszewski and Majka Leptusomorpha claudiae Klimaszewski, sp. n. Neotobia alberta Ashe Phanerota fasciata (Say)

NB, NF, NS, ON, PE, QC NB, NF, NS, ON, QC NB, NS, ON, QC ON, QC ON, QC AB, BC, NB, NF, NS, ON, QC, SK NB, NS, ON, QC

Phymatura blanchardi (Casey) Pleurotobia bourdonae Klimaszewski and Webster Pleurotobia brunswickensis Klimaszewski and Webster

AB, NB, ON

AB, MB, NB, NF, NS, ON

NB, NF, NS, ON PE, QC NS, QC

AK, AZ, CA, CO, FL, IA, ID, IN, MT, NY, OH, PA, TX, WA MA, NC, NH, NY, PA, VA, VT IA, IN, MS, OH, PA, TX NC, TN AL, FL, OH, PA, TN, VA AR, IN, MN, NJ, NY, PA, VT, VA

AR, GA, NC, NY, PA, RI, TN, WI NH, WV

BC AB, MB, NB, ON, QC ON

ME, NH, WI AR, DC, FL, GA, IA, IL, IN, KY, KS, LA, MD, MI, MO, MS, NC, NJ, NY, OH, PA, RI, TN, TX, VA, WI IA, IN, MO, NY

NB, QC NB (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

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Table A.2 (continued) Taxonomic name Silusa alternans Sachse Silusa californica (Bernhauer) Silusa densa Fenyes Silusa langori Klimaszewski Silusa opaca Fenyes Silusa prettyae Klimaszewski and Langor Silusa vesperis Casey Silusida marginella (Casey) Stictalia brevicornis Casey Stictalia californica (Casey) Stictalia carlottae Casey Stictalia kranabetteri Klimaszewski and Godin, sp. n. Thecturota capito Casey Thecturota tenuissima Casey Tribe Hoplandriini Casey Hoplandria klimaszewskii Génier Hoplandria laevicollis (Notman) Hoplandria laeviventris Casey

Canadian distribution NB, NS, ON, PE, QC AB, BC, LB, NB, NF, NS, ON, PE, QC, SK, YT AB, LB, NB, NF AB, NB BC NF

CA CA

BC NB, NF, NS, ON, PE, QC BC BC BC BC

CA, OR, WA KS, NY, PA, TN. CA CA, AK

ON ON, QC

IA, IN, TX, VA RI

ON, QC

AR, DC, FL, IL, MD, NC, NJ, NY, VA, WV

NB, ON, QC

DC, FL, GA, LA, NC, NJ, NY, VA

ON

AL, AR, CT, DC, GA, IL, IN, KY, LA, MA, MD, NJ, NY, NC, OH, PA, TN, TX, VA, WV AR, AZ, CA, DC, FL, GA, IA, IL, IN, KS, LA, MD, MI, MO, NC, NJ, NM, NY, OH, OK, PA, SD, TN, TX, VA, WV DC, GA, IA, IL, IN, KS, LA, NC, NE, NJ, PA, TN, VA

Hoplandria lateralis (Melsheimer)

NB, ON, QC

Platandria carolinae Casey

ON

Tribe Hypocyphtini Laporte Cypha crotchii (Horn) Cypha inexpectata Klimaszewski and Godin Holobus vancouveri Klimaszewski Oligota chrysopyga Kraatz{ Oligota inflata (Mannerheim) { Oligota parva Kraatz{ Oligota polyporicola Klimaszewski and Webster Oligota pusillima (Gravenhorst){ Oligota sevogle Klimaszewski and Webster Tribe Liparocephalini Fenyes Amblopusa brevipes Casey Diaulota densissima Casey Liparocephalus brevipennis Mäklin

United States distribution GA, NH, NY AK, CA, MN

AB, BC, SK ON, SK, YT

ID, OR, WA

BC NB SK

FL

BC [NPR], NB, PE NB

CA, MA, MO, NV, TX

NB

MA, NY

NB

BC BC BC

AK, CA AK, CA, OR, WA AK (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.2 (continued) Taxonomic name Liparocephalus cordicollis LeConte Paramblopusa borealis (Casey) Tribe Lomechusini Fleming Drusilla canaliculata (Fabricius){ Myrmedonota aidani Maruyama and Klimaszewski Myrmoecia canadensis Maruyama and Klimaszewski Myrmoecia lauta (Casey) Myrmoecia lugubris (Casey) Pella caliginosa (Casey) Pella carolinae (Casey) Pella criddlei (Casey) Pella gesneri Klimaszewski Pella glooscapi Klimaszewski and Majka Pella loricata (Casey) Pella schmitti (Hamilton) Xenodusa cava (LeConte)

Canadian distribution BC

United States distribution AK, CA, OR, WA

BC

AK, CA, OR, WA

NB, NF, NS, ON, PE, QC

AK, KY, MA, NY, PA, VT

ON

OH

AB ON, QC BC AB ON AB, MB, QC AB, MB, NB, ON NS

IN, MA, ME, MI, NH, NY, WI CO, WA CA, IN, NY NC

NS, ON ON, QC ON

OH MA, PA AR, CO, CT, IA, IL, IN, KS, KY, MA, ME, MI, MN, MO, NE, NH, NJ, NY, OH, OK, PA, RI, VA, WV, WI AZ, CA, CO, CT, IL, IA, MA, MD, MI, MN, MT, ND, NE, NH, NM, NV, NY, OR, RI, SC, UT, VT, WA, WI, WY MI, MO, NH, NY, OR

Xenodusa reflexa (Walker)

AB, BC, MB, NB, NS, ON, QC, SK

Zyras obliquus (Casey)

AB, BC, LB, MB, NB, NF, NS, ON, QC, SK ON

Zyras planifer (Casey) Tribe Myllaenini Ganglbauer Myllaena arcana Casey AB, LB, MB, NB, NF, NS, ON, QC, SK Myllaena audax Casey BC, LB, NB, NF, NT, ON, QC Myllaena cuneata Notman NS, ON Myllaena decreta Casey AB, BC Myllaena fenyesi Bernhauer BC Myllaena insomnis Casey AB, BC, LB, MB, NB, NF, NS, NT, ON, QC, SK, YT Myllaena kaskaskia NB Klimaszewski Myllaena ludificans Casey NB, ON, QC Myllaena potawatomi ON Klimaszewski Myllaena procidua Casey NB, NF, QC Myllaena vulpina Bernhauer NB, NS, ON Tribe Oxypodini C.G. Thomson Alfocalea montana AB, BC Klimaszewski Alisalia elongata NB Klimaszewski and Webster

DC, IN, NC AL, FL, IA, IL, MA, ME, NH, NJ, TN, WV AK, CA, GA, IL, LA, MA, NH, NJ, NY, OR, RI, UT, WA AR, FL, GA, IL, LA, MA, MD, NH, OK, TN, VA AZ, CA, NM, OR, UT, WA AZ, CA, NV, OR, UT, WA AK, ID, MA, MI, MN, MT, NY, WI GA, IL, OK, VA AR, MA, OK, RI, WI AL, AZ, CA, FL, GA, IL, IN, MA, OK, TX, VA, WI FL, MA, MD, NY, OK, VA AR, KY, PA, WV

(continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

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Table A.2 (continued) Taxonomic name Alisalia minuta Klimaszewski and Webster Alisalia testacea Casey Apimela canadensis Klimaszewski and Webster Apimela fusciceps (Casey) Betocalea pacifica Klimaszewski Blepharhymenus brendeli (Casey) Blepharhymenus illectus (Casey) Calodera bennetti Klimaszewski, sp. n. Calodera caseyi Assing Calodera parviceps (Casey) Crataraea suturalis (Mannerheim){ Devia prospera (Erichson)
Dexiogyia angustiventris (Casey) Gennadota canadensis Casey Gnathusa alfacaribou Klimazewski and Langor Gnathusa caribou Lohse Gnathusa eva Fenyes Gnathusa tenuicornis Fenyes Gyronycha pseudoobscura Klimaszewski and Webster Hylota cryptica Klimaszewski and Webster Hylota ochracea Casey Ilyobates bennetti Donisthorpe { Megocalea lemieuxi Klimaszewski Meotica exilis (Knoch){ Meotica pallens (Redtenbacher){ Meotica pseudowinkleri Klimaszewski and Langor Metocalea lindgreni Klimaszewski Mniusa minutissima (Klimaszewski and Langor) Mniusa odelli (Klimaszewski and Webster) Mniusa yukonensis (Klimaszewski and Godin) Neoisoglossa agnita (Casey, 1911)

Canadian distribution NB

United States distribution

NB, ON, QC NB

NC

NB BC

NC, NY

NB, NS, ON, QC

IA

BC [NPR]

OR

BC

AK

NB NB, NS, ON, YT BC, LB, NB, NS, ON, QC, SK, YT AB, BC, LB, MB, NB, NT, ON, QC, SK, YT NB, ON, QC

MA, ME, PA RI CA, IA, IL, IN, MA, MO, PA, RI, SC, VA, VT AK, CA, CO, MA, MI, MN, MT, NH, NM, OR, SD, UT, WA, WI, WY CA, FL, IA, NY, RI

NB, NS, ON, QC LB

PA

NT, YT AB, BC, SK, YT AB, BC, NB?, YT NB

AK CA AK, CA, OR

AB, NB NB, NS, NT, ON, QC, SK NB, NF, NS, ON, QC

NY, VT

BC NS BC, NB, NS, ON, YT [NPR] NF

ME? NJ, RI

BC NB, NF NB, NS?, QC? BC, NB, NS, QC, YT AB, BC [NPR]

AZ (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.2 (continued) Taxonomic name Neothetalia canadiana Klimaszewski Neothetalia columbiana (Klimaszewski) Neothetalia nimia (Casey) Neothetalia pallitarsis (Kirby) Neothetalia robergei Klimaszewski, sp. n. Neothetalia smetanai Klimaszewski Ocalea vancouveri Casey Ocyusa asperula Casey Ocyusa canadensis Lohse Oxypoda amica Casey Oxypoda brachyptera (Stephens){ Oxypoda canadensis Klimaszewski Oxypoda chantali Klimaszewski Oxypoda convergens Casey Oxypoda demissa Casey Oxypoda domestica Klimaszewski and Larson Oxypoda famula Casey Oxypoda frigida Bernhauer Oxypoda gatosensis Bernhauer Oxypoda glenorae Casey Oxypoda gnara Casey Oxypoda hiemalis Casey Oxypoda impressa Casey Oxypoda inimica Casey Oxypoda irrasa Mäklin Oxypoda lacustris Casey Oxypoda leechi Lohse Oxypoda longicarinata Klimaszewski Oxypoda lucidula Casey Oxypoda manitobae Casey Oxypoda nigriceps Casey Oxypoda nimbata Casey

Canadian distribution BC, LB, NB, QC, YT

United States distribution AK

BC

AK

BC AB, BC

AK

BC BC BC NB BC [NPR], NB, NF, ON, SK, YT MB, NB, NS, ON, QC, YT NB, NF, NS, ON, QC

IA, MA, RI AK IA, IN, KS, KY, LA, MO, NC, NJ, NY, TN

AB, BC [NPR], LB, MB, NF, NT, ON, QC, YT NS, ON, QC

AK, NH

AB, BC [NPR], LB, MB, NB, NF, NS, ON, QC LB, NB, NF, NS, ON, QC, SK, YT SK

IA, MO, NY

BC AB, BC, LB, NB, NF, NS, NT, ON, QC, YT BC BC MB, NB, ON, QC AB, LB, NB, NF, NS, NT, ON, QC, YT BC NB, NF, NT, ON, QC AB, BC [NPR], SK, YT AB, BC, LB, MB, NB, NF, NT, ON, QC, SK, YT YT BC AB, LB, MB, NB, NF, NT, ON, QC, YT BC, MB, SK NB, NS BC

AK, NH CA

RI AK, NH

MA AK, OR AK, WA

AK AK, IA, MO, NH, NY CO RI, VA CA (continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

619

Table A.2 (continued) Taxonomic name Oxypoda opaca (Gravenhorst){ Oxypoda operta Sjöberg{ Oxypoda orbicollis Casey Oxypoda perexilis Casey Oxypoda pseudoconvergens Klimaszewski and Godin Oxypoda pseudolacustris Klimaszewski Oxypoda regressa Casey Oxypoda rubescans Casey Oxypoda smithi Klimaszewski Oxypoda stanleyi Klimaszewski & McLean Oxypoda sunpokeana Klimaszewski and Webster Oxypoda sylvia Casey (¼grandipennis (Casey)) Oxypoda vancouveri Klimaszewski Oxypoda vockerothi Klimaszewski Oxypoda volkeri Klimaszewski Oxypoda yukonensis Klimaszewski and Godin Parocalea nearctica Lohse Parocalea pseudobaicalica Lohse Phloeopora arctica Lohse

Canadian distribution BC, LB, NB, NF, NS, ON, QC AB, LB, NS, ON, QC, YT AB, BC [NPR], LB, NB, NS, ON, QC, SK, YT NS, ON, QC YT AB, MB, NB, NF, NS, ON, QC, SK BC ON BC

United States distribution NC, NY, PA, SC, TN, VT NH WI IA, IN, MO, MS, NC, TX

NH

NY

BC NB AB, BC, LB, MB, NB, NF, NS, ON, QC, SK, YT BC

AK, ME, NH

NB, ON NT YT NT, YT BC [NPR], NT, QC, YT

AK AK, NH

AB [NPR], BC [NPR], NT, ON, YT BC [NPR], NB, NF

Phloeopora canadensis Klimaszewski and Langor Phloeopora gilbertae NB, NT Klimaszewski and Webster Phloeopora oregona Casey BC [NPR], NB Tetralaucopora americana NB, NF, ON (Casey) Tetralaucopora fuliginosa LB, NB, ON, SK (Casey) Tribe Placusini Mulsant and Rey Euvira micmac Klimaszewski NB, NS, ON and Majka Placusa canadensis NS, ON, QC Klimaszewski Placusa despecta Erichson ON, QC Placusa incompleta Sjöberg{ AB, BC, NB, NF, NS, ON, QC, SK Placusa petulans Casey BC

OR NY, PA MA?, NC, PA

MI, OH, RI MI, OH, WV AL, GA, SC, MI, WV WA CA, WA (continued)

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Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

Table A.2 (continued) Taxonomic name Placusa pseudosuecica Klimaszewski Placusa tachyporoides (Waltl){ Placusa tacomae Casey

Canadian distribution AB, BC, ON, QC, SK

AB, BC, NB, NS, ON, QC, SK AB, BC, NB, NF, NS, NT, ON, QC, SK, YT Placusa vaga Casey AB, BC, NB, NS, NT, ON, QC, SK, YT Tribe Tachyusini C.G. Thomson Brachyusa americana BC (Fenyes) [unsettled status] Brachyusa helenae (Casey) BC [NPR], LB, NB, NF, NT, ON, SK, YT Brachyusa saskatchewani SK Klimaszewski and Larson Gnypeta ashei Klimaszewski MB, NT, NU, QC, YT Gnypeta atrolucens Casey LB, NB, NF, QC Gnypeta baranowskii BC Klimaszewski, sp. n. Gnypeta brevicornis Casey BC Gnypeta brincki Palm
NT, QC, YT Gnypeta caerulea (C.R. AB, BC, LB, MB, NF, NS, Sahlberg)
NB, NT, ON, PE, QC, SK, YT Gnypeta canadensis AB, ON Klimaszewski Gnypeta carbonaria AB, MB, NB, NF, NT, ON, (Mannerheim)
QC, SK Gnypeta crebrepunctata BC (Casey) Gnypeta dentata AB, NT, SK Klimaszewski Gnypeta groenlandicae Lohse MB, NU, YT Gnypeta helenae Casey AB, BC, ON Gnypeta lohsei Klimaszewski AB, BC Gnypeta manitobae Casey MB Gnypeta minuta Klimaszewski LB, NB, SK and Webster Gnypeta nigrella (LeConte) NB, NF, ON

Gnypeta saccharina Klimaszewski and Webster Gnypeta sellmani Brundin
Gnypeta punctatula Casey [¼uteana Casey] Paradilacra densissima (Bernhauer) Tachyusa americana Casey Tachyusa americanoides Paśnik

United States distribution

CA, MA AZ, CO, MA, ME, MN, NY, WA CA

AK, MT

AK NH, NY, VT

AK AK

AK CA, OR

AK, Greenland AZ, CA, CO, MT, NV, NM, ND, OR AK, WA

MA, MD, NJ, NY, PA, VT, IL þ WV were added without records by Klimaszewski, Webster & al. (2018: 287)

NB, SK LB, MB, NF, NT QC, SK, YT AB

AK

AB, BC

CA, MT, NV, ND, OR, UT

NB, ON, QC

CO, DC, IA, IL, IN, MA, MD, MN, NH, NJ, NY, OH, PA, VT, WI IL, MA, MT, NH, NY, PA, VT

AB, BC, MB, NB, NF, NS, NT, ON, QC

(continued)

Appendix: Checklists of BC and all Canadian Species and Their Distribution in Canada

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Table A.2 (continued) Taxonomic name Tachyusa cavicollis LeConte

Canadian distribution BC

United States distribution AR, FL, IL, IN, KS, LA, MA, ME, MD, NE, NC, NH, NJ, NM, OH, PA, SD, TX, VA, VT, WY

Tachyusa faceta Casey Tachyusa obsoleta Casey

SK BC, NB, SK

Tachyusa smetanai Paśnik Tribe Taxicerini Lohse Halobrecta flavipes C.G. Thomson{

QC, MB

CA, CO, DC, IL, IN, KS, MT, NE, NM, NV, OK, SD, UT, WI, WY AZ, CO, KY, NM, TX

NB

NY, VA


Holarctic species, {adventive species, {species of uncertain distribution status—adventive in Canada or Holarctic, species without marks are native, species in bold are recorded from BC; NPR new provincial record, NSR new state record. Tribes, genera and species are listed in alphabetical order

Index

A Acrimea acerba, 64, 108 Acrotona, 20, 351, 432 A. brachyoptera, 602 A. horwoodae, 602 A. manitobensis, 602 A. onthophilides, 58, 605 A. pseudopygmaea, 602 A. pseudopygmaeides, 602 A. recondita, 602 A. sequestralis, 602 A. smithi, 602 A. sphagnorum, 602 A. subpygmaea, 602 Adota, 12, 54, 63, 67, 355 A. maritima, 67, 356, 463, 594, 603 Adoxopisalia, 304 Agaricochara, 613 A. aldersonae, 613 A. pulchra, 613 Agaricomorpha, 56 Agaricomorpha websteri, 613 Alaobia, 67, 361, 362 Aleochara, 2, 10–12, 18, 19, 56, 76, 78, 97–99, 108, 114–115, 142, 145, 146, 181 A. acerba, 108, 131, 593, 601 A. angustula, 414 A. assiniboin, 64, 99, 116, 593, 601 A. bilineata, 57, 58, 64, 103, 123, 593, 601 A. bimaculata, 28, 64, 103, 104, 124, 593, 601 A. carmanah, 56, 64, 67, 110, 111, 133, 593, 601 A. castaneimarmotae, 111, 112, 601 A. castaneipennis, 64, 110–112, 134, 593, 601 A. caviceps, 601 A. coprochara, 64, 97, 98, 103, 115 A. curtidens, 64, 106–107, 128, 593, 601 A. curtula, 19, 34, 58, 64, 98–99, 117, 593, 601 A. daviesi, 601 A. elisabethae, 601 A. emplenota, 64, 98, 106, 115 A. fumata, 57, 64, 110–112, 136, 593, 601 A. gracilicornis, 64, 80, 99, 100, 118, 593, 601 A. inexspectata, 113, 601 A. lacertina, 64, 110, 112–113, 137, 593, 601

A. lanuginosa, 64, 80, 110, 113, 138, 593, 601 A. laramiensis, 64, 101–102, 120, 593, 601 A. lata, 58, 601 A. litoralis, 34, 64, 106–108, 129, 593, 601 A. moerens, 112 A. morion, 64, 108, 109, 132, 593, 601 A. nitidum, 104 A. obscura, 574 A. ocularis, 601 A. pacifica, 64, 106–108, 130, 593, 602 A. pallitarsis, 178 A. quadrata, 12, 56, 64, 110–112, 135, 593, 602 A. rubricalis, 64, 101–102, 121, 593, 602 A. rubripes, 602 A. sculptiventris, 602 A. sekanai, 602 A. shelleyae, 602 A. speculicollis, 102, 602 A. suffusa, 35, 64, 103, 105, 125, 593, 602 A. sulcicollis, 64, 103, 105, 126, 593, 602 A. tahoensis, 64, 99–101, 119, 594, 602 A. thoracica, 602 A. tinotus, 12, 64, 76, 97, 108 A. trisecta, 602 A. tristis, 602 A. verna, 64, 103, 105–106, 127, 594, 602 A. villosa, 64, 102–103, 122, 594, 602 ALEOCHARINAE, 1–3, 5, 6, 9–14, 22–27, 46, 53–60, 63–70, 75 ALEOCHARINI, 2, 5, 12, 18, 54, 56, 57, 63, 97, 601 Aleodorus, 10, 19 A. bilobatus, 612 A. intricatus, 612 A. scutellaris, 612 Alevonota, 54 A. gracilenta, 603 Alfocalea, 12, 63, 65, 143, 169 A. montana, 65, 83, 169, 215, 598, 616 Alisalia A. elongata, 616 A. minuta, 617 A. testacea, 617 Alnus, 147, 148, 388, 392, 424, 442 Aloconota, 63, 67, 352, 354, 357

# Her Majesty the Queen in Right of Canada 2020 J. Klimaszewski et al., Aleocharine Rove Beetles of British Columbia: A Hotspot of Canadian Biodiversity (Coleoptera, Staphylinidae), https://doi.org/10.1007/978-3-030-36174-7

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624 Aloconota (cont.) A. cambrica, 57, 67, 85, 358, 464, 594, 603 A. carmanah, 358, 465, 594, 603 A. neocambrica, 603 A. sulcifrons, 58, 603 Amarochara, 56, 76, 97 A. brevios, 602 A. duryi, 602 A. fenyesi, 602 A. inquilina, 602 Amblopusa, 2, 12, 26, 57, 63, 66, 76, 283–285 A. borealis, 285 A. brevipes, 66, 81, 284, 289, 598, 615 Amelanchier, 309, 392 Amischa, 20, 63, 67, 351, 352, 359 A. analis, 57, 67, 85, 359, 466, 594, 603 A. praelonga, 603 Anaduosternum brevipennis, 455 Anatheta planulicollis, 603 Anepsiota A. quadricollis, 425 A. terminalis, 426 A. wickhami, 428 Anomognathus, 613 A. americanus, 613 A. athabascensis, 613 Anthomyiidae, 104, 106 Apalonia, 586 Apimela, 56 A. canadensis, 617 A. fusciceps, 617 Ashella, 283 A. giulanii, 283, 578 Atemeles reflexus, 584 Atheta, 12, 18, 20, 21, 54, 63, 67, 68, 177, 178, 351, 353, 355, 360, 361, 363–365, 371, 376, 380, 382, 391, 394, 400, 401, 406, 411, 423, 432, 438, 441 A. acadiensis, 603 A. aemula, 603 A. alesi, 67, 371, 474, 594, 603 A. alphacrenuliventris, 603 A. altaica, 603 A. amicula, 58, 603 A. angusticauda, 440 A. annexa, 603 A. asperiola, 372 A. atramentaria, 603 A. avalon, 603 A. basicornis, 68, 407, 510 A. borealides, 603 A. brumalis, 68, 384, 487, 594, 603 A. brunswickensis, 68, 404–406, 507, 594, 603 A. bubo, 603 A. burwelli, 387, 603 A. cadeti, 603 A. campbelli, 67, 369, 370, 372, 603 A. capsularis, 68, 385, 386, 389, 489, 594, 603 A. caribou, 387, 603 A. carlottae, 68, 382

Index A. celata, 67, 367, 382, 472, 594, 603 A. chaetida, 67, 361, 366 A. chartersensis, 603 A. cheersae, 68, 393, 496, 594, 603 A. circulicollis, 603 A. columbica, 67, 68, 369, 370, 373 A. concessa, 68, 401, 504, 594, 603 A. copleyi, 68, 394, 395, 497, 594, 604 A. cranberriensis, 604 A. crassicornis, 68, 369, 370, 375 A. crenuliventris, 604 A. crucialis, 412 A. cryptica, 67, 470, 594, 604 A. cursor, 68, 382 A. curtipenis, 604 A. curvipennis, 387, 604 A. dadopora, 67, 367, 368, 382, 473, 594, 604 A. densissima, 243 A. dentata, 418 A. moesta, 381 A. districta, 68, 370, 378, 482, 594, 604 A. euryptera, 604 A. fanatica, 68, 369, 370, 379, 380, 483, 594, 604 A. festinans, 604 A. finita, 604 A. formalis, 68, 405, 594, 604 A. formicaensis, 604 A. frosti, 12, 68, 404, 506, 594, 604 A. frugalis, 372 A. gelida, 604 A. giguereae, 604 A. globicollis, 454 A. godini, 68, 375, 478, 594, 604 A. graminicola, 35, 67, 86, 364, 468, 594, 604 A. hampshirensis, 68, 369, 370, 381, 382, 485, 594, 604 A. holmbergi, 68, 395, 396, 498, 594, 604 A. impressicollis, 412 A. iowensis, 418 A. irrita, 68, 379 A. irrupta, 68, 405, 406, 508, 594, 604 A. keeni, 68, 391, 494, 594, 604 A. klagesi, 12, 403, 604 A. larsonae, 604 A. laticollis, 409 A. lindrothides, 604 A. longiclava, 604 A. longicornis, 12, 54, 67, 86, 366, 435, 471, 594, 604 A. luctifera, 68, 395, 397, 499, 594, 604 A. macesi, 604 A. makepeacei, 604 A. manitobae, 604 A. martini, 605 A. mcalpinei, 605 A. meduxnekeagensis, 68, 390, 493, 594, 605 A. metlakatlana, 68, 375–377, 479, 594, 605 A. microelytrata, 605 A. modesta, 605 A. moesta, 381

Index A. mordax, 443 A. munsteri, 68, 380, 484, 594 A. nearctica, 605 A. nescia, 68, 400, 503, 595, 605 A. newfoundlandica, 605, 608 A. nigra, 605 A. nigritula, 404 A. nimia, 177 A. nova, 412 A. novaescotiae, 605 A. omissa, 379 A. pancota, 68, 400 A. particula, 605 A. parvipennis, 410 A. pecki, 605 A. pectorina, 412 A. pennsylvanica, 362, 605 A. petitcapensis, 605 A. picipennis, 68, 369, 371, 382, 486, 595, 605 A. platonoffi, 68, 395, 398, 400, 500, 595, 605 A. pratensis, 605 A. prudhoensis, 35, 68, 369, 371, 384, 386, 387, 389, 489, 595 A. pseudocrenuliventris, 605 A. pseudodistricta, 379, 605 A. pseudoklagesi, 68, 401–403, 505, 595, 605 A. pseudometlakatlana, 68, 376, 377, 480, 595, 605 A. pseudomodesta, 605 A. pseudopittionii, 605 A. pseudoschistoglossa, 68, 406, 407, 509, 595, 605 A. pseudosubtilis, 605 A. pseudovestita, 605 A. quebecensis, 605 A. recondita, 606 A. regissalmonis, 606 A. relicta, 68, 391, 392, 495, 595, 606 A. retrusa, 381 A. revoluta, 417 A. richardsoni, 606 A. ringi, 67, 364, 469, 595, 606 A. ripariides, 68, 395, 399, 501, 595, 606 A. rurigena, 68, 387, 490, 595, 606 A. savardae, 394, 606 A. sculptisoma, 606 A. smetanai, 606 A. sordida, 434 A. sparreschneideri, 58, 606 A. spermathecorum, 606 A. sphagnicola, 606 A. stercoris, 68, 373, 476, 595, 606 A. strigosula, 12, 68, 388, 491, 595, 606 A. subdepressa, 416 A. subrugosa, 407, 511 A. subsinuata, 57, 58, 606 A. subtilis, 606 A. sumpta, 401 A. surgens, 68, 395, 399, 400, 502, 595, 606 A. terranovae, 68, 386–389, 492, 595, 606 A. trinotata, 606

625 A. vaga, 380 A. vancouveri, 400 A. vasta, 391 A. ventricosa, 67, 85, 363, 467, 595, 606 A. vestita, 606 A. vicaria, 379 A. vincenti, 606 A. wheelerae, 68, 377, 481, 595, 606 A. whitehorsensis, 382, 606 A. winchesteri, 68, 374, 477, 595, 606 Athetalia, 177 ATHETINI, 2, 10–12, 18, 21, 26, 46, 53, 54, 56, 57, 63, 67, 77, 78, 351, 352, 594, 602 Athetota atriventris, 423 Autalia, 11, 63, 295–300 A. puncticollis, 28, 57, 66, 295, 298, 597, 611 A. rivularis, 66, 80, 295, 296, 299, 597, 612 A. truncatula, 66, 295, 297, 300, 597, 612 AUTALIINI, 56, 63, 75, 295, 597, 611 B Baryodma, 58, 102, 104, 105 B. ontarionis, 58, 104 B. rubricalis, 102 B. suffusa, 105 Bessobia, 67, 361, 365 Bessobia cryptica, 365 Betocalea, 12, 54, 63, 65, 143, 170 Betocalea pacifica, 65, 170, 216, 598, 617 Blepharhymenus, 63, 64, 141, 143–144 B. brendeli, 144, 617 B. illectus, 64, 144, 187, 598, 617 Boletus, 104, 240, 374, 379, 382, 435 Bolitochara, 146, 307, 308 B. californica, 308 B. notata, 307 B. suturalis, 146 Borboropora quadriceps, 612 Boreocypha websteri, 612 BOREOCYPHINI, 56, 57, 612 Boreomorpha, 70, 450 Boreophilia, 13, 63, 69, 351, 355, 380, 381, 407, 408 B. caseyi, 58, 607 B. davidgei, 606 B. eremita, 606 B. fusca, 606 B. hyperborea, 606 B. islandica, 606 B. nearctica, 607 B. neoinsecuta, 607 B. nomensis, 69, 86, 408, 512, 595, 607 B. ovalis, 607 B. subplana, 607 B. vega, 607 B. venti, 58, 607 Boreostiba, 63, 69, 351, 355, 408–411 B. campbelliana, 411, 607 B. frigida, 607 B. hudsonica, 410

626 Boreostiba (cont.) B. lagunae, 607 B. laticollis, 409 B. parvipennis, 69, 86, 409–411, 514, 595, 607 B. pseudolaticollis, 69, 409, 513, 595, 607 B. sibirica, 607 B. websteri, 607 Brachyusa, 63, 235, 236 B. americana, 56, 65, 236, 247, 600, 620 B. helenae, 34, 56, 65, 79, 236, 248, 600, 620 B. saskatchewanae, 620 Bryobiota, 63, 70, 571–573 B. bicolor, 70, 283, 572, 573, 577, 597, 612 B. giulianii, 573 Bryothinusa, 63, 279–281 Bryothinusa catalinae, 57, 66, 81, 279–281, 597, 612 Bubo virginianus, 102 C Callicerus, 54 C. obscurus, 607 C. rigidicornis, 607 Calliphoridae, 104 Calochara, 98, 101, 115 Calodera, 12, 63, 64, 141, 146 C. bennetti, 83, 146, 190, 598, 617 C. caseyi, 617 C. parviceps, 617 Camponotus, 584 C. herculeanus, 584 C. laevigatus, 584 C. modoc, 584 C. noveboracensis, 584 Carex, 147, 161, 237, 239, 271, 364, 373, 407, 424, 427, 442, 451–453 Castor canadensis, 100, 237 Chamaepsila rosae, 413 Cladina, 404, 456 Clusiota, 13, 18, 63, 69, 353, 411, 412 C. claviventris, 411 C. grandipenis, 607 C. impressicollis, 69, 411, 412, 515, 595, 607 Cordalia, 63, 70, 571, 573–574 C. obscura, 70, 79, 574, 579, 597, 612 Crataraea, 63, 142, 145 C. suturalis, 34, 58, 64, 83, 146, 189, 598, 617 Cypha, 63, 261–262 C. crotchii, 34, 66, 81, 262, 265, 598, 615 C. inexpectata, 262, 615 Cyphea curtula, 613 Cyphea wallisi, 613 D Dalotia, 63, 69, 351, 355, 412 D. coriaria, 13, 69, 412, 413, 516, 595, 607 Datomicra, 67, 362, 367, 382, 394, 400 D. decolorata, 368 D. inopia, 368 D. schematica, 368

Index D. stilla, 368 D. surgens, 399 D. vaciva, 417 D. wrangeli, 367 Deinopsis, 11, 75, 76, 89 D. canadensis, 612 D. harringtoni, 612 D. illinoisensis, 612 D. rhadina, 612 Delia radicum, 104, 413 Dendroctonus, 185 D. ponderosae, 184, 339 D. rufipennis, 339, 341 Devia, 63, 141, 142, 147 D. prospera, 35, 64, 83, 148, 191, 429, 598, 617 Dexiogyia angustiventris, 617 Diaulota, 2, 12, 26, 57, 63, 66, 76, 283, 286–287 D. densissima, 66, 81, 286, 291, 598, 615 Diglotta, 279 D. mersa, 57, 612 DIGLOTTINI, 56, 63, 66, 76, 279, 597, 612 Dinaraea, 13, 18, 63, 69, 354, 413 D. angustula, 57, 69, 414, 517, 595, 607 D. backusensis, 607 D. bicornis, 607 D. borealis, 607 D. curtipenis, 416, 607 D. inexpectata, 69, 414, 415, 518, 595, 607 D. longipenis, 607 D. nomensis, 408 D. pacei, 69, 414–416, 519, 595, 607 D. piceana, 607 D. planaris, 58, 608 D. quadricornis, 608 D. subdepressa, 69, 414, 416, 520, 595, 608 D. worki, 608 Dochmonota, 63, 69, 354, 417, 418 D. langori, 608 D. rudiventris, 69, 417, 521, 595, 608 D. simulans, 608 D. websteri, 608 Drosophila, 413 D. suzukii, 413 Drusilla canaliculata, 616 Dryas, 456 E Earota, 12, 63, 69, 351, 352, 355, 418 E. dentata, 418, 522, 595, 608 Echidnoglossa illecta, 144 Egregia, 288 Emmelostiba microptera, 58, 608 Encephalus, 63, 66, 303, 310 E. americanus, 67, 82, 310, 326, 597, 613 Eumicrota E. corruscula, 613 E. socia, 613 Eurypronota discreta, 433 Euryusa obtusa, 429

Index Euvira micmac, 619 Evanystes, 435 F Falagria, 63, 70, 571, 574–575 F. caesa, 612 F. dissecta, 70, 79, 575, 580, 597, 612 F. sulcata, 575 FALAGRIINI, 11, 56, 63, 70, 75, 283, 571–581, 597, 612 Formica, 26, 421, 584 F. integra, 421 F. rufa, 421 F. subpolita, 584 Fucus, 287 G Gansia, 6 Gatineauleptusa, 66, 304 Gennadota, 56 G. canadensis, 617 Geomys, 419 Geostiba, 12, 63, 69, 353, 419, 435 G. appalachigena, 608 G. circellaris, 608 G. horwoodae, 69, 419, 523, 596, 608 Gnathusa, 13, 63, 142, 148–150 G. alfacaribou, 617 G. caribou, 617 G. eva, 12, 64, 83, 149, 192, 598, 617 G. tenuicornis, 64, 149, 193, 598, 617 Gnypeta, 6, 12, 13, 20, 26, 56, 63, 235, 237, 238, 243 G. ashei, 58, 620 G. atrolucens, 620 G. baranowskii, 56, 66, 238, 253, 600, 620 G. brevicornis, 56, 66, 238, 242, 254, 600, 620 G. brincki, 239, 620 G. caerula, 238 G. caerulea, 28, 65, 79, 238, 239, 249, 600, 620 G. canadensis, 620 G. carbonaria, 239, 620 G. crebrepunctata, 65, 238, 240, 251, 600, 620 G. dentata, 620 G. groenlandica, 620 G. helenae, 66, 238, 240, 241, 252, 600, 620 G. lohsei, 35, 65, 238, 239, 250, 600, 620 G. manitobae, 620 G. minuta, 620 G. nigrella, 31, 66, 241, 620 G. saccharina, 242, 620 G. sellmani, 66, 242, 620 G. uteana, 620 Goniusa, 12, 63, 69, 353, 420, 421, 462 G. alperti, 608 G. carrorum, 608 G. caseyi, 69, 421, 524, 596, 608 Gymnusa, 11, 18, 63, 75, 76, 78, 89–91 G. atra, 63, 90, 93, 597, 612 G. brevicollis, 90, 612 G. campbelli, 612

627 G. grandiceps, 28, 34, 63, 80, 90, 91, 94, 597, 612 G. konopackii, 58, 613 G. lindrothi, 613 G. pseudovariegata, 63, 90, 91, 95, 597, 613 G. smetanai, 613 G. variegata, 91 GYMNUSINI, 56, 63, 76, 89, 597, 612 Gyronycha, 56 G. pseudoobscura, 617 Gyrophaena, 9, 11, 18, 21, 56, 63, 304, 310–311, 315 G. affinis, 67, 311, 312, 327, 597, 613 G. antennalis, 613 G. bihamata, 67, 312, 313 G. brevicollis, 613 G. californica, 67, 82, 311, 315, 331, 597, 613 G. caseyi, 613 G. chippewa, 613 G. criddlei, 613 G. dybasi, 613 G. egena, 613 G. flavicornis, 312, 613 G. fuscicollis, 613 G. gaudens, 313, 613 G. gilvicollis, 613 G. gracilis, 613 G. illiana, 613 G. insolens, 613 G. involuta, 613 G. keeni, 67, 312–314, 329, 597, 614 G. lacustris, 312 G. laetula, 614 G. lobata, 614 G. meduxnekeagensis, 614 G. michigana, 313, 614 G. modesta, 614 G. nana, 67, 311, 314, 330, 597, 614 G. nanoides, 614 G. neonana, 614 G. orientalis, 314 G. pacifica, 313 G. pseudocriddlei, 614 G. sculptipennis, 614 G. strictula, 67, 315 G. stroheckeri, 614 G. subnitens, 614 G. subpunctata, 312 G. uteana, 67, 312, 313, 328, 597, 614 G. vitrina, 614 G. wisconsinica, 614 Gyrophaenina, 6, 11, 66, 310 H Halobrecta flavipes, 621 Holobus, 57, 63, 261–263 H. chrysopygus, 615 H. vancouveri, 28, 57, 66, 81, 262, 266, 598, 615 Homalota H. cambrica, 358 H. celata, 367

628 Homalota (cont.) H. comparabilis, 448 H. coriaria, 412 H. geniculata, 461 H. laevicollis, 443 H. litoralis, 107 H. maritima, 356 H. nitens, 424 H. pallens, 145 H. picipennis, 382 H. plana, 58, 614 H. rudiventris, 417 HOMALOTINI, 2, 18, 21, 46, 54, 56, 63, 66, 76, 303, 597, 613 Hoplandria, 57, 615 H. klimaszewskii, 615 H. laevicollis, 615 H. laeviventris, 615 H. lateralis, 57, 615 HOPLANDRIINI, 56, 57, 615 Hydrosmecta, 351, 390 H. borealis, 608 H. caduca, 608 H. canadensis, 608 H. dulcis, 608 H. minutissimoides, 608 H. pseudodiosica, 608 Hydrosmectomorpha, 68, 361, 389, 390 Hylota, 617 H. cryptica, 617 H. ochracea, 617 HYPOCYPHTINI, 56, 57, 63, 66, 76, 261, 598, 615 Hypocyphtus crotchii, 262 I Ilyobates, 56 I. bennetti, 617 Ips pini, 339 Isoglossa, 12, 63, 65, 143, 173–175 I. agnita, 65, 174, 219, 599, 617 L Lamiota, 68, 362, 391, 402 Larix laricina, 185 Larus marinus, 155 Leptusa, 5, 12, 18, 63, 66, 303–305 L. brevicollis, 614 L. canonica, 614 L. carolinensis, 614 L. cribratula, 614 L. elegans, 614 L. gatineauensis, 66, 82, 305, 319, 598, 614 L. jucunda, 614 L. opaca, 614 L. pseudopaca, 614 Leptusomorpha, 56, 63, 66, 303, 305–306 L. claudiae, 56, 66, 82, 305, 306, 320, 598, 614 Liogluta, 13, 20, 27, 54, 63, 69, 351, 354, 401, 422, 423, 427 L. aloconotoides, 426 L. atriventris, 69, 422, 423, 525, 596, 608

Index L. castoris, 608 L. gigantea, 608 L. granulosa, 58, 608 L. intermedia, 608 L. microgranulosa, 69, 423, 609 L. nigropolita, 69, 423, 609 L. nitens, 69, 422–424, 526, 596, 609 L. pseudocastoris, 609 L. quadricollis, 69, 422, 425, 527, 596, 609 L. scudderi, 69, 422, 425, 528, 596, 609 L. terminalis, 69, 423, 425, 426, 428, 529, 596, 609 L. trapezicollis, 69, 422, 423, 427, 428, 530, 596, 609 L. vasta, 428 L. wickhami, 69, 422, 428, 531, 596, 609 LIPAROCEPHALINI, 11, 12, 26, 54, 56, 57, 63, 66, 76, 283–284, 573, 598, 615 Liparocephalus, 2, 11, 12, 26, 57, 63, 66, 287–293 L. brevipennis, 34, 66, 287, 288, 292, 598, 615 L. cordicollis, 28, 66, 81, 287, 288, 293, 598, 616 Lomechusa montana, 584 LOMECHUSINI, 56, 63, 70, 77, 351, 583–584, 590, 598, 616 Lorinota arizonica, 576 Lypoglossa, 12, 63, 69, 351–353, 429 L. angularis, 69, 429, 430, 532, 596, 609 L. franclemonti, 69, 429, 430, 533, 596, 609 L. manitobae, 609 M Macroterma alutacea, 418 Marmota monax, 372 Megocalea lemieuxi, 28, 65, 83, 172, 217, 598, 617 Meotica, 12, 18, 63, 64, 77, 142, 144 M. exilis, 617 M. pallens, 64, 84, 144, 188, 599, 617 M. pseudowinkleri, 617 Meronera, 351 M. venustula, 609 Metadimetrota, 68, 361, 393 Metaxya prognata, 443 Metocalea, 12, 54, 63, 143, 172–173 M. lindgreni, 65, 173, 218, 599, 617 Microdota, 68, 362, 371, 394, 399 Microglottina, 145, 272 Microtus, 435, 575 M. ochrogaster, 575 M. pennsylvanicus, 435, 575 Mniusa, 13, 63, 64, 142, 150–151 M. minutissima, 617 M. odelli, 617 M. yukonensis, 64, 84, 150, 194, 599, 617 Mocyta, 13, 20, 63, 69, 351, 353, 431, 433 M. amblystegii, 58, 69, 431, 433, 609 M. breviuscula, 69, 431, 432, 534, 596, 609 M. discreta, 69, 86, 431, 536, 596, 609 M. fungi, 12, 13, 57, 69, 431–433, 535, 596, 609 M. luteola, 609 M. sphagnorum, 609 Moluciba grandipennis, 166 Muscidae, 104, 113 Myllaena, 18, 63, 78, 152, 269–272

Index M. arcana, 616 M. audax, 66, 270, 273, 598, 616 M. cuneata, 616 M. decreta, 66, 270–271, 274, 598, 616 M. fenyesi, 34, 270–272, 276, 598, 616 M. insomnis, 28, 35, 66, 80, 270, 271, 275, 598, 616 M. kaskaskia, 616 M. ludificans, 616 M. potawatomi, 66, 271, 616 M. procidua, 616 M. vulpina, 616 MYLLAENINI, 11, 56, 63, 66, 76, 269, 598, 616 Myrmecocephalus, 63, 70, 571, 575–576 M. arizonicus, 79, 576, 581, 597, 612 M. cingulatus, 612 M. concinnus, 29, 58, 612 M. gatineauensis, 612 Myrmecopora, 572 M. californica, 572 M. vaga, 612 Myrmedonia obliqua, 586 MYRMEDONIINA, 583–585 Myrmedonota, 351 M. aidani, 616 Myrmoecia, 63, 70, 584–586 M. canadensis, 585, 589, 616 M. lauta, 616 M. lugubris, 70, 85, 585, 588, 598, 616 N Nehemitropia, 63, 69 N. lividipennis, 35, 57, 69, 434, 435, 537, 596, 609 Neothetalia, 54, 63, 65, 143, 175–180 N. canadiana, 54, 65, 168, 175, 176, 220, 599, 618 N. columbiana, 65, 175, 176, 221, 599, 618 N. nimia, 65, 84, 175–177, 222, 599, 618 N. pallitarsis, 65, 175, 176, 178, 223, 599, 618 N. robergei, 65, 176, 179, 224, 599, 618 N. smetanai, 65, 175, 176, 225, 599, 618 Neotobia alberta, 614 Nototaphra lugubris, 585 Noverota clemens, 442 O Ocalea, 12, 54, 63, 141–143, 168, 178, 180, 181 O. agnita, 174 O. picata, 181 O. vancouveri, 181, 226, 599, 618 Ocyusa, 12, 13, 63, 64, 142, 150–152 O. asperula, 618 O. canadensis, 84, 151, 195, 599, 618 O. yukonensis, 150 Oligota, 18, 63, 66, 261–264 O. inflata, 263, 615 O. parva, 57, 66, 263, 267, 615 O. polyporicola, 615 O. pusillima, 615 O. sevogle, 615 Ousipalia, 54, 63, 69, 353, 419, 435

629 O. pacifica, 69, 435, 538, 596, 609 Oxypoda, 12, 18, 19, 21, 54, 63, 64, 75, 76, 78, 141, 142, 152–156, 165, 166, 168, 366, 434, 435, 455 O. amica, 618 O. brachyptera, 618 O. canadensis, 64, 153, 154, 196, 599, 618 O. chantali, 618 O. convergens, 34, 64, 84, 153–156, 197, 435, 599, 618 O. demissa, 618 O. domestica, 618 O. famula, 64, 154, 160, 203, 599, 618 O. frigida, 65, 154, 163, 206, 599, 618 O. gatosensis, 64, 153, 157, 158, 200, 599, 618 O. glenorae, 65, 153, 167, 212, 599, 618 O. gnara, 618 O. hiemalis, 161, 618 O. impressa, 65, 153, 167, 168, 213, 599, 618 O. inimica, 618 O. irrasa, 64, 153, 159, 202, 599, 618 O. lacustris, 64, 154, 160, 161, 204, 599, 618 O. leechi, 618 O. lividipennis, 434 O. longicarinata, 64, 153, 158, 201, 599, 618 O. lucidula, 618 O. manitobae, 64, 153, 156, 198, 599, 618 O. nigriceps, 618 O. nimbata, 64, 154, 160, 162, 205, 599, 618 O. opaca, 13, 64, 153, 157, 199, 599, 619 O. operta, 619 O. orbicollis, 65, 152, 162, 163, 207, 599, 619 O. perexilis, 619 O. prospera, 148 O. pseudoconvergens, 619 O. pseudolacustris, 161, 619 O. regressa, 65, 154, 168, 214, 599, 619 O. rubescans, 619 O. smithi, 65, 154, 164, 208, 599, 619 O. spectabilis, 166 O. stanleyi, 13, 65, 152, 164, 165, 209, 599, 619 O. sunpokeana, 619 O. sylvia, 28, 35, 65, 84, 153, 165, 166, 210, 599, 619 O. vancouveri, 65, 153, 166, 167, 181, 211, 599, 619 O. vockerothi, 619 O. volkeri, 619 O. yukonensis, 619 OXYPODINI, 2, 18, 21, 46, 53, 54, 56, 64, 77, 141, 598, 616 P Paradilacra, 63, 235, 243 P. densissima, 66, 243, 255, 600, 620 Paragoniusa, 13, 63, 70, 351, 353, 462 P. myrmicae, 13, 70, 462, 565, 596, 609 Paraleptonia, 12, 13, 46, 54, 63, 69, 355, 437 P. pacei, 46, 69, 437, 539, 596, 609 Paramblopusa, 2, 12, 26, 57, 63, 66, 76, 283, 285–286 P. borealis, 66, 285, 290, 598, 616 Parocalea, 12, 63, 65, 143, 181–183

630 Parocalea (cont.) P. nearctica, 182, 619 P. pseudobaicalica, 65, 168, 182, 227, 599, 619 Pelioptera, 12, 63, 69, 353, 438 P. americana, 609 P. robusta, 69, 438, 439, 540, 596, 609 P. thujae, 609 Pella, 586 P. caliginosa, 616 P. carolinae, 616 P. criddlei, 616 P. gesneri, 616 P. glooscapi, 616 P. loricata, 616 P. schmitti, 616 Phanerota fasciata, 614 Philhygra, 26, 54, 63, 69, 351, 354, 407, 439, 442 P. angusticauda, 69, 440, 541, 596, 609 P. atypicalis, 610 P. botanicarum, 13, 69, 440, 441, 542, 596, 610 P. charlottae, 69, 440, 442, 543, 596, 610 P. clemens, 13, 69, 440, 442, 544, 596, 610 P. criddlei, 610 P. humivaga, 13 P. hygrotopora, 610 P. jarmilae, 610 P. junii, 610 P. laevicollis, 69, 440, 443, 444, 545, 596, 610 P. larsoni, 610 P. leechi, 610 P. luridipennis, 610 P. malleoides, 610 P. manitobae, 610 P. palustris, 610 P. proterminalis, 610 P. pseudoboreostiba, 610 P. pseudolarsoni, 610 P. pseudopolaris, 610 P. pseudoterminalis, 610 P. ripicoloides, 610 P. rostrifera, 610 P. satanas, 610 P. sinuipennis, 610 P. subpolaris, 610 P. terrestris, 69, 440, 444, 546, 596, 610 P. terrivaga, 69, 440, 445, 547, 596, 610 P. varula, 610 P. wisconsinica, 610 Phloeopora, 63, 65, 141, 183 P. arctica, 65, 183, 228, 599, 619 P. canadensis, 65, 183–185, 229, 600, 619 P. gilbertae, 619 P. oregona, 65, 183, 185, 230, 231, 600, 619 Phloeoporina, 182, 183 Pholiota, 166, 416 Phymatura blanchardi, 614 Phytosus P. bicolor, 572 P. fletcheri, 459 P. maritimus, 459 Pinus

Index P. contorta, 184, 185, 339–341 Placusa, 12, 18, 63, 67, 337–349 P. atrata, 340 P. canadensis, 337, 338, 619 P. complanata, 339 P. cribrata, 342 P. despecta, 340, 619 P. incompleta, 67, 338, 340, 343, 347, 600, 619 P. petulans, 67, 338, 339, 344, 600, 619 P. pseudosuecica, 67, 338, 341, 342, 348, 600, 620 P. suecica, 342 P. tachyporoides, 67, 337, 338, 340, 342, 343, 349, 600, 620 P. tacomae, 80, 338, 339, 345, 600, 620 P. vaga, 67, 337, 340, 346, 600, 620 PLACUSINI, 56, 63, 67, 76, 337, 600, 619 Platandria carolinae, 615 Pleurotobia, 56 P. bourdonae, 614 P. brunswickensis, 614 Pleurotus, 312, 313, 369, 380 Polistoma pacifica, 107 Polygraphus rufipennis, 341 Polyporus betulinus, 305 Pontomalota, 12, 54, 63, 70, 352, 446, 447 P. opaca, 28, 70, 86, 447, 548, 596, 610 Porphyra, 288 Psammostiba, 2, 12, 54, 63, 70, 355, 356, 447 P. comparabilis, 70, 448, 549, 596, 610 P. kenaii, 70, 448, 449, 550, 596, 611 P. yosii, 2 Pseudota, 68, 362, 401, 402 P. formalis, 405 P. irrupta, 406 P. miscella, 412 P. nanulina, 67, 367 P. nescia, 400 Purshia, 162, 175, 414, 415, 434, 436 R Russula, 387 S Salix, 148, 364, 366, 388, 424, 444 Scatella, 413 Schistoglossa, 12, 13, 63, 70, 354, 407, 408, 449–451 S. blatchleyi, 611 S. brunswickensis, 453, 611 S. campbelli, 70, 450, 452, 553, 596, 611 S. carexiana, 70, 450, 551, 596, 611 S. charlottae, 70, 450, 552, 596, 611 S. hampshirensis, 70, 450, 453, 554, 596, 611 S. pelletieri, 611 S. pseudocampbelli, 453, 611 S. sphagnorum, 611 S. viduata, 451 Seeversiella, 12, 63, 70, 353, 454 S. bispinosa, 11, 454 S. globicollis, 11, 70, 454, 555, 596, 611 Silusa, 12, 63, 67, 303, 315–316, 334 S. alternans, 615

Index S. californica, 66, 67, 316, 332, 598, 615 S. densa, 615 S. langori, 615 S. opaca, 67, 316, 317, 333, 598, 615 S. prettyae, 615 S. vesperis, 67, 82, 316–318, 334, 598, 615 Silusida marginella, 615 Silusina, 67, 315 Sphagnum, 145, 147, 155, 163, 164, 377, 382, 442, 451, 452 Staphylinus S. curtulus, 99 S. nana, 314 Stethusa, 54, 351, 376 S. klimschi, 611 S. spuriella, 611 Stictalia, 63, 66, 303, 306–307, 310 S. arcuata, 66, 307 S. brevicornis, 35, 66, 307, 322, 598, 615 S. californica, 307, 308, 323, 598, 615 S. carlottae, 56, 66, 307–309, 324, 598, 615 S. densicollis, 66, 308 S. kranabetteri, 56, 66, 82, 307, 309, 325, 598, 615 S. notata, 307 Strigota, 12, 63, 70, 351, 352, 454, 455 S. ambigua, 12, 70, 455, 456, 556, 597, 611 S. obscurata, 70, 455, 456, 557, 597, 611 Strophogastra penicillata, 382, 611 T Tachiona, 6 Tachyusa, 12, 56, 63, 66, 235 T. americana, 245, 620 T. americanoides, 28, 66, 79, 244, 245, 256, 600, 620 T. carolinae, 245 T. cavicollis, 66, 244–246, 257, 600, 621 T. crebrepunctata, 240 T. faceta, 621 T. obsoleta, 66, 244, 246, 257, 600, 621 T. smetanai, 621 TACHYUSINI, 2, 54, 56, 63, 65, 75, 235, 600, 620 Tarphiota, 12, 26, 54, 63, 70, 351, 352, 457 T. fucicola, 70, 457, 558, 597, 611 T. geniculata, 70, 457, 461, 559, 597, 611 T. insolita, 457 T. iota, 457

631 T. litorina, 457 T. pallidipes, 457 T. seditiosa, 458 TAXICERINI, 56, 57, 621 Tetralaucopora, 619 T. americana, 619 T. fuliginosa, 619 Tetralina helenae, 236 Thamiaraea, 611 T. brittoni, 611 T. claydeni, 611 T. corverae, 611 Thecturota, 25, 56 T. capito, 615 T. tenuissima, 615 Thinusa, 12, 26, 54, 63, 70, 352, 458, 459 T. divergens, 458 T. fletcheri, 70, 458–460, 560, 597, 611 T. maritima, 70, 458–460, 561, 562, 597, 611 T. nigra, 458 T. robustula, 458, 459 Tomoglossa, 54 T. decora, 611 Trichiusa, 13, 63, 70, 353, 460 T. columbica, 70, 460, 461, 563, 597, 611 T. compacta, 611 T. geniculata, 461 T. hirsuta, 461, 611 T. pilosa, 35, 70, 460, 461, 564, 597, 611 T. pseudopostica, 611 T. robustula, 611 Tricholoma, 363 Tropimenelytron, 69, 438 T. robustum, 438 X Xenodusa, 11, 18, 63, 70, 583–584 X. cava, 616 X. reflexa, 28, 35, 70, 85, 584, 587, 598, 616 Z Zyras, 63, 70, 585–586 Z. obliquus, 70, 85, 586, 590, 598, 616 Z. planifer, 616 Z. pseudohaworthi, 586