Ladybird Beetles of the Australo-Pacific Region: Coleoptera: Coccinellidae: Coccinellini 9781486303878, 9781486303885, 9781486303892, 9781472978660

Citation preview

LADYBIRD BEETLES OF THE AUSTRALO-PACIFIC REGION Coleoptera: Coccinellidae: Coccinellini

Adam Slipinski, Jiahui Li and Hong Pang

LADYBIRD BEETLES OF THE AUSTRALO-PACIFIC REGION Coleoptera: Coccinellidae: Coccinellini

We dedicate this book to our distinguished elders, who, by their excellent work and dedication have paved the way to a better understanding of the ladybird beetles: Xiong-fei Pang (China), Hiroyuki Sasaji (Japan), Ryszard Bielawski (Poland), Robert D. Gordon (USA) and Helmut Fürsch (Germany).

LADYBIRD BEETLES OF THE AUSTRALO-PACIFIC REGION Coleoptera: Coccinellidae: Coccinellini

Adam Slipinski, Jiahui Li and Hong Pang

© Adam Ślipiński, Jiahui Li and Hong Pang 2020 All rights reserved. Except as permitted by applicable copyright laws, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO Publishing for all permission requests. The authors assert their moral rights, including the right to be identified as an author. A catalogue record for this book is available from the National Library of Australia and the British Library. Published in print in Australia and New Zealand, and in all other formats throughout the world, by CSIRO Publishing. CSIRO Publishing Locked Bag 10 Clayton South VIC 3169 Australia Telephone: +61 3 9545 8400 Email: [email protected] Website: www.publish.csiro.au ISBN: 9781486303878 (hbk) ISBN: 9781486303885 (epdf) ISBN: 9781486303892 (epub) Published in print only, throughout the world (except in Australia and New Zealand), by BLOOMSBURY WILDLIFE. BLOOMSBURY WILDLIFE Bloomsbury Publishing Plc 50 Bedford Square, London, WC1B 3DP, UK BLOOMSBURY, BLOOMSBURY WILDLIFE and the Diana logo are trademarks of Bloomsbury Publishing Plc ISBN: 978-1-4729-7866-0 (hbk) Front cover: (main image) Harmonia conformis (photo: Nick Monaghan); (top, left to right) Cleobora mellyi (photo: Jiri Lochman); Coccinella transversalis (photo: Marie Lochman); Illeis galbula (photo: Steve Axford); Harmonia octomaculata (photo: David McClenaghan, CSIRO); Micraspis frenata (photo: Paul Zborowski) Back cover: Harmonia testudinaria (photo: Jiri Lochman) Cover design by Andrew Weatherill Typeset by Thomson Digital Printed in China by Leo Paper Products Ltd. CSIRO Publishing publishes and distributes scientific, technical and health science books, magazines and journals from Australia to a worldwide audience and conducts these activities autonomously from the research activities of the Commonwealth Scientific and Industrial Research Organisation (CSIRO). The views expressed in this publication are those of the author(s) and do not necessarily represent those of, and should not be attributed to, the publisher or CSIRO. The copyright owner shall not be liable for technical or other errors or omissions contained herein. The reader/user accepts all risks and responsibility for losses, damages, costs and other consequences resulting directly or indirectly from using this information. The paper this book is printed on is in accordance with the standards of the Forest Stewardship Council® and other controlled material. The FSC® promotes environmentally responsible, socially beneficial and economically viable management of the world’s forests.

Oct19_01

CONTENTS

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vi Preface and acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii I. Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 II. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 III. The Coccinellini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 1. Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 3. Economic importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 IV. Classification of Australo-Pacific Coccinellini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 V. Keys to identification of Australo-Pacific genera of Coccinellini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 VI. Keys to identification of Australian species of Coccinellini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 VII. Review of the Australo-Pacific Coccinellini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Colour plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 Index of scientific names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229

ABSTRACT

The genera and species of the Australo-Pacific Coccinellini (Coleoptera: Coccinellidae: Coccinellinae) are reviewed. In total 22 genera and 95 valid species are recognised. All genera and species are diagnosed, described and illustrated and keys to their identifications are provided. An additional key to the Australian species of Coccinellini based on external characters is also provided. The following new species are described: Harmonia ohlmusi sp. nov. (the island of New Guinea); Harmonia boothi sp. nov. (Indonesia); Harmonia micra sp. nov. (Solomon Islands); Harmonia tobleri sp. nov. (Solomon Islands); Heterocaria bella sp. nov. (Australia); Heterocaria decemmaculata sp. nov. (Indonesia); Heterocaria riedeli sp. nov. (the island of New Guinea); Heterocaria transversoguttata sp. nov. (the island of New Guinea); Illeis wyliei sp. nov. (the island of New Guinea); Phrynocaria ambonina sp. nov. (Indonesia); Phrynocaria georgia sp. nov. (Solomon Islands); Phrynocaria kolombangara sp. nov. (Solomon Islands). The following new generic synonymies are proposed (junior synonym listed first): Antineda Iablokoff-Khnzorian, 1982 (= Docimocaria Crotch, 1874); Oiocaria Iablokoff-Khnzorian, 1982 (= Heterocaria Timberlake, 1943); Bothrocalvia Crotch, 1874 (= Microcaria Crotch, 1871). The following new specific synonymies are proposed (junior synonym listed first): Caria thoracica Weise, 1892a (= Coelophora pulchra Crotch, 1874); Harmonia uninotata Bielawski, 1964 (Harmonia basinotata Bielawski,

1964); Harmonia rhopalocera Iablokoff-Khnzorian, 1982 (= Harmonia nigromarginata Bielawski, 1968); Harmonia incognita Chazeau, 1990 (= Callineda decussata Crotch, 1874); Coelophora extensa Weise, 1902 (= Halyzia funerea Crotch, 1874); Coelophora auberti Chazeau, 1978 (= Coelophora vivida Crotch, 1874). Of the previously described species, Coelophora aruensis Crotch, 1874 and Neda princeps Mulsant, 1850 are transferred to Docimocaria Crotch. Coelophora tricolor Korschefsky, 1944; Synharmonia fijiensis Sicard, 1929a are transferred to Harmonia Mulsant. Cycloneda barronensis Blackburn, 1895; Lemnia kaszabi cinerea Chazeau, 1990; Egleis delta Weise, 1898; Halyzia funerea Crotch, 1874; Lemnia (Lemnia) kaszabi Iablokoff-Khnzorian, 1982 and Oiocaria samuelsoni Iablokoff-Khnzorian, 1982 are transferred to Heterocaria Timberlake. Callineda principalis Weise, 1895 and Lemnia grata Iablokoff-Khnzorian, 1982 are transferred to Heteroneda Crotch. Coelophora salomonensis Korschefsky, 1940; Coelophora versipellis Mulsant, 1850 and Coelophora vivida Crotch, 1874 are transferred to Microcaria Crotch. Lemnia (Microlemnia) aurita Iablokoff-Khnzorian, 1982; Coelophora moseri Weise, 1902; Coelophora quadrivittata Fauvel, 1903 are transferred to Phrynocaria Timberlake. To stabilise the taxonomic status of some species, the lectotypes of Synharmonia fijiensis Sicard, 1929a and Coelophora extensa var. mysolensis Weise, 1902, and the neotype of Callineda principalis Weise, 1895 and Oiocaria samuelsoni Iablokoff-Khnzorian, 1982, are designated.

PREFACE AND ACKNOWLEDGMENTS

The ladybirds are iconic beetles that we are all familiar with since our childhoods, having observed colourful beetles wandering on leaves and flowers and feeding on aphids on plants in our gardens. Most of these commonly observed ladybirds belong to the true ladybirds, currently classified in the tribe Coccinellini. They are easily recognisable by their relatively large and glabrous shiny bodies, bearing contrasting colour patterns on their elytra and leaving droplets of a slightly smelly, yellow substance on our hands when handled. The Coccinellini is one of the most widely studied groups of beetles. They have been the subject of thousands of research papers and many high-impact contributions featured in textbooks (Majerus 1994, 2016; Hodek and Honěk 1996; Hodek et al. 2012). They are often discussed in the popular media, which comment on their apparent scarcity or sudden abundance. They are important model objects of biological and ecological research, and the polymorphic colour patterns of many species have been a subject of genetic research for several years (Hodek 1973; Hales 1976; Houston and Hales 1980; Majerus 1994, 2016). Many species of Coccinellini are also of great economic importance, either as biological control agents, or as unwanted invaders on a scale of entire continents (e.g. the Asian harlequin ladybird beetle, Harmonia axyridis (see Brown et al. 2011)). The idea of a modern illustrated guide to the Australian species of Coccinellini was conceived by A. Ślipiński over 10 years ago, during his research on the genera of the Australian ladybirds (Ślipiński 2007). The original intention was to produce an online resource allowing reliable identification of most Australian species. However, over the years the project grew, as it became evident that the generic classification of the Australian taxa needed a thorough revision in a broader geographic context by using both morphology and modern molecular methods. We started working on molecular phylogeny of Coccinellini during the PhD tenure of Jiahui Li at CSIRO, under the supervision of H. Pang and A. Ślipiński. The project has been further developed and expanded at the Australian National Insect Collection (ANIC), CSIRO, by Hermes Escalona, Xingmin Wang, Wioletta Tomaszewska and Andreas Zwick – leading to the publication of the first underlying phylogeny and worldwide generic classification of the Coccinellini (Escalona et al. 2017; Tomaszewska et al., pers. comm.). Recently, we decided to expand the geographical scope of the project to include the island of New Guinea and neighbouring Pacific islands, to incorporate the new generic concepts developed during molecular research, and to publish in book form. It took several more years and many overseas trips to examine the type specimens and prepare the text and illustrations to finish this much expanded research. The expansion of the geographic range covered by the book from Australia to the entire Australo-Pacific region took its toll as

instead of 33 species, we now include almost 100 species, with many of them new. We hope this effort was worthwhile, and the book will be a valuable contribution to the taxonomy of the ladybirds and to the knowledge of the biodiversity of this unique biogeographic region. Over the years, generous support from the Australian National Insect Collection (CSIRO), the Australian Biological Resources Study (Department of the Environment and Energy) and the Department of Agriculture and Water Resources (G. Maynard, D. Kerruish and C. Mathenge) provided A. Ślipiński with research facilities and funding, enabling us to continue this research to its successful conclusion. Additional funding for this study was provided by a grant from the Polish National Science Centre (Narodowe Centrum Nauki), No. 2012/07/B/ NZ8/02815, to W. Tomaszewska and A. Ślipiński, and the National Natural Science Foundation of China, Grant No. 31572052, National Key R & D Program of China, Grant No. 2017YFD0201000, Science-Technology Basic Condition Platform from The Ministry of Science and Technology of the People’s Republic of China, National Specimen Infrastructure (NSII) (Grant No. 2005DKA21402) to H. Pang. The doctoral research of Jahui Li in Australia was funded by the China Scholarship Council (CSC). We thank many individuals and institutions for making available type material and other specimens used in this study: D. Smith and C.A.M. Reid (AM); R.G. Booth (BMNH); A. Ramsdale (BPBM); F. Gernier (CMN); P. Bouchard and A. Zmudzinska (CNC); W. Tomaszewska, T. Huflejt, D. Iwan (MIIZ); B. Viklund (NRM); P. Gillespie (NSWDA); G.B. Monteith (QMB); E.G. Matthews and P. Hudson (SAMA); N. Vandenberg (USNM); G. Daniels (UQIC); and P. Perkins (MCZ). Special thanks are due to Roger G. Booth (BMNH) for his support, friendship and unlimited access to his vast knowledge of ladybirds, as well as for extensive loans of material under his care, without which it would have been impossible to complete this study. Wioletta Tomaszewska and Hermes Escalona are warmly acknowledged for their support and collaboration on various papers leading to the production of this book. ANIC technical staff, Cate Lemann and Anne Hastings, are sincerely acknowledged for their support, digital imaging work and assistance with various aspects of the production of this book. Colour photographs of live ladybird beetles in their native environments (Figs 1–6) have been provided generously at no cost by Jiří and Marie Lochmann (Lochmann Transparencies, Perth); Steve Axford (Melbourne); Paul Zborowski (Close-UpPhotolibrary); Nick Monaghan (Life Unseen); and Yun Li (Sun Yat-sen University, Guangzhou). We would particularly like to acknowledge the Australian Biological Resources Study for continuous interest and support of our research, especially Haylee Weaver for editing the book, and Brigitte Kuchlmayr for her work on the graphics.

This page intentionally left blank

I. MATERIAL AND METHODS This book is based on more than 10 000 specimens of the Australian, New Guinean and Pacific Coccinellini examined during the past decade. Specimens representing most of the world genera were also examined for comparative purposes. Some of the generic concepts are based on still unpublished molecular research led by Wioletta Tomaszewska, Hermes Escalona and Andreas Zwick (Tomaszewska et al., pers. comm.). The following abbreviations are used for the names of the institutions where the specimens used in the project are held. The names in parentheses are the curators who arranged the loans: AM Australian Museum, Sydney (D. Smith, C. Reid) ANIC Australian National Insect Collection, Canberra BMNH Natural History Museum, London (R.G. Booth) BPBM Bernice Pauahi Bishop Museum, Honolulu (S. Myers, A. Samuelson, N. Evenhuis) CMN Canadian Museum of Nature, Ottawa (F. Gernier) CNC Canadian National Insect Collection, Ottawa (P. Bouchard, A. Zmudzinska) DARI Insect Collection, New South Wales Department of Agriculture, Orange, New South Wales (P. Gillespie) DEI Deutsches Entomologisches Institut, Müncheberg (S. Blank, L. Behne) ELKU Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka (S. Kamitani) MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Mass. (P. Perkins) MHNG Muséum d’histoire Naturelle, Genève (G. Cuccodoro, I. Löbl) MIIZ Museum and Institute of Zoology, PAS, Warsaw (W. Tomaszewska, T. Hufjelt, D. Iwan) MNHN Muséum National d’histoire Naturelle, Paris (A. Mantillieri, T. Deuve, R. Nattier) NAQS Northern Australia Quarantine Strategy, Department of Agriculture and Water Resources, Mareeba (J.F. Grimshaw) NBCL Naturalis Biodiversity Center, Leiden (J. Krikken) NHMB Naturhistorisches Museum, Basel (Isabelle ZürcherPfander) NKME Naturkunde Museum, Erfurt (M. Hartmann) NRM Naturhistoriska Riksmuseet, Stockholm (B. Viklund) NTDA Northern Territory Department of Primary Industry and Fisheries, Darwin (H. Brown) OMNH Oxford University Museum of Natural History, Oxford (J. Hogan) QDPIB Entomology Collection, Queensland Primary Industries and Fisheries, Indooroopilly (J. Donaldson, J. Bartlett) QMB Queensland Museum, Brisbane (G. Monteith, S. Wright) SAMA South Australian Museum, Adelaide (E.G. Matthews and P. Hudson) SYSBM The Museum of Biology, Sun Yat-sen University, Guangzhou (H. Pang) UCCC University of Cambridge Crotch Collections, Cambridge (R.C. Stebbings, W. Foster) UQIC University of Queensland Insect Collection (now deposited in QMB)

USNM United States National Museum, Washington DC (N. Vandenberg) ZMB Museum für Naturkunde, Berlin (M. Uhlig, B. Jäeger) In order to examine the morphological characters for generic and species diagnoses, many specimens were completely cleared, disarticulated and mounted in glycerine on slides for further examination. The structural illustrations were based on dry specimens, and from the glycerine slides using digital cameras attached to compound and stereo microscopes. The habitus images were generated in ANIC using the BK PLUS Laboratory System (Dun Inc., USA). Digital images have been enhanced and assembled in plates using Adobe Photoshop. Measurements. In species descriptions, several specimens of extreme size and variability were usually measured, always including the largest and the smallest available specimens. The numbers presented in the measurement sections include the ranges of these measurements in millimetres, and the mean and standard deviation. Standard ratios calculated on the pronotum and elytra are also provided. Measurements were made using a micrometer attached to a dissecting (stereo) microscope and are abbreviated as follows: ● (TL) total length (= body length), from apical margin of the clypeus to apex of the elytra; ● (PL) pronotal length, from the middle of the anterior margin to the margin of the basal foramen; ● (PW) the widest point of the pronotum; ● (EL) elytral length along the suture, including the scutellum; ● (EW) elytral width across both elytra at the widest part; ● (GD) greatest depth from the highest point of the elytra in a straight line to the lowest point of the metaventrite. Terminology (Figs 7–9) used for adult and larval morphology follows Ślipiński (2007). Ślipiński (2007) departed from a traditional nomenclature describing genitalia of male coccinellid beetles and applied general beetle terminology with the penis (= sipho) and tegmen forming jointly the beetle aedeagus. The tegmen consists of the phallobase (= basal piece), penis guide (= basal lobe, medial lobe) and parameres (= lateral lobes). Male and female genitalia in routine identifications were dissected, cleared in a 10% solution of potassium hydroxide and mounted in glycerine for examination and photography. After examination, the preparations were stored in a microvial in glycerine, or mounted permanently on a card in a drop of DMHF (dimethyl hydantoin formaldehyde), and pinned with the specimen. Taxonomic conventions. A diagnosis and complete description is presented for each genus and species. The generic diagnostic combination is a summary of the critical characters of the adult ladybird beetle that should be checked, without a lengthy description, once a specimen has been successfully keyed out using the generic key. Identified beetles should possess all, or most of, the mentioned character states. Under ‘Remarks’ there is a statement regarding genera that are so similar and could be confused with the genus in question, and a list of the critical distinguishing features is provided. The species diagnosis lists the characters that separate species from the most similar taxa. Geographic coverage and conventions. The geographic area covered in this book (Fig. 10) encompasses three provinces of

2

Ladybird Beetles of the Australo-Pacific Region

Oceania, Australasia (Australia and New Zealand), Melanesia (New Guinea, Solomon Islands, New Caledonia, Vanuatu and Fiji) and Polynesia (limited here to Tonga and Samoa). The north-west boundary of the study area is mostly set by the Lydekker’s Line (Lydekker 1896) which runs along the border of Australia’s continental shelf (the Sahul Shelf). However, in the species treatments, we have also included species known from the islands of Aru, Kai, Seram and Ambon of the Maluku

Archipelago (Indonesia), as many species described from those islands have been subsequently found on the island of New Guinea. Within Australia and the island of New Guinea, the records are subsequently divided by states or countries respectively. The western part of New Guinea comprises the Indonesian Provinces of Papua and West Papua, (collectively listed as Indonesia), and the eastern part is the country of Papua New Guinea (PNG).

II. INTRODUCTION The ladybirds were very familiar to the earliest European entomologists and Carolus Linnaeus (1758) was the first to recognise the genus Coccinella, describing 36 species. Since Linnaeus’ time, many new species have been described by authors from all over the world. However, it was ~100 years before two major works by the French entomologist, Etienne Mulsant (1846, 1850), set the foundations for the modern classification of the Coccinellidae. Mulsant’s publications included the first ever supra-generic categories (tribes) in the Coccinellidae, keys to genera and descriptions of all (then) known world species. In 1866 Mulsant followed with a monographic volume entirely devoted to the species of world Coccinellini. Mulsant’s monographs were revised by George Crotch (1874), who was the last author to treat the world species of ladybird beetles in a single volume. The 20th century was a period of intensive research on the Coccinellidae, starting with mostly descriptive papers by Julius Weise (1892–1927) and Albert Sicard (1907–1929) and later enhanced by many important contributions on taxonomy and higher classification of ladybirds by A.P. Kapur (1963), Hiroyuki Sasaji (1968– 1981), Ryszard Bielawski (1957–1968), Xiong-fei Pang (1979), Robert Pope (1989), Helmut Fürsch (1964–1990), Jean Chazeau (1978–1990) and Robert Gordon (1985–1987). Richard Korschefsky (1931, 1932) published the Coccinellidae portion of the ‘Coleopterorum Catalogus’, to date the only complete world catalogue of the Coccinellidae to be published. These rich traditions of taxonomic research on ladybirds have continued in Europe, Asia and the USA, with new generations of researchers contributing new data on the taxonomy, biology and evolution of ladybirds. This has culminated in many comprehensive faunal (Sasaji 1971; Pang and Mao 1979; Hoang 1982, 1983; Iablokoff-Khnzorian 1982; Gordon 1985; Kuznetsov 1997) and ecological treatments (Hodek 1973; Hodek and Honěk 1996; Majerus 1994, 2016). Chazeau (1978, 1990) treated some of the New Guinean and Pacific taxa of Coccinellini, which were then included in the book by Iablokoff-Khnzorian (1982). The Australian genera and species were treated by Pope (1989), and Ślipiński (2007), who described the larvae of almost all the Australian genera.

The tribe Coccinellini (formerly subfamily Coccinellinae) has a long history of consistent recognition as ‘genuine Coccinellidae’ since the earliest classifications of Mulsant (1850) and Crotch (1874). Mulsant (1850) split the Coccinellinae into many smaller units (among them Halyziini, Discotomini and Tytthaspidini). Sasaji (1968) resurrected the subfamily Coccinellinae in a modern sense and included the tribes Coccinellini and Halyziini. Miyatake (1972) proposed the new tribe Singhikaliini for the peculiar genus Singhikalia Kapur (1963) from Nepal and subsequently found in Taiwan, which was originally included in the subfamily Coccidulinae. Singhikalia has posed a particular challenge to classification because of its distinctly hairy body, combined with simple mandibles and pretarsal claws without a basal tooth. It was returned to Coccidulinae by Fürsch (1990) but subsequently Jadwiszczak (1990), Yu (1994) and Kovář (1996b) argued for its position in the Coccinellini, which was confirmed by the work of Ślipiński (2007) and Escalona et al. (2017). The last person who comprehensively studied taxa of the Coccinellini was Stepan M. Iablokoff-Khnzorian, who published several papers and a book on Coccinellini from the Palaearctic and Asia. Iablokoff-Khnzorian (1979, 1982, 1986) used male and female genitalia as the principal characters for generic definitions, but unfortunately neither his definitions nor illustrations were of sufficient quality to follow up his reasoning. Recent papers on molecular phylogeny of the family Coccinellidae (Giorgi et al. 2009; Magro et al. 2010; Aruggoda et al. 2010; Seago et al. 2011; Escalona et al. 2017) confirmed monophyly for the family Coccinellidae, and strongly supported a monophyletic tribe Coccinellini with the tribe Chilocorini identified as the sister taxon. Escalona et al. (2017) performed molecular analyses of a large sample of world Coccinellini and recovered three main clades among the studied taxa. The native Australo-Pacific taxa are mostly of the Oriental origin, and they represent by far the greatest diversity of Coccinellini genera and species. They have also found that the previously recognised tribes Discotomini, Halyziini, Singhikaliini, and Tytthaspidini are not valid clades and should not be retained.

III. THE COCCINELLINI 1. Morphology Adult beetle (Figs 7, 8)

The body of a true coccinellid beetle is usually oval or rounded in outline and quite convex. The dorsal surfaces are usually polished and apparently glabrous, and are only exceptionally covered with short but dense setae (e.g. the genera Subepilachna and Singhikalia) (Fig. 6F). The head is at least partly withdrawn into a prothorax and to varying extent covered from above by the pronotum. The articulation between the prothorax and the elytral bases is always broad and rigid. Head. The head is usually flattened and directed anteriorly (prognathous). Many important characters are situated on the head and its appendages. The size of the eyes and their separation can vary between genera and species but in most Coccinellini the eyes are large and occupy a large portion of head. The anterior margin of the eye is slightly indented by a cuticular process (ocular canthus). Eye facets are usually fine and flat in most groups, but in some taxa (e.g. Illeis) the eye facets are coarse and convex. The interocular distance is the minimum distance between the inner orbits of the eyes. The width of the head is defined as the maximum width with the eyes included. The clypeus in the Coccinellini is defined as a structure in front of the frons, below the lower margins of the eyes. It is usually short and straight between two lateral projections (Fig. 13B), rarely distinctly arcuate anteriorly (e.g. Synona, Fig. 13A). The antennae are relatively long, 10 or 11-segmented, each usually with distinct antennal club. The mouthparts of the Coccinellini are fairly uniform, some showing slight modifications or adaptations to the particular habits of a group, for example predation, fungivory or herbivory. The labrum is a simple transverse sclerite about as wide as the clypeus. Both mandibles are opposable and mostly symmetrical, and the basal molar area is concave and is armed with a tooth. The mandibular apex is often bidentate, with the inner, ventral tooth slightly or distinctly below the apex. In the mould-feeding genera of Coccinellini, there are additional minute denticles or teeth at either the apex or along the incisor edge of the mandible (Fig. 11M). The maxilla is a large, freemoving structure comprising a subquadrate or transverse cardo and elongated stipes. The stipes bears a four-segmented maxillary palp, a broad apically setose galea and a much smaller lacinia. The shape of the terminal palpomere ranges from weakly to strongly securiform (= axe-shaped; Figs 11E,F). The labium consists of a broad, mentum that narrows posteriorly, articulated with a solidly fused gula-submentum. The labial palps have three segments. The ligula is usually membranous and expanded anteriorly and laterally. The prothorax is always transverse with the head partially withdrawn inside it. In some Coccinellini (e.g. Microcaria salomonensis), it is withdrawn so far that the head is barely visible from above (Fig. 95B). The pronotal disc is convex, with or without punctures, but otherwise devoid of sculpture. Anterior margin of the pronotum is usually emarginate with anterior angles projecting forward weakly. There is a bordering line on the lateral margin of the pronotum, forming a lateral bead. In many genera the lateral borders are weakly to moderately explanate and usually upturned, forming a shallow gutter. The ventral surface of the prothorax consists of the deflexed sides of the notum, called the hypomere (singular, hypomeron) and a central part called the prosternum (Fig. 8B). This is separated

from the hypomeron by a notosternal suture. The prothoracic part of the hypomeron often bears the foveae near the anterior angles, seen as crescent-shaped grooves near the junction of the hypomeron and the anterior margin of the prosternum (Figs 11A–C). The prosternal sclerite lies between and in front of the coxae. It is usually moderately long and T-shaped, with broad lateral wings that meet the hypomeron laterally. The prosternal process normally has parallel sides and is rounded apically. Its dorsal surface often has a pair of carinae. The lengths of these vary considerably. The procoxal cavity is always open externally and closed internally. The procoxa is transverse, has a long internal extension and a hidden trochantin and is mostly flat. Pterothorax. The mesoventrite is always short. Its anterior edge is on a much lower plane than its posterior edge and it slopes steeply up and back towards the mesocoxae. The lower part of the mesoventrite forms a supporting collar behind the procoxae. It often bears a distinct fossa in its centre to receive the tip of the prosternal process (Fig. 11N). The front of the raised portion of the mesoventrite is straight or variously emarginate and often bordered entirely by a weak carina. The outline of the mesocoxal cavity is always circular. It is open externally, with the triangular mesepimeron closing the cavity laterally. This completely separates the meso- and metaventrites. The mesotrochantin is always hidden. The meso-meta ventral junction is usually flexible, with both sclerites meeting along a straight line. The metaventrite is flat or slightly convex, often with a medial discrimen and always with metaventral postcoxal lines. The metaventral postcoxal lines usually join in the middle; laterally the line is curved, making a distinct arch behind the posterior margins of the mesocoxae and descending with its end being the lowest point located far from the mesocoxae (Fig. 7B). The metanepisternum is narrow and often vertical but clearly visible from below. Alternatively, the metepimeron is very short and often barely visible posteriorly where it articulates with the hind coxae. The hind coxae are transverse and flat and are separated by the intercoxal process of abdominal ventrite 1. Together, the bases of the elytra are usually slightly broader than the pronotum (Fig. 7C). In winged forms the elytra are convex at the humeral angles. Their dorsal surfaces are mostly irregularly punctured, but sometimes appear almost smooth and polished. The epipleuron is an inflexion of the lateral edge of the elytron. It is defined externally by a longitudinal ridge. Internally, near the base, it is quite broad, gradually narrowing towards the apex. The epipleural surface between the inner edge and the external ridge may be completely flat, or it may descend towards the apex or even disappear altogether before it reaches the apex. In many taxa the epipleuron bears shallow foveae housing the tips of the femora. The scutellum (Fig. 7A) is always exposed and usually triangular, pointed posteriorly. Its width compared to the maximum width of the elytra is used as a generic character for the genus Micraspis, where the scutellum is exceptionally small. Functional wings are always present in Coccinellini. They have strongly simplified cucujoid venation (Lawrence et al. 2011) with a radial cell and the medial fleck always absent; there are usually two cubito-anal cells and an anal lobe. All three pairs of legs are similar. The femora and tibiae are usually slender and not flattened. The tibial apex of the mid and hind legs may have paired tibial spurs protruding from a fringe of apical setae (Fig. 11N). The tarsus is always four-segmented

III. The Coccinellini

with tarsomeres 1 and 2 lobed below. Tarsomere 3 is minute and often very hard to distinguish from 4. The tarsal claws are moderately long and usually have a basal tooth (appendiculate; Fig. 11O). Sometimes there is a subapical tooth or the claw is split apically (Fig. 11P). Abdomen and terminalia. The abdomen of all Coccinellini consists of five or six visible sternites called ventrites; two basal ventrites are connate and the following three to four are freely articulated. Ventrite 1 is usually the longest and has a pair of abdominal postcoxal lines of various form and length. The abdominal postcoxal lines may be associated with an oblique postcoxal line (Figs 11K,L). On the dorsal side of the abdomen tergites 7 and 8 are lightly sclerotised. The terminal segments of the abdomen consist of segments 9 and 10, forming genital capsule to which the genitalia proper is attached (Fig. 8C). Male genitalia. The male genitalia (Figs 8D–F) comprise a complex tegmen and relatively simple and sclerotised penis (sipho). The penis is usually enlarged at the base forming a capsule (Fig. 8D). The tegmen (Figs 8E,F) bears a ring-like phallobase that projects into a medial structure called the penis guide. The base of the guide articulates with the tegminal strut (trabes) and the apically setose parameres. Female genitalia. The ovipositor consists of a pair of lateral paraprocts (sternite IX) and a pair of coxites. These usually bear narrow apical styli. On the dorsal side of the ovipositor there is an additional sclerite called a proctiger. The female genitalia of all examined Coccinellini bear membranous colleterial glands (Fig. 8A), associated with the ventral or lateral side of each coxite. Internally, the female genital tract consists of a distally enlarged vagina. This forms a membranous bursa copulatrix at the apex that receives the common oviduct and the sperm duct. The spermatheca (sperm reservoir) is usually a C-shaped structure connected to the bursa copulatrix through the long and membranous sperm duct. In some groups, the sperm duct widens into a tube called the infundibulum (Fig. 8B). In Coccinellini the spermatheca is often divided into a basal nodulus connecting to the sperm duct, a swollen ramus (attachment of the accessory gland), and a curved apical portion at the apex called the cornu.

Larva (Figs 9A–H)

The mature larvae of Coccinellini (Gage 1920; Kamiya 1965a; Rees et al. 1994; Ślipiński 2007) are elongate fusiform with well-developed legs (Fig. 9A). The body is without wax covers and has a granulose or spinulose integument, with more or less distinct sclerotised plates developed on most of the thoracic and on smaller parts of the abdominal tergites. Dorsal sclerotisations are usually associated with complex, cuticular armature (Fig. 9C). Head. The head capsule in ladybird larvae is strongly sclerotised and rectangular or transverse. There are usually three stemmata on each side, forming a regular triangle behind the antennal foramen. Frontal arms are visible in most larvae and are lyriform (in the shape of an inverted omega symbol) and meet at their bases with a short epicranial stem, which is rarely visible in some Coccinellini. The frontoclypeal suture is always absent. The gula is broad, entirely membranous and well separated from the genae by the complete gular sutures (Fig. 9F). The hypostomal rods are usually short and parallel or weakly divergent posteriorly; epicranial ridges are absent. The antenna (Fig. 9B) is extremely variable, usually short and one- to three-segmented, with the third antennomere usually strongly reduced and shorter than the conical sensory appendix. The mandibles (Fig. 9G) are symmetrical, triangular and without asperate mola or prostheca, but usually with a distinct

5

molar projection (sometimes misinterpreted as a retinaculum), bearing rows of microtrichia. The mandibular apex is usually unidentate or bidentate, but in the mycophagous genus Illeis it bears additional denticulation along the outer edge, forming a scoop-like structure (Fig. 9H). The ventral mouth parts are retracted. The maxilla is fleshy and partially sclerotised with the cardo and stipes always fused together, without a distinct suture. The mala is blunt or narrowed apically but never narrowly falcate or angulate at the apex, without an uncus but often with a double stylus. The maxillary palps are three- or rarely two-segmented. The labium has the submentum and mentum completely fused, the prementum is sometimes distinct, but the suture is usually obliterated. The ligula is short and usually not strongly expanded. The labial palps are moderately separated, two-segmented. The hypopharynx is usually well developed, transversely spinulose and has a sclerotised transverse bridge. Thorax. The three thoracic segments are well defined and larger than the following 10 abdominal segments. The dorsal side of the segment (tergite) is weakly convex and sclerotised, forming one to several smaller plates. The prothoracic plate is usually the largest and is divided in two halves by a narrow medial line; the meso- and metathoracic plates are smaller and their admedian separation is much broader. The thoracic and abdominal segments are covered by various elements of armature, ranging from simple setae to long multiple-branched processes (Fig. 9D). The nomenclature of the armature was developed by Gage (1920) and Rees et al. (1994). The legs are well developed in the highly mobile larvae of Coccinellini, with the front legs slightly longer than the mid and hind pairs. The legs consist of five elements: the coxa, trochanter, femur, tibiotarsus and pretarsus (claw). The apex of the tibiotarsus usually has several specialised clubbed or flattened setae; the claw (Fig. 9E) bears a single seta, and at the base an appendiculate tooth that is often blunt. Abdominal segment 9 is always simple, never bearing true apical urogomphi; segment 10 forms a cylindrical pygopod surrounding the anus and is rarely exposed. The spiracles are annular, situated at the anterior end of the mesothorax and laterally on abdominal segments 1–8; the dorsal glandular openings are always absent.

Pupa (Figs 9I–K)

Body is smooth or with sparse and very short setae; colour is brown or yellow with dark maculae (Figs 9I–K). The larval exuvium, the last larval skin, folds back and covers the tip of abdomen (Fig. 9K). The wings are usually lightly sclerotised, membranous, bulbous and tapered apically; their surface is usually smooth and glabrous, rarely roughly spinulose. Abdominal pleura 3–5 are usually projecting and sometimes bearing long lateral processes (Fig. 9J); abdominal terga 4–7 are sclerotised and their much lower intersegmental membranes are exposed (Phuoc and Stehr 1974). This structure makes the pupal abdomen much more flexible, allowing it to ‘flick’ when disturbed by parasites (Majerus 1994). The sclerotised opposable edges may also serve as a ‘gin trap’, and are analogous to similar but more elaborate structures in dermestid or tenebrionid beetles used to pinch mites and other predators (Hinton 1946). Abdominal spiracles are present on terga 1–8 in all pupae. The urogomphi that anchor the pupa to the substrate within the larval skin are short and apically blunt.

2. Biology (Figs 1–6) Various aspects of the biology of ladybird beetles, mostly focussed on the relatively well-known European and North

6

Ladybird Beetles of the Australo-Pacific Region

American species of Coccinellini, are covered in several books and reviews (e.g. Clausen 1940; Hagen 1962; Hodek 1973, 1996a,b; Hodek and Honěk 1996; Kuznetsov 1997). Two books by Majerus (1994 and 2016) provide excellent summaries of the evolutionary biology of the Coccinellidae with a strong emphasis on the British fauna. On the other hand, Dixon (2000) discusses the ecology of predatory Coccinellidae and their interactions with various prey groups. Ślipiński (2007) provides a brief summary of the general biology of ladybirds relevant to the Australian fauna. The Coccinellini, like all beetles, undergo a complete metamorphosis with four discrete life stages: egg, larva, pupa and adult beetle. The larval stage takes ~14–20 days and the pupal stage a further 7–14 days to eclosion of an adult beetle. However, the length of the life cycle of ladybirds is strongly influenced by their prey and environmental factors, particularly temperature and humidity (Hodek and Honěk 1996). Adults and immatures co-occur in the same habitat and very often can be found on a single leaf with a colony of aphids. After copulation a female will lay batches of eggs on leaf surfaces close to colonies of aphids, or other preferred prey (Iperti 1999). The eggs are usually elongate-oval, yellow with smooth surfaces, and are always laid upright and very close together. Newly hatched larvae of Coccinellini often consume infertile eggs in their clutch, and eat the unhatched eggs with slowdeveloping embryos (Hagen 1962; Hodek 1967, 1973). Larvae of Coccinellini have long legs and are agile predators, quickly moving around to locate new prey. Their conspicuously coloured dorsum, with sclerotised tubercles and/or spiniferous processes, is not covered by waxes and does not have defensive gland openings on the abdomen, but some larvae are capable of excreting droplets of haemolymph (reflex bleed) on the intersegmental abdominal membranes (Nedvěd 2014). The larvae have strong, apically bidentate mandibles, and are usually capable of masticating and chewing the entire prey, differing from the many other ladybirds that use extraoral digestion to suck out the juices from prey and then discard the exoskeleton. The fungivorous larvae graze on leaves covered by mildew growth, with the help of multidentate apices of mandibles to scrape fungal tissue and conidia that are then digested internally. There are four stages (instars) in larval life, each separated by moulting periods where the larva casts off an old exoskeleton and grows a larger new one, considerably increasing its size. The final moult transforms the larva into a pupa, preceded by a quiescence period called pre-pupa. Teneral adult ladybirds emerging from the pupal skin are soft and pale, and it takes several hours for the cuticle to harden and to develop typical colour patterns. Univoltinism (one generation per year) is common in many species of Coccinellini in temperate climates, while bivoltinism is observed in those aphidophagous species that have two generations per year (spring and autumn) with a summer interval of aestivation. Multivoltinism, with at least three generations per year, has been observed in Coccinellini inhabiting warmer climates. An extreme case of this phenomenon was reported from India, where Synona melanaria feeds on the plataspid bug genus Coptosoma (see Afroze and Uddin 1998). Afroze and Uddin (1998) reported that S. melanaria can complete nine generations per year, and a female on average consumes 6600 nymphs of Coptosoma during the 46 days of its life. The Coccinellini are mostly aphid predators, but their diet is diverse and includes other hemipterous insects (heteropterans, psyllids, whiteflies), beetle and moth larvae, pollen, fungi

or even plant tissue (Schilder and Schilder 1928; Balduf 1935; Evans 2009; Giorgi et al. 2009). Pollen and nectar are consumed by various species of Coccinellini but some (e.g. Micraspis or Coleomegilla) are known to feed extensively on pollen (Hawkeswood and Turner 2002; Lundgren et al. 2005; Lundgren 2009), and are capable of completing their larval development on pollen alone (Shepard and Rapusas 1989; Lundgren and Wiedenmann 2004). Pollen grains often mixed with fungal spores are consumed by predatory species, but the members of the former tribe Halyziini (e.g. Illeis) feed exclusively on the conidia and hyphae of powdery mildews, fungi classified as Erisyphales (Ascomycota) (Anderson 1981; Sutherland and Parrella 2009). Escalona et al. (2017) analysed the food preferences of this group and concluded that there were many independent food preference transitions from ancestral aphidophagy to other food sources such as mycophagy, plant tissue, pollen and specialised predation on Acari, psyllids, plataspid bugs or larvae of chrysomelid beetles (Bashford 1999).

3. Economic importance Ladybirds were praised as beneficial insects in gardens and orchards in 19th century England (Kirby and Spence 1822), but their use as biological control agents dates from the spectacular control of the cottony cushion scale (Icerya purchasi) by the Australian vedalia beetle Rodolia cardinalis in citrus plantations in California in 1888 (Caltagirone and Doutt 1989). Since then, many coccinellid species have been tested and used as biological control agents to suppress various, usually exotic, pest species. Most of the successful applications of ladybird beetles as biological control agents have involved the control of scale insects (DeBach 1964), with R. cardinalis and Cryptolaemus montrouzieri now distributed almost worldwide. The aphidophagous Coccinellini have been less successful in biological control, being somehow harder to establish and mostly unable to effectively suppress rapidly growing aphid populations to acceptable levels. Apparently, the first species of Coccinellini intentionally used as a control agent was Coccinella undecimpunctata, introduced from Britain to New Zealand in 1874 (Majerus 2016; but see Galbreath and Cameron 2015) to control introduced European aphids. The same species and Adalia bipunctata were subsequently introduced into Australia around 1895, but the introductions were not well documented (Pope 1989). There have been several deliberate introductions of species from Australia to New Zealand with two Coccinellini, Harmonia conformis and Cleobora mellyi, established as biological control agents of psyllids, with the latter being a primary predator of a eucalyptus pest, the introduced leaf beetle Paropsis charybdis. Two Palaearctic species, Harmonia axyridis and Coccinella septempunctata, have a long history in North America as effective and desirable biological control agents of psyllids and aphids, but subsequently became large-scale invaders displacing native ladybirds and causing other environmental problems (Hodek and Michaud 2008). The harlequin ladybird beetle (Harmonia axyridis) presents a particular case of a biological control agent causing problems on a global scale (Brown et al. 2011; Roy et al. 2016; Camacho-Cervantes et al. 2017), due to the initial intentional introductions, rapid geographic range expansions and further spread via commerce. This species was recently established in New Zealand (Martin 2017) but is already very common in the Auckland area (Fig. 4B). It is unknown what the

III. The Coccinellini

environmental consequences of the establishment of H. axyridis could be in New Zealand, which has only two native species of Coccinellini, and where Coccinella leonina is already in jeopardy, being displaced by the apparently more successful introduced species Coccinella undecimpunctata (see Watt 1986). Australia has recently experienced an invasion of the Palaearctic Hippodamia variegata, an aphidophagous species

7

that was able to colonise the entire continent within 10 years of its initial discovery near Brisbane in 2000 (Franzmann 2002). The Australian biosecurity services have frequently intercepted adults of Harmonia axyridis on packing materials and external surfaces of shipping containers from shipments from Asia and Europe, but so far this species has not yet established in Australia.

IV. CLASSIFICATION OF AUSTRALO-PACIFIC COCCINELLINI Distribution of the species in the study area: AU – Australia (including Torres Strait Islands); NG – New Guinea and adjacent islands (PNG; Indonesian provinces Papua, West Papua and Maluku including the Aru and Kei Islands); NZ – New Zealand; PAC – Pacific Islands (including New Caledonia, Fiji, Tonga and Samoa); SOL – the Solomon Archipelago. Family Coccinellidae Latreille, 1807 Subfamily: Coccinellinae Latreille, 1807 Tribe: Coccinellini Latreille, 1807 ADALIA MULSANT, 1846 (1) Adalia bipunctata (Linnaeus, 1758) AU, NZ ARCHEGLEIS IABLOKOFF-KHNZORIAN, 1984b (2) Archegleis duplicata (Crotch, 1874) AU (3) Archegleis edwardsi (Mulsant, 1850) AU (4) Archegleis interrupta (Weise, 1927) AU (5) Archegleis kingi (Macleay, 1826) AU AUSTRALONEDA IABLOKOFF-KHNZORIAN, 1984b (6) Australoneda bielawskii Li, Ślipiński & Pang, 2009 NG (7) Australoneda bourgeoisi (Gadeau de Kerville, 1884) AU (8) Australoneda fuerschi (Bielawski, 1963a) NG (9) Australoneda horni (Korschefsky, 1934b) NG (10) Australoneda karubakana (Bielawski, 1963a) NG (11) Australoneda maai (Bielawski, 1963a) NG (12) Australoneda ruitong Li, Ślipiński & Pang, 2009 NG (13) Australoneda taengana (Bielawski, 1963a) NG CHEILOMENES CHEVROLAT, 1836 (14) Cheilomenes polynesiae Crotch, 1874 PAC (15) Cheilomenes samoensis Arrow, 1927 PAC (16) Cheilomenes sexmaculata (Fabricius, 1781) NG, PAC, NZ, AU, SOL CLEOBORA MULSANT, 1850 (17) Cleobora mellyi (Mulsant, 1850) AU, NZ COCCINELLA LINNAEUS, 1758 (18) Coccinella leonina Fabricius, 1775 NZ (19) Coccinella transversalis Fabricius, 1781 NG, AU, SOL (20) Coccinella undecimpunctata Linnaeus, 1758 AU, NZ COELOPHORA MULSANT, 1850 (21) Coelophora bisellata Mulsant, 1850 NG, SOL (22) Coelophora inaequalis (Fabricius, 1775) NG, AU, SOL (23) Coelophora vesiculata Chazeau, 1990 NG DOCIMOCARIA CROTCH, 1874 (24) Docimocaria aruensis (Crotch, 1874) NG (25) Docimocaria princeps (Mulsant, 1850) AU (26) Docimocaria pulchra (Crotch, 1874) NG HARMONIA MULSANT, 1850 (27) Harmonia antipodum (Mulsant in White, 1848) NZ (28) Harmonia areolata Bielawski, 1968 NG (29) Harmonia axyridis (Pallas, 1773) NZ (30) Harmonia basinotata Bielawski, 1964b NG

(31) Harmonia bicolor (Blackburn, 1892) AU (32) Harmonia boothi sp. nov. NG (33) Harmonia conformis (Boisduval, 1835) AU, NZ (34) Harmonia decussata (Crotch, 1874) NG (35) Harmonia fijiensis (Sicard, 1929a) PAC (36) Harmonia manillana (Mulsant, 1866) NG (37) Harmonia micra sp. nov. NG (38) Harmonia nigromarginata Bielawski, 1968 NG (39) Harmonia novaehebridensis (Korschefsky, 1943) PAC (40) Harmonia octomaculata (Fabricius, 1781) NG, AU (41) Harmonia ohlmusi sp. nov. NG (42) Harmonia testudinaria (Mulsant, 1850) AU, NG (43) Harmonia tricolor (Korschefsky, 1944) SOL (44) Harmonia tobleri sp. nov. SOL HETEROCARIA TIMBERLAKE, 1943 (45) Heterocaria barronensis (Blackburn, 1895) AU (46) Heterocaria bella sp. nov. AU (47) Heterocaria cinerea (Chazeau, 1990) NG (48) Heterocaria decemmaculata sp. nov. NG (49) Heterocaria delta (Weise, 1898) AU (50) Heterocaria funerea (Crotch, 1874) NG (51) Heterocaria kaszabi (Iablokoff-Khnzorian, 1982) NG (52) Heterocaria papuana Timberlake, 1943 NG (53) Heterocaria riedeli sp. nov. NG (54) Heterocaria samuelsoni (Iablokoff-Khnzorian, 1982) NG (55) Heterocaria transversoguttata sp. nov. NG HETERONEDA CROTCH, 1871 (56) Heteroneda grata (Iablokoff-Khnzorian, 1982) NG (57) Heteroneda mimetica Chazeau, 1990 NG (58) Heteroneda principalis (Weise, 1895) comb. nov. NG HIPPODAMIA DEJEAN, 1836 (59) Hippodamia variegata (Goeze, 1777) AU HYSIA MULSANT, 1850 (60) Hysia endomycina (Boisduval, 1835) NG ILLEIS MULSANT, 1850 (61) Illeis flava Pope, 1989 AU (62) Illeis galbula (Mulsant, 1850) NG, AU, NZ (63) Illeis gressitti Bielawski, 1961c NG (64) Illeis wyliei sp. nov. NG MEGALOCARIA CROTCH, 1871 (65) Megalocaria rufitarsis (Sicard, 1929b) NG (66) Megalocaria singularis (Weise, 1902) NG (67) Megalocaria tricolor (Fabricius, 1787) PAC MICRASPIS CHEVROLAT, 1836 (68) Micraspis allardi (Mulsant, 1866) PAC (69) Micraspis aphidectoides (Korschefsky, 1934a) AU (70) Micraspis bennigseni (Weise, 1903) NG (71) Micraspis confusa Pope, 1989 AU

IV. Classification of Australo-Pacific Coccinellini

(72) Micraspis flavovittata (Crotch, 1874) AU (73) Micraspis frenata (Erichson, 1842) AU, NC (74) Micraspis furcifera (Guérin-Méneville, 1835) AU (75) Micraspis lineata (Thunberg, 1781) NG (76) Micraspis lineola (Fabricius, 1775) PAC, AU MICROCARIA Crotch, 1871 (77) Microcaria jansoni (Crotch, 1874) AU (78) Microcaria mulsanti (Montrouzier, 1861) NG, PAC, AU (79) Microcaria salomonensis (Korschefsky, 1940) SOL (80) Microcaria versipellis (Mulsant, 1850) NG (81) Microcaria vivida (Crotch, 1874) PAC OENOPIA MULSANT, 1850 (82) Oenopia guttata (Blackburn, 1892) AU, NG OLLA CASEY, 1899 (83) Olla v-nigrum (Mulsant, 1866) NC, AU

9

PHRYNOCARIA TIMBERLAKE, 1943 (84) Phrynocaria ambonina sp. nov. NG (85) Phrynocaria aurita (Iablokoff-Khnzorian, 1982) NG (86) Phrynocaria astrolabiana (Weise, 1902) NG, AU (87) Phrynocaria crotchi Li, Tomaszewska, Pang & Slipinski, 2014 NG (88) Phrynocaria georgia sp. nov. SOL (89) Phrynocaria gratiosa (Mulsant, 1853) AU (90) Phrynocaria kolombangara sp. nov. SOL (91) Phrynocaria moseri (Weise, 1902) NG (92) Phrynocaria quadrivittata (Fauvel, 1903) NC (93) Phrynocaria wallacii (Crotch, 1874) NG SUBEPILACHNA Bielawski, 1963b (94) Subepilachna latemarginata Bielawski, 1963b NG SYNONA POPE, 1989 (95) Synona melanaria (Mulsant, 1850) AU, NG

V. KEYS TO IDENTIFICATION OF AUSTRALO-PACIFIC GENERA OF COCCINELLINI (FIGS 11–16) 1. – 2(1). –

Dorsum distinctly pubescent (Fig. 11A)...........................................................................................................................Subepilachna Dorsum apparently glabrous (Fig. 11B)������������������������������������������������������������������������������������������������������������������������������������������������2 Abdominal postcoxal line recurved (Fig. 11I)...................................................................................................................................... 3 Abdominal postcoxal line approaching the posterior border of ventrite 1, not recurved (sometimes with associated oblique postcoxal line – Figs 11J,K)....................................................................................................................................................5 3(2). Tarsal claws split apically (Fig. 11P); prosternal process very narrow [Australia only]....................................................Hippodamia – Tarsal claws appendiculate (with a basal tooth, Fig. 11O); prosternal process broad.......................................................................... 4 4(3). Body distinctly elongate (Fig. 11D); legs long with half of the femora visible dorsally. Anterior margins of the pronotum almost straight, anterior angles indistinct [New Guinea]...................................................................................................................Hysia – Body elongate-oval (Fig. 11C); legs mostly under the elytra, at most tips of the femora visible dorsally. Anterior margin of the pronotum deeply emarginate, with projecting anterior angles [New Zealand]..................................................................... Adalia 5(2). Maxillary palps with terminal segment exceptionally broad (Fig. 11E); mandible with denticulation along the incisor edge (Fig. 11M); anterior margin of the pronotum straight or weakly arcuate (Fig. 11Q).............................................................. Illeis – Maxillary palps with the terminal segment securiform (Fig. 11F); mandible without denticulation along the incisor edge; anterior margin of the pronotum usually deeply emarginate....................................................................................................................... 6 6(7). Hind tibiae without apical spurs............................................................................................................................................................ 7 – Hind tibiae with apical spurs (Fig. 11N)..............................................................................................................................................11 7(6). Abdominal oblique postcoxal line always present and usually connected to the postcoxal line (Figs 11K,L); terminal antennomere expanded, strongly securiform (Fig. 11H)...................................................................................................................... Harmonia – Abdominal oblique postcoxal line absent or barely visible (Fig. 11J); the terminal antennomere usually slightly wider than the penultimate and rounded apically (Fig. 11G)................................................................................................................................ 8 8(7). Prothoracic hypomera foveate anteriorly (Fig. 12B); frons very narrow, usually much narrower than eye width (Fig. 12P); elytral epipleuron with foveae receiving apices of folded legs.............................................................................................. Phrynocaria – Prothoracic hypomera flat or concave, but without delimited foveae anteriorly (Fig. 12A); frons usually much broader than eye width (Fig. 12O); elytral epipleuron without foveae receiving apices of folded legs................................................................... 9 9(8). Scutellum very small, usually less than 1/15 width of elytra at the bases................................................................... Micraspis [part] – Scutellum larger, at least 1/10 width of elytra at the bases................................................................................................................. 10 10(9). Abdominal postcoxal line very close to the posterior margin of ventrite 1, usually extending laterally to the lateral margin of the ventrite (Fig. 12Q); mesoventrite with anterior margin deeply emarginate (Fig. 12N); spermatheca weakly sclerotised (Fig. 12H) [the island of New Guinea, and the Solomon Islands]................................................................................Heterocaria – Abdominal postcoxal line short, and distinctly separated from the posterior border of ventrite 1 (Fig. 12R); anterior margin of mesoventrite straight or weakly emarginate (Fig. 12M); spermatheca sclerotised and bicameral (Fig. 12G) [Australia]............................Archegleis 11(6). Scutellum very small, usually less than 1/15 width of the elytra at the base............................................................... Micraspis [part] – Scutellum normal, usually between 1/10–1/12 as broad as elytral width........................................................................................... 12 12(11). Elytron with carina delimiting the epipleuron, extending to the apex (Fig. 12F); large ladybirds, length more than 8 mm; elytral margins broadly explanate with a strongly reduced marginal bead............................................................................................. 13 – Elytron with carina delimiting the elytral epipleuron, disappearing before reaching the apex of the elytron (Fig. 12E); smaller ladybirds, length usually less than 5 mm; elytral margins narrowly explanate and almost always with a marginal bead.......... 15 13(12). Antenna longer than head width, with a very weak club (Fig. 12J); prosternal process without distinct admedian carinae (Fig. 12A); elytra dome-shaped in lateral view, strongly explanate laterally............................................................................... Australoneda – Antenna shorter than head width, with a distinct compact club (Fig. 12I); prosternal process with distinct carinae (Figs 12B,C); elytra convex, lateral margins variously explanate...................................................................................................................... 14 14(13). Penis very long and thin, pointed apically (Fig. 12L); elytra very narrowly explanate with a weak marginal bead visible at least in the anterior corners; smaller species less than 9 mm long.......................................................................................... Docimocaria – Penis short and stout with expanded apical section (Fig. 12K); elytra broadly explanate without traces of a marginal bead; larger species, over 10 mm (except for M. tricolor from Fiji and Samoa, ~8 mm long)...................................................... Megalocaria 15(12). Mesoventrite straight anteriorly, without distinct emargination or notch (Fig. 12M)........................................................................ 16 – Mesoventrite deeply emarginate anteriorly (Fig. 12N)....................................................................................................................... 17 16(15). Elytra black with yellow markings (Fig. 13R) or orange with black markings (Fig. 13K)...................................................Coccinella – Body colour pale grey, with fine black markings (Fig. 13S) [New Caledonia, Australia].............................................................. Olla 17(15). Clypeus strongly arcuate between lateral projections (Figs 13A,C); antennal scape flat and markedly asymmetrical (Figs 13U,V)...... 18 – Clypeus straight between lateral projections (Fig. 13B); antennal scape at most convex.................................................................. 19 18(17). Elytra almost always orange or yellow, with a black colour pattern (Fig. 13H); if elytra black then the pronotum is black with yellow converging fasciae as in Fig. 13M; antenna very short with spindle-shaped club (Fig. 13V); lateral margins of elytra without a marginal bead.............................................................................................................................................................. Cheilomenes – Elytra always black (Fig. 13I); antenna slender with a broad antennal club (Fig. 13V); lateral margins of elytra with distinct a marginal bead.......................................................................................................................................................................Synona

V. Keys to identification of Australo-Pacific genera of Coccinellini

11

19(17). Small (length 3.2–4.0 mm), almost circular species with a characteristic colour pattern (Fig. 13L); abdominal oblique postcoxal line weak but always present; pronotum almost as broad as elytra at the base, widest at the posterior angles, distinctly narrowing anteriorly; mesoventrite shallowly emarginate anteriorly (Fig. 13G)..............................................................................Oenopia – Mostly larger species with the pronotum narrower at the base and elytra with different colour patterns; abdominal oblique postcoxal line absent; mesoventrite deeply emarginate anteriorly (Fig. 13F)............................................................................................20 20(19). Prosternal process without carinae (Fig. 13D) [Australia, New Zealand]............................................................................. Cleobora – Prosternal process with distinct carinae (Fig. 13E)............................................................................................................................21 21(20). Antennal club very weak (Fig. 13T) [Solomon Islands].......................................................................................... Microcaria [part] – Antennal club distinct........................................................................................................................................................................22 22(21). Prothoracic hypomera not foveate anteriorly (Fig. 13D)................................................................................................... Heteroneda – Prothoracic hypomera foveate anteriorly (Fig. 13E).........................................................................................................................23 23(22). Spermatheca with a long and irregular cornu as in Fig. 13O; penis with two hinged sclerites (Fig. 13N)....................... Coelophora – Spermatheca stout with a short cornu (Fig. 13Q); penis consists of a single medially expanded sclerite and a short apical pouch (Fig. 13O)..................................................................................................................................................... Microcaria [part]

VI. KEYS TO IDENTIFICATION OF AUSTRALIAN SPECIES OF COCCINELLINI The following key is based almost entirely on external, easily observable characters. It relies heavily on dorsal colour patterns known to occur in the species in mainland Australia. The key does not include Norfolk Island, which has a very limited fauna. It should allow for easy and correct identification of most of the species; however, some species of Phrynocaria, Microcaria

and Coelophora produce very similar colour variants that can be hard for non-specialists to distinguish. In any case, the identification obtained from this key should always be confirmed by examination of the discrete generic and species characters and, if necessary, by comparing the dissected male and female genitalia with the illustrations.

1. Elytra uniformly black (Figs 14A–H)................................................................................................................................................... 2 – Elytra with a colour pattern................................................................................................................................................................... 5 2(1). Eyes separated by less than the eye width (Fig. 11P); pronotum entirely orange (Fig. 14C) [northern Australia] ............................... �������������������������������������������������������������������������������������������������������������������������������������������������������������� Phrynocaria astrolabiana – Eyes smaller and broadly separated; pronotum bicoloured (Figs 14E–H)........................................................................................... 3 3(2). Pronotum orange or brown with admedian black fasciae (Fig. 14D); smaller species, length usually less than 6 mm......................... ������������������������������������������������������������������������������������������������������������������������������������������������������������������� Coelophora inaequalis – Pronotum black, with anterior and anterolateral areas yellow (Figs 14A,E–H); larger species, usually 5.0–6.5 mm long................. 4 4(3). Frons black (female) or bright yellow (male); if frons yellow then pronotum with extensive yellow anterolateral areas (Figs 14F,H); anterior clypeal margin strongly arcuate............................................................................................................ Synona melanaria – Frons and narrow stripe along anterior edge of the pronotum orange-brown in both sexes (Figs 14B,E,G); anterior clypeal margin straight.................................................................................................................................................................Harmonia bicolor 5(1). Large species, 8–11-mm length with characteristic colour pattern and domed elytra (Figs 14I,Q)...............Australoneda bourgeoisi – Large species, 7–10-mm length; pronotum and elytra orange red, bearing broad black margins (Fig. 14J)...... Docimocaria princeps – Usually smaller species, always with different elytral colour pattern.................................................................................................. 6 6(5). Elytra red, orange or pale yellow, with black isolated spots only (Figs 14K–P,R,S)............................................................................ 7 – Elytra with other colour pattern.......................................................................................................................................................... 15 7(6). Each elytron with a single black spot (Fig. 14P).......................................................................................................Adalia bipunctata – Elytron with multiple spots................................................................................................................................................................... 8 8(7). Elytra with 17 large spots including a scutellar and a preapical spot occupying both sides of suture (Fig. 14O)....... Harmonia conformis – Elytra with 16 spots, mostly situated at the basal half (Figs 99A–C)..............................................................................Olla v-nigrum – Elytra with less than 16 spots................................................................................................................................................................ 9 9(8). Elytra with a scutellar spot (Figs 14M,N)........................................................................................................................................... 10 – Elytra without a scutellar spot (Figs 14R,S)........................................................................................................................................11 10(9). Elytra with 11 spots (Fig. 14N); pronotum black medially but light anteriorly...................................... Coccinella undecimpunctata – Elytra with nine spots (Fig. 14M); pronotum with extensive white or yellow markings...................................Hippodamia variegata 11(9). Elytra with spots situated mostly laterally except a spot near the scutellum (Figs 14K,L)................................................................ 12 – Elytra with distinct spots at disc in posterior half (Figs 14R,S)......................................................................................................... 13 12(11). Pronotum with four or five black spots; each elytron with three small black spots and a reddish stripe along the lateral margin (Fig. 14K).............................................................................................................................................................. Archegleis kingi – Pronotum with two black spots; each elytron with six black spots, without a reddish border (Fig. 14L)............ Archegleis duplicata 13(11). Elytron with an apical spot very close to suture; pronotum usually with two dark spots (Fig. 14R)................Coelophora inaequalis – Elytron with an apical spot distinctly separated from the suture; pronotum without spots (Fig. 14S) [northern Australia].............. 14 14(13). Eyes narrowly separated (Fig. 12P).............................................................................................................. Phrynocaria astrolabiana – Eyes broadly separated (Fig. 12O)........................................................................................................................ Microcaria mulsanti 15(6). Elytra uniformly yellow (Fig. 14T)........................................................................................................................................Illeis flava – Elytra not uniformly yellow................................................................................................................................................................ 16 16(15). Elytra bright yellow, with black cross-like pattern (Fig. 14U); anterior margin of the pronotum straight; terminal maxillary palpomere very broad...................................................................................................................................................................Illeis galbula – Elytra and pronotum different to above; terminal maxillary palpomere securiform.......................................................................... 17 17(16). Each elytron with at least one continuous longitudinal black stripe or narrow fascia on disc (not counting the black stripe along the suture) (Figs 15A–J).................................................................................................................................................................... 18 – Elytra with other colour pattern than above........................................................................................................................................ 27 18(17). Eyes narrowly separated (Fig. 12P).............................................................................................................. Phrynocaria astrolabiana – Eyes broadly separated (Fig. 12O)...................................................................................................................................................... 19 19(18). Elytra with a complete stripe close to and parallel to the suture; suture blackened but without broad sutural stripe (Figs 15B,C)......... 20 – Elytra with a broader sutural stripe, not closely parallel to the suture (Figs 15A,D–J)...................................................................... 21 20(19). Elytron with a slimmer branched pattern as in Fig. 15B....................................................................................... Archegleis edwardsi – Elytron with broader and reduced stripes as in Fig. 15C..................................................................................... Archegleis interrupta 21(20). Elytron with a distinct black discal spot between the sutural and sublateral stripes (Figs 15H, 16C)................... Micraspis furcifera – Elytron without isolated discal spots.................................................................................................................................................. 22

VI. Keys to identification of Australian species of Coccinellini

13

22(19). Elytron with two stripes laterally and a third incomplete dorsal stripe connected to first lateral one (Fig. 15D)...........Heterocaria delta – Elytron with three somewhat irregular stripes (Fig. 15J).............................................................................Phrynocaria astrolabiana – Elytron with single stripe (Figs 15F,G,J) or branched stripe only (Figs 15A,E)............................................................................... 23 23(22). Discal stripe bifid from the beginning (Figs 15A,E, 16A,B)............................................................................................................. 24 – Discal stripe simple, sigmoid (Figs 15F,G) or irregular, but not bifid............................................................................................... 25 24(23). From Queensland; bifid dorsal stripe narrow, reaching the base of the elytron but not connected to the black sutural area (Figs 15A, 16B)�����������������������������������������������������������������������������������������������������������������������������������������������Micraspis confusa – From Victoria; bifid dorsal stripe broad, does not reach the base of the elytron but usually joins the black sutural area (Figs 15E, 16A)......................................................................................................................................... Micraspis flavovittata 25(22). Discal stripe on the elytra sigmoid or z-shaped (Figs 15G, 16F).............................................................................Micraspis frenata – Discal stripe on the elytra linear (Figs 15F,I, 16G)............................................................................................................................ 26 26(25). Pronotum with a black area at base limited to the middle; sutural and discal stripes on elytra very narrow (Fig. 15F) ...................... �������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Micraspis lineola – Pronotum with a black region larger and occupying most of basal area; sutural and discal stripes on elytra broad, sometimes interrupted (Fig. 15I).........................................................................................................................................Coelophora inaequalis 27(17). Body small (3.2–3.8 mm long), and rounded; elytra black with four orange spots not extending to the lateral margins (Fig. 15K) ���������������������������������������������������������������������������������������������������������������������������������������������������������������������������� Oenopia guttata – Body and elytral colour pattern different than above........................................................................................................................ 28 28(27). Pronotum amber or reddish; elytra black, with small red apical spot, and rarely additional humeral spots (Fig. 15L) ....................... ����������������������������������������������������������������������������������������������������������������������������������������������������������������������� Microcaria jansoni – Pronotum and elytral colour pattern different than above................................................................................................................. 29 29(28). Pronotum with complete medially angulate admedian stripes (Figs 15M,N); elytra with variably developed reticulate network-like colour pattern............................................................................................................................................ Harmonia testudinaria – Pronotum with a different colour pattern than above........................................................................................................................ 30 30(29). Dorsum uniformly orange or brown (Figs 15O,P)............................................................................................................................. 31 – Colour pattern different than above................................................................................................................................................... 32 31(30). Eyes narrowly separated (Fig. 12P).................................................................................................................... Phrynocaria gratiosa – Eyes broadly separated (Fig. 12O)....................................................................................................................... Microcaria mulsanti 32(30). Eyes narrowly separated, found in northern Australia (Fig. 12P)...................................................................................................... 33 – Eyes broadly separated (Fig. 12O)..................................................................................................................................................... 34 33(32). Prosternal process with converging admedian carinae....................................................................................... Phrynocaria gratiosa – Prosternal process without admedian carinae..............................................................................................Phrynocaria astrolabiana 34(32). Elytra black with 18 yellow spots, found in northern Queensland (Fig. 16U)......................................................... Heterocaria bella – Elytra different than above................................................................................................................................................................. 35 35(34). Elytra orange, each with a single medially interrupted stripe (Figs 16E,R,S)................................................................................... 36 – Elytra with other colour pattern......................................................................................................................................................... 37 36(35). Pronotum almost entirely orange, dorsal stripes not very distinct, found in the Northern Territory (Figs 16E,R)............................... ��������������������������������������������������������������������������������������������������������������������������������������������������������������� Micraspis aphidectoides – Pronotum with large black areas, elytral stripes broader and distinctly black (Fig. 16S)................................Coelophora inaequalis 37(35). Pronotum mostly black with yellow anterior margin forming converging medial branches reaching almost the middle of the ­pronotum, found in Western Australia (Figs 16 I,J)............................................................................. Cheilomenes sexmaculata – Pronotum with colour pattern other than above................................................................................................................................. 38 38(37). Elytra orange or yellow, with four transverse fasciae in addition to the black tips (Fig. 16V)...................................Cleobora mellyi – Elytra at most with three fasciae or irregular sets of spots in addition to the black tips.................................................................... 39 39(38). Body elongate-oval; elytra orange with three transverse fasciae or closely set rows of partially separate and/or interconnecting spots in addition to the usually black tips (Figs 16K,L); pronotum distinctly narrower than the elytra..........Harmonia octomaculata – Body short and more convex; elytra with at most two transverse fasciae and black apical spot (Figs 16N,T)................................. 40 40(39). Elytra with a large black spot around the scutellum (Fig. 16T); colour pattern consists of uneven transverse fasciae not extending to the elytral margins (Fig. 16H)..............................................................................................................Coccinella transversalis – Elytra variable without a distinct black spot around the scutellum; usually with a black sutural stripe separated from the scutellum (Figs 16M–Q); transverse fasciae, if present, extending to lateral margins of the elytra (Figs 16M,N).................................... 41 41(40). Eyes broadly separated; elytra bright yellow with extensive black colour pattern as in Fig. 16M; elytra with large humeral black fasciae (Fig. 16M)...................................................................................................................................Heterocaria barronensis – Eyes broadly separated; elytra orange or yellow with spots or combined spots and longitudinal fasciae (Figs 16Q,S)....................... ������������������������������������������������������������������������������������������������������������������������������������������������������������������Coelophora inaequalis – Eyes narrowly separated; elytra yellow with black colour pattern as in Figs 16N–P......................................... Phrynocaria gratiosa

VII. REVIEW OF THE AUSTRALO-PACIFIC COCCINELLINI Coccinellini Latreille, 1807: 70. Halyziini Mulsant, 1846: 147. Bulaeini Savoiskaya, 1969: 69 Discotomini Mulsant, 1850: 4 Tytthaspidini Crotch, 1874: xiii Psylloborini Casey, 1899: 73 Singhikaliini Miyatake, 1972: 96.

Diagnosis Body medium-sized or large (3–18 mm), moderately to strongly convex; dorsum almost always glabrous, smooth and shiny, usually with contrasting markings. Head: clypeus straight or rarely arcuate anteriorly between distinct lateral projections, ventral side without or with a strongly reduced antennal groove. Eyes large, mostly finely facetted, weakly emarginate anteriorly. Mandible large, triangular with a double apical tooth and sclerotised basal mola bearing a prominent tooth; maxillary palp long, terminal palpomere securiform to varying degrees. Ligula usually expanded anteriorly, labial palps slender, moderately separated at base and inserted on ventral side of the prementum. Antenna 11-segmented with distinct one- to three-segmented club. Pronotum simple. Prosternal process moderately broad, completely separating procoxae, usually with parallel carinae. Elytral punctures absent or faint and irregular; epipleuron variable usually incomplete apically, without or with weak cavities to receive the apices of the femora. Tarsi foursegmented; claws appendiculate. Abdomen almost always with six ventrites; ventrite 1 hardly longer than 2. Postcoxal line at abdominal ventrite 1 variable, sometimes with an associated oblique postcoxal line. Tegmen symmetrical, parameres well developed and multi-setose apically. Ovipositor broad and well sclerotised; coxal plates shaped like a club handle, bearing short styli; colleterial glands always present; spermatheca large and usually with well-differentiated cornu and ramus.

Review of genera and species Adalia Mulsant, 1846 (Fig. 17)

Idalia Mulsant, 1846: 44. Type species designated by Crotch 1874: 99, Coccinella bipunctata Linnaeus, 1758 (junior homonym). Adalia Mulsant, 1846: addenda et errata [p. 2] (replacement name).

References: Gordon 1985: 776; Iablokoff-Khnzorian 1982: 429; Pope 1989: 648; Poorani 2002a: 320. Diagnostic combination Length 4.0–6.7 mm, dorsum glabrous. Elytra colour pattern variable but usually red, with a single black central spot each (Fig. 17A). Antenna shorter than head width with a compact, three-segmented club. Anterior clypeal margin straight. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere strongly securiform (Fig. 17J). Prothoracic hypomeron without a fovea. Prosternal process without carinae. Anterior border of mesoventrite straight with a raised ridge. Elytral margin with a complete bead; epipleuron narrow, without foveae. Tibial spur formula 0–2–2. Tarsal claws with a large subquadrate basal tooth. Abdominal postcoxal lines separate medially, strongly recurved but incomplete;

oblique postcoxal line absent. Penis consists of a single sclerite (Fig. 17D). Infundibulum sclerotised, tubular (Figs 17H,I). Spermatheca C-shaped with well-developed ramus and nodulus.

Description Body elongate-oval, weakly convex; dorsum glabrous. Head (Fig. 17B). Eyes largely exposed with the inner orbits almost parallel. Frons about twice as broad as the eye. Antenna (Fig. 17K) 11-segmented; length 0.6–0.8 times the width of the head capsule; scape not expanded, pedicel subequal to antennomere 3; antennal club compact, three-segmented (Fig. 17K), penultimate antennomere about as long as the terminal segment; the terminal antennomere truncate or weakly arcuate apically. Anterior clypeal margin straight. Terminal maxillary palpomere strongly securiform (Fig. 17J). Mandible bifid apically, molar part with basal teeth. Prothorax. Pronotum weakly convex; the pronotal base without a marginal bead; lateral margins very narrowly upturned. Prothoracic hypomeron without foveae. Prosternal process not extending beyond the coxae, smooth without carinae. Pterothorax. Scutellum ~1/15 times as broad as elytra at the shoulders. Elytra slightly broader than the pronotum at base; lateral margin narrowly upturned but without a bead; epipleuron 1.0–1.2 times the width of the metanepisternum, obliquely inclined without delimited foveae. Mesoventrite anteriorly straight and with a complete raised ridge; mesal surface without clear articulation with the prosternal process. Mesoventral process width about half the mesocoxal diameter. Metaventral postcoxal lines joined medially, arcuate, weakly descending and complete. Middle and hind tibia each with two apical spurs; tarsal claws with a basal subquadrate tooth. Abdomen with six ventrites in both sexes. Abdominal postcoxal lines separate medially, recurved but incomplete (Fig. 17C); oblique postcoxal line absent. Male terminalia (Figs 17D–G). Penis guide slender and pointed apically; penis stout, consists of a single sclerite, expanded apically (Fig. 17E). Female terminalia (Figs 17H,I). Coxites elongate-oval, narrowing posteriorly, styli strongly reduced. Bursa copulatrix apically and tubular infundibulum strongly sclerotised. Spermathecal capsule weakly curved; ramus and nodulus distinct; spermathecal accessory gland adjacent to sperm duct.

Biology Adult and larvae of Adalia are aphid and adelgid predators, with very broad host ranges (Gordon 1985).

Distribution There are over 35 species of Adalia distributed worldwide (Gordon 1985). Six species are known from the Palaearctic region (Iablokoff-Khnzorian 1982). The Holarctic species Adalia bipunctata was introduced to most continents, including Australia and New Zealand.

Remarks Adalia is easily distinguished from the other genera of Coccinellini in the Australo-Pacific Region in having strongly recurved abdominal postcoxal lines and straight anterior margin of the mesoventrite.

VII. Review of the Australo-Pacific Coccinellini

Adalia bipunctata (Linnaeus, 1758) (Fig. 17)

Coccinella bipunctata Linnaeus, 1758: 364. Type locality: Europe. Idalia bipunctata: Mulsant 1846: 51 Adalia bipunctata: Mulsant 1846: Addenda et errata [second]; Crotch 1874: 102; Korschefsky 1932: 385; Iablokoff-Khnzorian 1982: 438; Gordon 1985: 780; Pope 1989: 649; Poorani 2002a: 320; Ślipiński 2007: 146.

Description Colour pattern (Fig. 17A). Typical form: head black, with a semicircular yellow area on either side adjacent to the inner borders of eyes; labrum black, sometimes yellowish apically. Pronotum cream-yellow with a black M-shaped area medially. Scutellum black. Elytra red, each with a single black spot near the middle. Ventral side black, except for yellow prothoracic hypomera, elytral epipleura and, sometimes, lateral and apical margins of abdominal ventrites. Legs black. Colour variation: In melanic colour forms, black colouration on the pronotum and elytra is expanded; in extreme forms, the head and pronotum are almost completely black, and the elytra are black with humeral and preapical red spots. Morphology. Eyes finely facetted. Frons flat, distinctly punctured; punctures smaller than eye facets and 1–2 diameters apart; interspaces densely reticulate, weakly shiny. Pronotum regularly punctate, the punctures similar to those on the head, the interspaces less densely reticulate, and shiny. Scutellum an equilateral triangle, at base ~1/15 times as broad as elytra at the shoulders. Elytral punctures larger and more deeply impressed than those of the pronotum, the interspaces shiny and polished. Abdomen with the posterior border of ventrite 5 straight in males, slightly arcuate medially in females. Male terminalia (Figs 17D–G). Female terminalia (Figs 17H,I). Measurements (n = 5). TL 4.60–5.30 (5.04 ± 0.32); PL 0.80– 1.00 (0.88 ± 0.08); EL 3.80–4.40 (4.16 ± 0.25); PW 2.10–2.40 (2.32 ± 0.13); EW 3.20–4.10 (3.66 ± 0.34); GD 1.70–2.10

15

(1.94 ± 0.15). Ratios: PL/PW 0.35–0.42 (0.38 ± 0.02); EL/EW 1.05–1.19 (1.14 ± 0.06); EL/PL 4.30–5.00 (4.74 ± 0.27); EL/ GD 2.05–2.24 (2.15 ± 0.07); TL/EW 1.27–1.44 (1.38 ± 0.07).

Specimens examined AUSTRALIA: Tasmania: Hobart (BMNH). Western Australia: Dongara, xii. 1935 (BMNH). NEW ZEALAND: South Island: Nelson, 20.iv.1969, J. E. Tobler (CAS); Nelson, 18.ix.1957, M. J. Esson (NZAC); CO Cromwell,1978, R. Olds (NZAC); MK Te Kohai I. L Pukaki, 27.i.1976, C. Smith (NZAC); AK Woodhill, 25.i.1978, J. Forster (NZAC); Christchurch, 25.xi.1950, V. Cottier (NZAC).

Distribution This Holarctic species was introduced to Chile, Argentina, New Zealand and Australia. It was introduced to Tasmania and Western Australia in 1901–1902 to control the cabbage aphid (Brevicoryne brassicae (L.)) but the Western Australian population did not successfully establish. Hobart had flourishing populations in cultivated areas from 1972, and abundant specimens were still seen in 1981 (Pope 1989). However, we were not able to confirm occurrence of this species in Tasmania with recently collected material and it is now considered as not established in Australia. It is a common species throughout New Zealand, represented there by several colour forms (Martin 2016a).

Archegleis Iablokoff-Khnzorian, 1984b (Figs 18–21)

Archegleis Iablokoff-Khnzorian, 1984b: 113. Type species by original designation, Coccinella kingi Macleay, 1826.

References: Pope 1989: 692; Ślipiński 2007: 149. Diagnostic combination

Length 4.8–7.3 mm; dorsum glabrous. Elytra colour pattern variable, but with a yellowish or green background and black spots or fasciae. Antenna shorter than head width; the antennal club weak and loosely articulated. Anterior clypeal margin straight. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron without a fovea. Prosternal process with weak carinae not extending to the anterior margin of the prosternum. Anterior border of the mesoventrite shallowly emarginate medially, with a raised ridge. Tibial spur formula 0–0–0. Tarsal claws with a subquadrate basal tooth. Abdominal postcoxal lines short, incomplete laterally, straight or angulate; oblique postcoxal line absent. Penis long and slender, consisting of two hinged sclerites. Infundibulum absent; spermatheca sclerotised and bicameral.

Description

Distribution of Adalia bipunctata

Body elongate-oval, moderately convex, dorsum glabrous. Head. Eyes largely exposed with the inner orbits almost parallel. Frons about twice as broad as eye width. Anterior clypeal border straight. Antenna 11-segmented, very slender, length 0.6–0.8 times the head capsule width; scape not expanded, pedicel subequal to antennomere 3; antennomeres 9–11 form a very weak antennal club; the terminal antennomere is distinctly longer than the penultimate and rounded apically. Terminal labial palpomere much slimmer and shorter than

16

Ladybird Beetles of the Australo-Pacific Region

the penultimate palpomere. Terminal maxillary palpomere moderately securiform. Mandible bifid apically; molar area with basal teeth. Prothorax. Pronotum weakly convex; the pronotal base without a marginal bead; lateral margins very narrowly upturned, without a bead. Prothoracic hypomeron without foveae. Prosternal process ~0.2 times as broad as the coxal width, with pair of weak carinae extending beyond the process but not reaching the anterior margin of the prosternum. Pterothorax. Scutellum 1/10–1/12 times as broad as the elytra at the shoulders. Elytra moderately convex, much broader than the pronotum at the base, explanate laterally with a marginal bead present at least along the basal half of elytron. Elytral epipleuron 1.2–2.1 times the width of the metanepisternum, slightly inclined, without foveae. Mesoventrite with a weak medial notch and an entirely raised anterior margin; mesal surface with a shallow fossa receiving the prosternal process. Metaventral postcoxal line joined medially, complete laterally. Middle and hind tibiae without apical spurs; tarsal claws with a large subquadrate basal tooth. Abdomen with six ventrites in both sexes. Postcoxal lines separate medially, arcuate, short and incomplete laterally and not reaching the posterior margin of the ventrite; the oblique postcoxal line almost always absent (visible in some specimens of A. interrupta; Fig. 20D). Male terminalia. Penis slim and much longer than abdomen length, consists of two hinged sclerites; capsule with weak inner and outer arms; penis guide strongly curved apically; parameres slim, bearing long and dense hairs apically. Female terminalia. Coxites slim and long, broad at the base with distinct styli; infundibulum absent; bursa copulatrix long. Spermatheca entirely sclerotised and bicameral, with the ramus and nodulus forming two balls connected by a narrow stalk; cornu absent.

Biology The specimen label data associated with various species (Pope 1989) indicate aphids and other Hemiptera are the primary food source of Archegleis. Adults are usually collected from flowering bushes and trees and on light. Ślipiński (2007) described the larva and pupa of A. kingi and A. edwardsi based on specimens reared at QDPI Brisbane by K. J. Houston, using Hyperomyzus aphids on Sonchus.

Distribution Endemic to northern and eastern Australia.

Remarks Six Australian species of Archegleis were reported by Pope (1989), who noticed two different types of male genitalia within the genus. The division in the penis morphology is paralleled by even more pronounced differences in structure of spermatheca in female terminalia of some species, and the genus Archegleis is here restricted to four Australian species. Typical Archegleis can be diagnosed by the penis, consisting of two hinged sclerites, and the bicameral, entirely sclerotised, spermatheca in female terminalia. Two species of Archegleis not conforming to the definition above (A. barronensis and A. delta) are here transferred to closely related Papuan genus Heterocaria Timberlake (= Oiocaria Iablokoff-Khnzorian). Both Heterocaria and Archegleis are externally very similar in appearance. In the distinguishing characters given in the key to genera, the shorter abdominal postcoxal lines in Archegleis are not easy to observe and may prove to be variable if more species are discovered.

Key to species of Archegleis 1. – 2(1). – 3(1). –

Elytral pattern of round black spots on a pale background (Fig. 18A)................................................................................................ 2 Elytral pattern of black stripes on a pale background (Figs 19A, 20B)............................................................................................... 3 Pronotum with 4 or 5 black spots; each elytron with 3 small black spots and a reddish stripe along lateral margin (Figs 21A–C)..... ������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������A. kingi Pronotum with 2 black spots; each elytron with 6 black spots, without a reddish border (Figs 18A–C).......................... A. duplicata Elytra with slim black stripes forming a pattern as in Fig. 19A..........................................................................................A. edwardsi Elytra with distinctly broad stripes forming a pattern as in Fig. 20B...............................................................................A. interrupta

Archegleis duplicata (Crotch, 1874) (Fig. 18)

Callineda duplicata Crotch, 1874: 161. Type locality: N Australia. Korschefsky 1932: 282. Harmonia duplicata: Iablokoff-Khnzorian 1982: 461. Archegleis duplicata: Pope 1989: 695 (lectotype); Ślipiński 2007: 151.

Diagnosis Very similar to A. kingi, but both species can easily be distinguished by the number of dark spots on the pronotum and elytra, and the lack of reddish elytral fascia along the external border in A. duplicata.

Description Colour pattern (Figs 18A–C). Head and appendages yellow. Pronotum peach-yellow, with two black elongated oval spots

close to the base. Scutellum pale with black lateral borders. Elytra pale, testaceous with six small black spots, suture and external borders black. Ventral side body yellow, except for the metaventrite which ismostly black. Legs yellow. Morphology. Length 5.6–6.0 mm. Frons flat, width ~1.8–2.0 times the eye width, distinctly punctured and setose. Pronotum moderately convex, ~0.6 times as broad as elytra at the shoulders. Prosternal process with longitudinal carinae slightly convergent apically, extending anteriorly to the middle of the prosternum. Scutellum at base 1/12 times as broad as the elytra. Elytra convex, broadest in anteriorly; punctures larger than eyes’ facets; epipleuron broad, ~1.8–2.0 times as wide as the metanepisternum. Abdominal postcoxal line short and weakly arcuate (Fig. 18D). Posterior margin of ventrite 5 emarginate in male, broadly arcuate in female. Male terminalia (Figs 18E,F,I). Female terminalia (Figs 18G,H,J). Measurements (n = 2): TL 5.60–5.90; PL 1.00–1.20; EL 4.00–5.30; PW 2.90–3.10; EW 5.00–5.20; GD 2.60–2.70.

VII. Review of the Australo-Pacific Coccinellini

17

Description

Distribution of Archegleis duplicata

Ratios: PL/PW 0.34–0.39; EL/EW 0.80–1.02; EL/PL 4.00– 4.42; EL/GD 1.54–1.96; TL/EW 1.08–1.18.

Types C. duplicata: Type/ Callineda duplicata, (Type) Crotch/ duplicata (BMNH, lectotype).

Specimens examined AUSTRALIA: Western Australia: 14.38°S 125.48°E, Mitchell Plateau, ex. Acacia sp. beating flowers, 4.v.1992, P.J. Gullan (4, ANIC).

Distribution Archegleis duplicata is known only from the type specimen and four specimens collected in the Kimberley region of Western Australia.

Remarks The taxonomic status of this species remains unclear, as the male and female genitalia are almost identical in its eastern vicariant (A. kingi), but the differences in external colouration between the two populations are very consistent.

Colour pattern (Figs 19A–C). Body greenish-yellow with black markings. Frons and head appendages yellow, vertex with two transverse black marks. Pronotum yellow, with black stripes medially. Scutellum black. Elytra greenishyellow with longitudinal and oblique black stripes, suture and external borders black. Prothoracic hypomeron yellow with a black patch posteriorly. Prosternum yellow, with black external borders in males and completely black in females; the mesoventrite black in females and black with yellow centre in males; mesanepisternum and mesepimeron pale; metaventrite and metanepisternum black; abdomen mostly yellow. Legs mostly yellow, partially black. Morphology. Length 5.0–6.7 mm. Frons width ~2.0–2.3 times the eye width, flat, distinctly punctured and setose. Pronotum moderately convex, ~0.7–0.8 times as broad as elytra at the shoulders. Prosternal process with longitudinal carinae parallel, extending to the middle of the prosternum. Scutellum 1/14 times as broad as elytra at the humeri. Elytra convex, broadest anteriorly; punctures larger than facet of the eyes; epipleuron broad, ~2.0–2.2 times as wide as the metanepisternum. Abdominal postcoxal line short and sometimes angulate (or with faint oblique postcoxal line, Fig. 19D). Posterior margin of ventrite 5 truncate in male, broadly arcuate in female. Male terminalia (Figs 19E, G–I). Female terminalia (Figs 19F, J–L). Measurements (n = 8): TL 5.00–6.70 (5.88 ± 0.48); PL 1.00–1.10 (1.05 ± 0.05); EL 4.00–5.60 (4.79 ± 0.46); PW 2.50– 3.330 (2.88 ± 0.24); EW 4.10–5.30 (4.68 ± 0.41); GD 2.00–2.90 (2.44 ± 0.27). Ratios: PL/PW 0.33–0.41 (0.37 ± 0.03); EL/EW 0.98–1.12 (1.02 ± 0.05); EL/PL 4.00–5.09 (4.59 ± 0.42); EL/ GD 1.84–2.14 (1.97 ± 0.08); TL/EW 1.20–1.40 (1.26 ± 0.06).

Types E. edwardsi: ‘Museum Paris, Australia 2–47/ 670/ 57/ 2.47/ Egleis edwardsi Muls. auct. det.’ (lectotype, MNHN). E. pascoei: ‘Type Pascoei/ TYPE’ (lectotype, UCCC).

Specimens examined AUSTRALIA: Queensland: Lamington NP, 20.v.1964, D.F. O’Sullivan (1, UQIC); Saddletree Creek, via Maidenwell, 12– 14.ii.1976, A. Postle (1, UQIC); Bunya Mts, 5.vi.1959, I. C.

Archegleis edwardsi (Mulsant, 1850) (Fig. 19)

Egleis edwardsi Mulsant, 1850: 158. Type locality: Australia. Weise 1898: 226; Korschefsky 1932: 576; Timberlake 1943: 27. Egleis pascoei Crotch, 1874: 131. Type locality: Australia. Synonymised by Weise 1898: 226. Halyzia pascoei: Masters 1888: 90; Lea 1902: 490. Archegleis edwardsi: Pope 1989: 701 (lectotypes); Ślipiński 2007: 151. Archegleis delta: Iablokoff-Khnzorian 1984b: 117 [misidentification].

Diagnosis Archegleis edwardsi has a similar yellowish-green background and black fasciae to those of A. interrupta and Heterocaria delta, but each of these species can be easily distinguished by a slightly different pattern of black stripes and spots (see Figs 19, 20, 65).

Distribution of Archegleis edwardsi

18

Ladybird Beetles of the Australo-Pacific Region

Yeo (1, UQIC); Cunningham’s Gap, 2484 ft., 1–2.vi.1966, Z. Liepa (1, ANIC); Cunningham’s Gap, xi.1953, JTA (1, ANIC); National Park Macpherson Range, i.1928, H.J. Cater (2, ANIC); D’Aguilar NP., 7.viii.2004, M. Wanat (2, ANIC). New South Wales: Audley NP, 33 mi. S. Sydney, 11.xii.1960, J. Schenk (1, ANIC); Mt. Keira, 1.ix.1979, G.A. Holloway (1, AM); Mt. Tomah, Blue Mountains, 22.ii.1992, N. W. Rodd (1, ANIC); 1km W Barrengarry, on Ac. melanoxylon flower, 27.ix.1986, C. Reid (1, ANIC); 28.29S 152.23E, Beaury S.F., 700M, 15–17. ii.1983, T. Weir & A. Calder (1, ANIC); Eden, E.W. Ferguson (2, ANIC); Richmond R., 1909–174 (1, ANIC); Rivertree, E. Sutton (1, ANIC); Long Flat, J. Armstrong (1, ANIC); Wauchope (3, ANIC); Putty Road, Colo River crossing, on flowers of Leptospermum, 26.xi.1994, A. Sundholm (1, ANIC).

Distribution The known distribution of this species is limited from southern Queensland to Eden in New South Wales. Several undated records from Victoria have not been confirmed with new material and appear to be beyond natural range of the species (Pope 1989).

Archegleis interrupta (Weise, 1927) (Fig. 20)

Egleis interrupta Weise, 1927: 12. Type locality: Queensland, Cedar Creek. Bielawski 1965: 233. Archegleis interrupta: Iablokoff-Khnzorian 1984b: 115; Pope 1989: 700 (lectotype); Ślipiński 2007: 151.

Diagnosis Archegleis interrupta can be distinguished from similar species A. edwardsi and Heterocaria delta by having a very broad and complete first stripe along the suture that continues along the lateral margin of the elytron (Figs 19, 20, 65).

Description Colour pattern (Figs 20A–C). Body greenish-yellow with a black pattern. Head pale yellow with two small black marks across the vertex. Scutellum black. Elytra greenish-yellow, with black pattern, suture and external borders are black. Prosternum pale, with the anterior and posterior borders black in males, completely black in females. Mesoventrite, mesepimeron and mesanepisterna pale; metaventrite, metepimeron and metanepisterna dark. Anterior and middle legs pale with some dark areas, hind legs dark. Elytral epipleuron yellow, except for narrow, black internal and external borders. Morphology. Length 5.0–5.9 mm. Frons width about twice the eye width, flat, distinctly punctured and setose. Pronotum moderately convex, ~0.7–0.8 times as broad as elytra at the shoulders. Prosternal process with longitudinal carinae parallel, extending forward to the middle of the prosternum. Scutellum at the base 1/16 times as broad as the elytra. Elytra convex, broadest anteriorly; punctures larger than eye facets; epipleuron broad, ~2.0–2.1 times as wide as the metanepisternum. Abdominal postcoxal line short and not recurved (Fig. 20D). Posterior margin of ventrite 5 truncate in males, broadly arcuate in females. Male terminalia (Figs 20E–G). Female terminalia (Figs 20H–J). Measurements (n = 4): TL 5.00–5.90 (5.42 ± 0.33); PL 0.90– 1.10 (1.01 ± 0.06); EL 4.00–4.80 (4.39 ± 0.28); PW 2.60–3.00

Distribution of Archegleis interrupta

(2.81 ± 0.18); EW 4.10–4.80 (4.51 ± 0.25); GD 2.00–2.60 (2.34 ± 0.18). Ratios: PL/PW 0.33–0.38 (0.36 ± 0.02); EL/EW 0.91–1.02 (0.97 ± 0.04); EL/PL 4.00–4.70 (4.35 ± 0.23); EL/ GD 1.78–2.00 (1.88 ± 0.07); TL/EW 1.13–1.27 (1.20 ± 0.04).

Types E. interrupta: Cedar Creek/ Queensl. Mjoberg/ mars/ Egleis interrupta m (NRMS, lectotype).

Specimens examined AUSTRALIA: Queensland: 17.19S 145.29E, Wongabel SE, botanic walk, beating trees and bushes, 12.vi.1992 & 8.xi.1992, C. Reid (5, ANIC); 17.10S 145.39E, 1 km N Cathedral Fig., beating trees/bushes by stream, 8.xi.1992, Danbulla SF, C. Reid (2, ANIC); Yungaburra (state Forest 452), 29.iv.1967, D.H. Colless (1, ANIC); Lake Barine, 530m, 1.ii.1964, J. Sedlacek (2, BPBM); Cairns distr., J.A. Anderson (1, ANIC); 7 mi. N.E. Ravenshoe, 975m, 7.xi.1962, E.S. Ross & D. Q. Cavagnaro (2, CAS).

Distribution This species is known only from the Atherton Tableland in northern Queensland.

Archegleis kingi (Macleay, 1826) (Fig. 21)

Coccinella kingi Macleay, 1826: 454. Type locality: Australia. Mulsant 1950: 1049; Crotch 1874: 310; Masters 1888: 89; Blackburn 1892: 238; Lea 1902: 488. Egleis kingi: Blackburn 1892: 238; Korschefsky 1932: 576; Timberlake 1943: 27. Archegleis kingi: Iablokoff-Khnzorian 1984b: 115; Pope 1989: 694 (lectotype); Ślipiński 2007: 151. Egleis varicolor Mulsant, 1850: 154. Type locality: Australia. Mulsant 1866: 120; Crotch 1874: 131. Synonymised by Blackburn 1892: 238. Halyzia varicolor: Masters 1888: 90.

Diagnosis Archegleis kingi is very similar to A. duplicata in size and in general colouration, but the two species are readily distinguished by different colour patterns on the pronotum and elytra, and

VII. Review of the Australo-Pacific Coccinellini

19

Queensland: Port Denison (2, ANIC); Mulgowie,16.ix-7.xi.1983, M.D. Peart (8, ANIC); Pine Creek near Bundaberg, 10.xii.1975, H. Frauca (1, ANIC); 25.48S 148.15E, Great Diving Range, 600–650 m, 35 km NW Injune, road to Mt. Moffatt, 3.iv.2002, A. Steiner & A. Zwick (1, ANIC); 26.50S 151.33E, Burtons Well, Bunya Mts Nat. Pk, 6–7.x.1984, I. Naumann & J. Cardale (2, ANIC); Cemetery Point, Vernon via Maryborough, 24.xii.1970, I.F.B. Common (1, ANIC); 19 km S. of Bundaberg, Gorge at Burnett R., 20–24.xi.1975, H. Frauca (1, ANIC); Herberton, 12.vi.1992, C. Reid (1, ANIC); Brisbane,13.i.1965, E. B. Tay (1, ANIC); Brisbane, 6.iii.1932, Wassell (2, ANIC). New South Wales: 11km SSE Narrandera, 2.x.1990, C. Reid (1, ANIC); University Armidale, 27.xi.1971, C. W. Frazier (1, ANIC).

Distribution Distribution of Archegleis kingi

A. kingi having distinct reddish fascia along the external elytral margin.

Description Colour pattern (Figs 21A–C). Head yellow, except for two transverse black markings on the vertex; appendages yellow, terminal palpomere slightly darker. Pronotum pale yellow with four or five black markings medially. Scutellum black, except for a small pale area at the base. Elytra creamy-yellow with a reddish stripe along the lateral border and three rounded black spots on each elytron (spot near the scutellum often strongly reduced or absent); the suture and external borders are narrowly black. Ventral side yellow, except for a black metaventrite. Legs with femora and tibiae grey, partially blackened. Morphology. Length 4.8–7.3 mm. Frons width 1.5–1.8 times the eye width, flat or weakly concave, distinctly punctured and setose. Pronotum moderately convex, ~0.7 times as broad as elytra at the shoulders. Prosternal process with longitudinal carinae slightly convergent apically, extending close to the anterior border of the prosternum. Scutellum at the base 1/12 times as broad as elytra at the shoulders. Elytra convex, broadest anteriorly; discal punctures larger than eyes’ facet; epipleuron broad, about twice as wide as the metanepisternum. Abdominal postcoxal line is short and weakly arcuate (Fig. 21D). Posterior margin of ventrite 5 truncate in male, broadly arcuate in female. Male terminalia (Figs 21E–G). Female terminalia (Figs 21H–J). Measurements (n = 12): TL 4.80–7.30 (5.88 ± 0.68); PL 0.80–1.20 (0.97 ± 0.12); EL 4.00–6.10 (4.95 ± 0.60); PW 2.30– 3.50 (2.95 ± 0.32); EW 3.90–6.00 (4.91 ± 0.58); GD 2.10–2.90 (2.43 ± 0.23). Ratios: PL/PW 0.30–0.37 (0.33 ± 0.02); EL/EW 0.94–1.08 (1.01 ± 0.04); EL/PL 4.50–5.70 (5.13 ± 0.35); EL/ GD 1.88–2.26 (2.04 ± 0.12); TL/EW 1.12–1.26 (1.20 ± 0.04).

Types C. kingi: ‘Coccinella Kingii McL, Capt. King Australia’ (lectotype, ANIC). E. varicolor: ‘TYPE/ Type varicolor Reich’ (holotype, UCCC).

Specimens examined AUSTRALIA: Northern Territory: Hamilton Downs Station, 60km NW Alice Springs, 9.vi.1976, G. Griffin (2, ANIC).

This species is found from Atherton and Kuranda in northeastern Queensland to Armidale and Narrandera in northern New South Wales. The inland locality near Alice Springs is probably an accidental introduction or temporary extension of the normal range of that species (Pope 1989).

Australoneda Iablokoff-Khnzorian, 1984 (Figs 22–29)

Australoneda Iablokoff-Khnzorian, 1984b: 107; Type species, original designation, Neda bourgeoisi Gadeau de Kerville, 1884;

References: Bielawski 1963a: 469; 1965: 229; IablokoffKhnzorian 1984b: 107; Li et al. 2009: 223. Diagnostic combination Length 8–12 mm; dorsum glabrous. Elytra colour pattern variable, from uniformly brown to black with a yellow or red pattern. Antenna about as long as or slightly longer than head width; antennal club weak and loosely articulated. Anterior clypeal margin straight. Mandible bifid apically, molar part with a basal tooth. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron without a fovea. Prosternal process extending behind the procoxae, pointed apically with lateral margins; carinae sometimes continue anteriorly. Anterior border of the mesoventrite deeply emarginate medially. Tibial spurs very short, formula 0–2–0. Tarsal claws with a subquadrate basal tooth. Abdominal postcoxal (Fig. 23I) lines long, complete laterally, very close to the posterior margin of ventrite 1; oblique postcoxal line absent. Penis long and slender, consisting of two hinged sclerites. Infundibulum absent; spermatheca moderately sclerotised, C-shaped.

Description Body broadly rounded, disc-shaped, strongly convex and usually pointed in profile; dorsally glabrous; elytral margins very broadly explanate. Head. Eyes only partially exposed. Frons 1.5–1.7 times broader than the eye. Antenna 11–segmented; length 1.0–1.3 times the width of the head; scape not expanded; pedicel shorter than antennomere 3; antennal club weak, three-segmented, terminal antennomere arcuate apically. Anterior clypeal margin straight. Terminal maxillary palpomere distinctly securiform. Mandible bifid apically, molar part with basal teeth. Prothorax. Pronotal disc broadly, evenly convex; hypomera without delimited foveae; prosternal process ~0.5 times width of coxal diameter, triangular apically with carinate borders and rarely with the carinae extending in front of the procoxae.

Ladybird Beetles of the Australo-Pacific Region

20

Pterothorax. Mesoventrite deeply excavate to accommodate the apex of the prosternal process. Elytra convex, raised medially, much broader than the pronotum at the base, broadly explanate laterally, without a marginal bead. Elytral epipleuron very broad, slightly inclined, without foveae. Metaventral postcoxal line complete, descending. Legs usually long, slender, tibial spurs formula 0–2–0 in all species. Abdomen with six ventrites in both sexes; ventrite 6 partially or fully hidden in females; apical margin of ventrite 6 in males emarginate medially, while impressed and evenly arcuate in females. Abdominal postcoxal line not recurved, almost reaching the lateral margin of the first ventrite, posteriorly very close to the hind margin of the ventrite; transverse postcoxal line absent. Male terminalia. Penis simple, consisting of two hinged sclerites; penis capsule short. Penis guide usually shorter, or as long as parameres, with a hook at the apex in most species. Female terminalia. Coxites large and sclerotised with subterminal styli; sperm duct length variable, but without infundibulum; spermatheca large and usually C-shaped, always

with deep spiral lines on surface; nodulus indistinct; ramus short with spermathecal gland.

Biology Nothing is known on biology of several species from the island of New Guinea (Figs 1A–C). The single Australian species, Australoneda bourgeoisi, has been collected regularly on leaves of small eucalypts close to rainforest edges. It was reared in captivity on Hyperomyzus aphids on Sonchus sp. by K. J. Houston at QDPI Brisbane (Pope 1989). Ślipiński (2007) described and illustrated larva of A. bourgeoisi.

Distribution Island of New Guinea and north-eastern Australia.

Remarks Australoneda is a very distinctive genus, easily recognised by its large, almost circular body, with elytra domed and bearing strongly explanate margins.

Key to the species of Australoneda 1. – 2(1). – 3(2). – 4(1). – 5(4). – 6(4). – 7(6). –

Explanate, lateral part of elytron with red or yellow spots (Fig. 26A)................................................................................................ 2 Explanate, lateral part of elytron without colour spots (Fig. 22A)...................................................................................................... 4 Elytra weakly convex, regularly rounded, not pointed in lateral view (Fig. 28B)................................................................ A. ruitong Elytra strongly convex, in lateral view pointed at distal 1/3 length (Fig. 29A)................................................................................... 3 Elytron (Figs 26A–C) with a large triangular spot adjacent to the scutellum, and usually with a much larger apical spot (Fig. 26B); pronotum almost always with a small black spot at the centre of each large yellow lateral marking (Figs 26A–C); elytra strongly convex and pointed in lateral view; male and female terminalia as in Figs 26D–H............................... A. karubakana Elytron at most with a very small yellow spot near scutellum (Fig. 29B); pronotum without a black spot in the yellow lateral marking (Fig. 29A); elytra moderately convex and weakly angulate at highest point (Fig. 29A); male and female terminalia as on Figs 29C–G................................................................................................................................................................. A. taengana Elytra black with isolated spots (Figs 25, 26, 27)................................................................................................................................ 5 Elytra yellow or reddish, without distinct spots (Fig. 24).................................................................................................................... 6 Elytral humeral spots large and orange and yellow, almost extending to the suture; apical spot triangular, large (Fig. 25A)...... A. horni Elytral humeral and apical spots small, yellow, and situated near the lateral margin only (Fig. 27B)..................................... A. maai Body yellow or brown, without black markings (Fig. 24A)................................................................................................A. fuerschi Pronotum and elytra with black margins (Figs 22A, 23A).................................................................................................................. 7 Elytral disc yellow with black colour limited to a very narrow lateral margin and fine line separating the disc and explanate margins (Fig. 22A)...................................................................................................................................................................A. bielawskii Elytra disc orange and yellow with black colour forming very broad lateral margins (Fig. 23A).................................. A. bourgeoisi

Australoneda bielawskii Li, Ślipiński & Pang, 2009 (Fig. 22)

Australoneda bielawskii Li, Ślipiński & Pang, 2009: 228. Type ­locality: Western New Guinea: Testega, Meydoudga.

Diagnosis Due to the yellowish-brown body and non-maculate pronotum and elytra, this species is most similar to A. maai and, to a lesser extent, A. bourgeoisi. Australoneda bielawskii differs from both these species by a presence of a small black spot on the lateral pronotal areas. The dark longitudinal pronotal markings and narrow, black margins of the elytra distinguish it further from A. maai. Australoneda bourgeoisi has elytra with very broad black margins and a pronotum usually with dark areas connected into a complex network.

Description Colour pattern (Fig. 22A). Head entirely yellowish; pronotum yellow, with the lateral and posterior margins black; disc with two black admedian longitudinal bands, and a small spot on

each lateral area; scutellum yellow with black borders; elytra yellowish, with black narrow external margins and suture; additionally each elytron with a narrow black line separating the disc from the broadly explanate margins, the line does not reach the anterior margins or the elytra; large and dense black punctures on the explanate margins and along with suture; epipleuron yellow with black margins. Prosternum mostly black with a yellow area laterally in front of each coxa; pterothorax and abdomen black except for the lighter mesepimeron. Legs mostly black, tibiae often pale-coloured. Morphology. Length 9.0–11.5 mm. Prosternal process with carinate borders but carinae not extending beyond procoxae. Male terminalia (Figs 22D–G). Penis with apical sclerite short, simple and very slim; penis guide as long as parameres, with hook at the apex. Female terminalia (Fig. 22H). Coxite twice as long as wide, with a short subapical stylus; sperm duct thick walled, externally smooth, 0.3 times as long as bursa copulatrix; spermatheca as in Fig. 22H. Measurements (n = 5): TL 9.00–11.50; PL 1.51–1.70; EL 8.00–8.13; PW 4.20–4.40; EW 9.20–9.50; GD 3.80–4.01.

VII. Review of the Australo-Pacific Coccinellini

Distribution of Australoneda bielawskii

Ratios: PL/PW 0.33–0.38; EL/EW 0.84–0.90; EL/PL 4.70– 5.00; EL/GD 1.91–2.00; TL/EW 1.03–1.10.

Types

21

black borders. Front and middle legs pale-coloured in males, predominantly black in females; hind legs of both sexes black. Morphology. Length 8.5–11.0 mm. Elytra in lateral view strongly convex and pointed. Prosternal process with carinate borders but carinae not extending in front of the coxae. Male terminalia (Figs 23E–G). Penis with apical sclerite slim and very long; penis guide slightly shorter than parameres without apical hook. Female terminalia (Figs 23D,H). Coxite twice as long as broad, with a short apical stylus; sperm duct about as long as the bursa, membranous, slim and smooth; surface of bursa copulatrix smooth, with a few transverse wrinkles; spermatheca as in Fig. 23D. Measurements (n = 10): TL 8.50–10.85; PL 1.80–1.92; EL 7.00–8.53; PW 4.20–4.82; EW 8.12–9.25; GD 4.00–5.10. Ratios: PL/PW 0.38–0.43; EL/EW 0.80–0.92; EL/PL 3.82– 4.12; EL/GD 1.70–1.78; TL/EW 1.06–1.16.

A. bielawskii: Irian Jaya, Testega, Meydoudga, 1100–1350m, 10.vi.1993, A. Riedel (NHMS, holotype).

Types

Specimens examined

Specimens examined

INDONESIA: Manokwari, Gn. Meja, 200m, 18.iii.1993, A. Riedel (2, NHMS); Meydoudga, 1200–1400m, 5.iv.1993, A. Riedel (NHMS); Iba, 1300m, 7–8.iv.1993, A. Riedel (BMNH); Manokwari Prov., Mokwam, Arfak Mts, 1300–1650m, 17.iv.1993, A. Riedel (ANIC); Manokwari Prov., Mokwam (Siyoubirig), 1400–1800m, 24–28.II.2007, A. Weigel (NKME); Manokwari Prov., 18km NE Ransiki, Cutting area, 02.-06. III.2007, A. Skale (1, NKME); Testega, 1100–1300m, 30.iii-2. iv.1993, A. Riedel (NHMS).

AUSTRALIA: Queensland: Bunya MTs, 60km NE Dally, 900 m, 17.vi.–19.viii. 1982, S. & J. Peck (ANIC); Bunya MT., Nat. Park, 5.v.1978, B. S. Selman (ANIC); Bunya Mtns., (26.50S, 151.33E), 3 km from summit, on Kingaroy road, 6.i.1970, Britton, Holloway, Misko (1, ANIC); Joalah N.P., Tamborine Mt. (27.56°S, 156.12°E), 18–21.x.1978, Lawrence & Weir (1, ANIC); Mt. Glorious St. Forest, i–ii.1986, Y. Basset (ANIC); St Bernard’s, Mt Tamborine, 27.i.1961, C. W. Frazier (ANIC); Maryborough, 29.i.1978, B. J. Selman (1, ANIC); Brisbane, xi–xii.1978, K. Houston (ANIC); Lamington National Park, Macpherson Range, H. J. Carter (2, ANIC); Bunya Mt. N.P., 5.v.1978, B.S. Selman (1, BMNH); Tambourine Mountains, 1–9.v.1935, R. E. Turner (1, BMNH). New South Wales: Comboyne, J. Armstrong (3, ANIC); Upper Tweed. R., xii.1978, G.M. (1, ANIC); Beaury, SF, 13.xii.1984, G. Williams, ex rainforest foliage (1, ANIC); Dorrigo National Park, 11.xi.1961, C. W. Frazier (2, ANIC); N. P. Lookout. 5.200 ft, 14.x.1961, C. W. Frazier (1, ANIC); Eve Ck. 34.5 km W of Coramba, (30.14S, 153.01E), 26.vi.1976, W. Allen (1, ANIC); Gibraltar Range National Park (29.28S, 152.21E), 950 m, 10.x.1974, I. F. B. Common & E. D. Edwards (2, ANIC); Cabridge Plateau, Richmond Ranges SF, WNW Casino, 15.xi.1996, mv light, SG.

Distribution Known only from the Indonesian provinces of New Guinea.

Australoneda bourgeoisi (Gadeau de Kerville, 1884) (Fig. 23)

Neda bourgeoisi Gadeau de Kerville, 1884: 70. Type locality: Australia. Korschefsky 1932: 279; Bielawski 1963a: 469; 1965: 229. Australoneda bourgeoisi: Iablokoff-Khnzorian 1984b: 109; Pope 1989: 643 (lectotype); Ślipiński 2007: 151; Li et al. 2009: 225.

N. bourgeoisi: Australie coll. Sicard (MNHN, lectotype).

Diagnosis This is the only species of this genus occurring along the eastern coast of Australia. The non-maculate elytra with black external margins are similar to A. bielawskii but A. bourgeoisi can easily be distinguished from that species by a different pronotal colour pattern, lacking a black spot on the lateral pronotal areas, and by broad black margins of the elytra.

Description Colour pattern (Figs 23A–C). Head yellow in males and much darker in females; pronotum yellow, with a black central pattern and lateral and posterior borders; scutellum black; elytral explanate external margins and narrow sutural area black; disc orange, separated from black margins by a narrow yellow band. Prosternum mostly black, with yellow area laterally in front of the procoxae; hypomeron yellow, except for black lateral and hind margins; mesoventrite, metaventrite and abdomen black; mesanepisternum and mesepimeron usually yellowish with

Distribution of Australoneda burgeoisi

22

Ladybird Beetles of the Australo-Pacific Region

Watkins (1, ANIC); Wiangaree S.F. 8.xii.1982. G. T. Williams (1, BMNH); Dorrigo N. P. 18.xi.1973, A. & M. W. WalfordHuggins (2, BMNH); Richmond River, 1909 (1, BMNH).

Distribution This species is isolated from the centre of diversity of this genus in New Guinea, being distributed along the eastern coast of southern Queensland and northern New South Wales.

Australoneda fuerschi (Bielawski, 1963a)

Distribution of Australoneda fuerschi

(Fig. 24)

Neda fuerschi Bielawski, 1963a: 467. Type locality: New Guinea, Mount Otto. Australoneda fuerschi: Iablokoff-Khnzorian 1984b: 111; Li et al. 2009: 226.

The uniformly light brownish body with fine yellowish elytral markings but without any contrasting spots or dark markings will distinguish this species from other Australoneda species.

2000–2150m, 29. viii.1991, A. Riedel (1, ANIC). Sentani Cyclops Mts. 1000–1200m, 9.viii.1992, A. Riedel (ANIC). PAPUA NEW GUINEA: Wau Kuper Range, Biaru, 1700– 2000m, 10.x.1992, A. Riedel (1, NHMS); Wau, Mt. Kaindi, 1650m, 7.x.1992, A. Riedel (3, NHMS; 1, BMNH); Western Highlands, Mt. Hagen Range, Murmur Pass, 8700ft., 27.x-20. xii.1961, W. W. Brandt (1, ANIC); Upper Manki, 4.viii.1982, F. R. Wylie & P. Shanahan (4, ANIC); Baiyer R., 1100 m, 26.xii.78–25.i.79, J. Sedlacek (1, BMNH).

Description

Distribution

Colour pattern (Fig. 24A). Head yellow or light brown; pronotum usually brown with lighter lateral explanate margins; scutellum dark brown or black; elytra with a darker suture, brown with a yellowish explanate region and with a light band extending from the highest point of the elytra to the humeral angle. Venter from yellow to brown, mesepimeron light. Legs yellow. Morphology. Length 7.5–10.0 mm. Elytral disc in lateral view strongly convex and pointed. Prosternal process with carinate borders but carinae not extending beyond the procoxae. Male terminalia (Figs 24F–I). Penis slim with apical sclerite very short and tube-like; tegmen with penis guide as long as parameres, apex with apical hook and subapical crenulations. Female terminalia (Figs 24D,E). Coxite 2.3 times as long as wide, with subapical stylus; sperm duct slim and long, almost as long as the bursa copulatrix, very densely wrinkled; spermatheca relatively straight, C-shaped (Fig. 24D). Measurements (n = 4): TL 7.51–9.55; PL 1.10–1.41; EL 6.00–6.80; PW 3.51–4.12; EW 6.91–7.52; GD 3.52–3.61. Ratios: PL/PW 0.31–0.34; EL/EW 0.86–0.90; EL/PL 4.80– 5.41; EL/GD 1.71–1.88; TL/EW 1.08–1.12.

Known from the islands of New Guinea and New Britain.

Diagnosis

Types Neda fuerschi: New Guinea: Mt. Otto, 2200 m, June 24 1955, coll. J.L. Gressitt/ Neda fuerschi sp. n., det. R. Bielawski 1962 (BPBM, holotype); Neth, Katsime, West of Swart Val., 1500m. N.E. Mt. Otto, 2200 m, 23.vi.1955/ J.L. Gressitt/ Paratype/ Neda fuerschi sp. n., det. R. Bielawski 1962 (ANIC; BPBM, BMNH, 7 paratypes).

Specimens examined INDONESIA: Prov. Jayawijaya, Naica, 1900–2100m, 8.ix.1992, A. Riedel (1, NHMS); Prov. Jayawijaya, Bommela, 1750m, 30.viii-1.ix.1992, A. Riedel (1, NHMS; 1, ANIC); Jayawijaya, Bime, 1650–2000m, 10.ix.1993, A. Riedel (1, NHMS); Manokwari. Gn. Meja, 200m, 18.iii.1993, A. Riedel (2, NHMS); Minyambou, 1500–1900m, 13–14.iv.1993, A. Riedel (2, NHMS); Prov. Manokwari. Anggi. Gn. Dishehey,

Australoneda horni (Korschefsky, 1934b) (Fig. 25)

Neda horni Korschefsky, 1934b: 270. Type locality: Papua New Guinea, Southern Highlands, Tigibi. Bielawski 1963a: 460. Australoneda horni: Iablokoff-Khnzorian 1984b: 110; Li et al. 2009: 226.

Diagnosis This is a very distinctive species due to its elytra bearing broadly explanate, black borders and the disc with a black cross, and yellow and orange markings (Fig. 25A).

Description Colour pattern (Figs 25A–C). Head yellow, with a black central marking that is distinctly larger in females than in males; pronotum black with a broad yellow longitudinal area, separated from the lateral margin and extending from the anterior margin to ~2/3 of the pronotal length; scutellum black; elytra tricoloured, with broad, black explanate margins; disc with broad longitudinal and transverse markings forming roughly a cross, with the anterior quarters yellowish and orange or brown, and posterior quarters yellow; epipleuron yellow and black, the yellow area close to the pterothorax and surrounded by black borders. Hypomeron black, with a yellow marking corresponding to a yellow dorsal area; prosternum mostly black with a yellow area laterally in front of the procoxae; pterothorax and abdomen black, except for light mesepimeron. Morphology. Length 7.5–10.0 mm. Prosternal process with external borders carinate but not extending in front of the procoxae; elytral disc in profile strongly convex and pointed (Fig. 25C). Male terminalia (Figs 25F–H). Apical sclerite of penis slim and very long; penis guide much shorter than parameres. Female terminalia (Figs 25D,E). Coxite twice as long as broad with a short stylus; sperm duct slim about twice as long

VII. Review of the Australo-Pacific Coccinellini

Distribution of Australoneda horni

as bursa, with surfaces densely wrinkled; surfaces of bursa copulatrix wrinkled and uneven (Fig. 25E); spermatheca as in Fig. 25D. Measurements (n = 2): TL 7.51–9.95; PL 1.80–1.90; EL 5.20–6.51; PW 3.51–4.12; EW 6.91–7.52; GD 3.52–3.61. Ratios: PL/PW 0.31–0.34; EL/EW 0.86–0.90; EL/PL 4.80– 5.41; EL/GD 1.71–1.88; TL/EW 1.08–1.12.

23

separated from the lateral spot or suture. The dorsal markings may vary considerably from an absent apical spot (Fig. 26A), to a form with apical spots enlarged and joined with lateral spots. Hypomeron yellow with a broad black lateral margin; prosternum mostly black with yellow area laterally in front of the procoxae; pterothorax and abdomen black except for a lighter mesepimeron. Legs yellow. Morphology. Length 7.1–10.0 mm. Elytra strongly convex and pointed in lateral view. Prosternal process with prosternal carinae extending anteriorly beyond the procoxae. Male terminalia (Figs 26E–H). Penis thickened and crenulate near the middle, apical piece curved, short and pointed in the apex; penis guide as long as parameres, with hook at apex. Female terminalia. With a wrinkled bursa copulatrix; spermatheca as in Fig. 26D. Measurements (n = 5): TL 7.21–9.88; PL 1.30–1.85; EL 5.70–6.90; PW 3.21–4.22; EW 6.91–8.52; GD 3.22–4.51. Ratios: PL/PW 0.40–0.45; EL/EW 0.78–0.83; EL/PL 3.72– 4.38; EL/GD 1.58–1.78; TL/EW 1.04–1.06.

Types Types N. horni: ‘D. Neu-Guinea, Wahnes, Franklin Müller/ Neda Hornii m. det. R. Korschefsky 1934/ Paratypus’ (1, NHMB; 1, MNHN; 1, BMNH, syntypes).

Specimens examined PAPUA NEW GUINEA: Bosavi, xi.1973, H. Ohlmus (1, ANIC); Bulolo, 26.xii.1971, H. Ohlmus (1, ANIC); D.N. Guinea, Stattelberg (6, ZMB).

Distribution Endemic to the southern part of Papua New Guinea.

Australoneda karubakana (Bielawski, 1963a) (Fig. 26)

Neda karubakana Bielawski, 1963a: 462. Type locality: New Guinea Australoneda karubakana: Iablokoff-Khnzorian 1984b: 112; Li et al. 2009: 228.

Diagnosis Australoneda karubakana is very similar to A. taengana and can only be distinguished by the presence of a black spot at the centre of the yellowish lateral area on the pronotum, by the more numerous dorsal elytral spots, and by details of male terminalia (see also diagnosis of A. taengana).

Neda karubakana: NETH., Kutsime, West of Swart Val., 1500 m, Nov. 14 1958, coll. J.L. Gressitt (BPBM, holotype).

Specimens examined INDONESIA: NE. Swart Val. Karubaka, 1450m, 5.xi.1958, J. L. Gressitt (1, ANIC); N.E. Swart Valley, Karubaka, 1500 m, 11.xi.1958, J.L. Gressitt (1, BMNH); Jayawijaya. Bime, 1600–1900 m, 22.ix.1993, A. Riedel (2, NHMS); Biak-Is., Suneri, 4.x.1990, A. Riedel (3, NHMS); Prov. Manokwari. Membey, 800–1200m, 31.viii.1991, A. Riedel (1, ANIC); Testega, 1200m, 31.iii-12.iv.1993, A. Riedel (16, NHMS, ANIC); Iba, 1300 m, 7–8.iv.1993, A. Riedel (1, NHMS); Anggi, Tetaho Kosmena, 1400–1750m, 26–28.iii.1993, A. Riedel (1, NHMS); Nabire. N. Mapia. Km177. Ugida, 1400m, 29.vii.1996, Schule, Stuben (1, NHMS). PAPUA NEW GUINEA: Prov. Morobe, Aseki, 1000–1300m, 13.x.1992, A. Riedel (10, NHMS); Wau, Mt. Kaindi, 1550–2150 m; 7–8.x.1992, A. Riedel (ANIC); Wau, Mt. Kaindi, 1650 m; 7.x.1992, A. Riedel (2,NHMS); Upper Manki, logging area, Bulolo, Dist, 16.vi.1972, 30.vi.1972, 13.vii.1972, F. R. Wylie & P. Shanahan (5, ANIC); NE Wau, 1200m, 21–25.xii.1965, J. Sedlacek (BPBM); Bulolo, 26.xii.1971, H. Ohlmus (1, ANIC); Miyama, xii.1971, H. Ohlmus (2, ANIC).

Distribution Endemic to the island of New Guinea.

Description Colour pattern (Figs 26A–C). Head yellow or brown; pronotum medially black, with two large yellow lateral areas, almost always with a black central dot, joined by a narrow stripe along the anterior margin and extending posteriorly to almost the posterior margin, and clearly separated from the lateral margins; scutellum black. In the typical form (Fig. 26B), the elytra are black with one reddish to yellowish crescentshaped area along the explanate margin, a triangular basal spot extending from the scutellum to almost the humeral angle and narrowing posteriorly, and an elongate apical spot clearly

Distribution of Australoneda karubakana

24

Ladybird Beetles of the Australo-Pacific Region

Australoneda maai (Bielawski, 1963a) (Fig. 27)

Neda maai Bielawski, 1963a: 458. Type locality: Papua New Guinea, Kassam. Australoneda maai: Iablokoff-Khnzorian 1984b: 110 (as synonym of A. horni); Li et al. 2009: 227.

Remarks This species is known from two females only and its status remains unclear. It was synonymised with A. horni by IablokoffKhnzorian (1984b) but was resurrected by Li et al. (2009) based on characters of the spermatheca, but these distinctions should be confirmed by the collection of males for further examination of the male terminalia.

Diagnosis This species is somewhat similar to A. horni, sharing black a explanate region of the elytra and yellowish sublateral pronotal markings without dark central spots. Both species can be separated by the elytral colour pattern, consisting of four spots in A. horni and two much smaller and narrower spots in A. maai.

Description Colour pattern (Fig. 27B). Head yellowish brown, with black frontal area; pronotum black, with a broad sublateral yellow area extending from the anterior margin to ~2/3 of the pronotal length; scutellum black; elytra black, each with two light yellow spots near the explanate margins; epipleuron black with one yellow area. Prosternum completely black; hypomeron yellow with a broad black lateral margin; pterothorax and abdomen black, with a lighter mesepimeron. Legs black. Morphology. Length 8.4 mm. Elytra strongly convex and pointed in profile. Prosternal process with carinate borders but carinae do not extend beyond the procoxae. Abdominal postcoxal line does not reach the lateral margin, with one discontinuity in the middle. Male. Unknown. Female terminalia (Figs 27C–E). Coxite 2.5 times as long as wide, with subapical stylus; sperm duct slim and densely wrinkled, ~1.5 times as long as the bursa copulatrix; spermatheca (Fig. 27D) with base enlarged and spiral elements relatively coarser than in other species. Measurements. Length 9.4 mm; width 8.9 mm.

Australoneda ruitong Li, Ślipiński & Pang, 2009 (Figs 1A,B, 28)

Australoneda ruitong Li, Ślipiński & Pang, 2009: 226. Type locality: Papua New Guinea, Eastern Highlands, Tigibi.

Diagnosis This is the only species of the genus to have a lateral elytral profile that is rounded and not pointed at the highest point; it may also easily be distinguished from remaining Australoneda by each elytron bearing two discal and a single marginal, deep red spots.

Description

A species currently known from two separate localities at the opposite ends of the island of New Guinea.

Colour pattern (Figs 28A,B). Head brown in males, brown with single black marking in females; pronotum black, with a sublateral longitudinal elliptical wite or yellow area extending from the anterior margin to ~2/3 of the pronotal length; one small black insertion in the middle of the external border of each yellow area; scutellum black; elytra black with one reddish crescent-shaped area on the explanate margins and two red spots on the disc along suture; epipleuron red and black, with bright yellow spots close to the anterior angle of the epipleuron in males. Prothoracic hypomeron yellow and black; prosternum mostly black with a yellow area laterally in front of the procoxae; pterothorax and abdomen black except for the lighter mesepimeron. Front and middle legs pale-coloured in male, black in female; hind legs black in both sexes. Morphology. Length 8.0–8.5 mm. Prosternal process with carinate borders, but carinae not extending beyond the procoxae. Male terminalia (Figs 28D–G). Apical sclerite of the penis short and curled, pointed at apex; penis guide as long as parameres with hook at the apex. Female terminalia (Figs 28C,H). Coxite twice as long as wide, arcuate apically with a short, subapical stylus; sperm duct thick and twisted with margins almost smooth, ~0.6 times as long as the bursa; bursa copulatrix densely wrinkled (Fig. 28H); spermatheca as in Fig. 28C. Measurements (n = 2): TL 8.0–8.5; PL 1.40–1.71; EL 6.01– 6.52; PW 3.51–3.73; EW 6.00–6.49; GD 2.60–3.00. Ratios: PL/PW 0.40–0.46; EL/EW 0.80–0.82; EL/PL 3.82–4.28; EL/ GD 2.17–2.30; TL/EW 1.05–1.06.

Distribution of Australoneda maai

Distribution of Australoneda ruitong

Types Neda maai: Papua New Guinea, N.E. Kassam, 1350 m, 48 km E. of Kainantu, 7.xi.1959, coll. T.C. Maa/ Holotype/ Neda maai sp. n. det. R. Bielawski 1962/ Holotypus (BPBM, holotype).

Specimens examined INDONESIA: Testega, Meydoudga, 1100–1350 m, 10.iv.1993, A. Riedel. (1, NHMS).

Distribution

VII. Review of the Australo-Pacific Coccinellini

25

Types A. ruitong: ‘Terr Papua & New Guinea: Tari subdistrict, Tigibi, native gardens, 28.v-12.vi.1966, 1600m, W Vink’ (NBCL, holotype and paratype).

Distribution So far, this species was collected from a single locality in the mountainous region of Papua New Guinea but the photographs of live specimens taken by Paul Zborowski (Figs 1A,B) near Komo confirm its slightly broader range.

Australoneda taengana (Bielawski, 1963a) (Fig. 29)

Neda taengana Bielawski, 1963a: 465. Type locality: Papua New Guinea, Taenga. Australoneda taengana: Iablokoff-Khnzorian 1984b: 113.

Diagnosis Australoneda taengana can rather easily be recognised by its pronotal lateral area without a black spot, and entirely black elytra with a red area along the explanate margin only. It can only be confused with the rather variable A. karubakana, however, the latter can be distinguished by the pronotum bearing a black spot in the centre of the lateral area and much more extensive elytral dorsal maculation. However, we have seen three specimens of A. karubakana with extensive elytral maculation but without a black spot on the pronotal lateral area.

Description Colour pattern (Fig. 29B). Head yellow in females with black marking in males; pronotum centrally black, with two large yellow lateral areas joined by a narrow stripe along the anterior margin and extending posteriorly to almost the posterior margin, clearly separated from lateral margins; scutellum black; elytra black with one reddish crescent area along the explanate margin (a single male examined had a very small yellow marking close to each side of scutellum). Hypomeron yellow, with a broad black lateral margin; prosternum mostly black with a yellow area laterally in front of theprocoxae; pterothorax and abdomen black except for the lighter mesepimeron. Legs yellow. Morphology. Length 7.5–9.6 mm. Elytral disc strongly convex and pointed in profile. Prosternal process with carinae anteriorly beyond the procoxae. Male terminalia (Figs 29C–F). Penis with apical sclerite very short, narrowing apically. Penis guide as long as the parameres with hook at the apex. Female terminalia. Coxite 2.3 times as long as wide, with a subapical stylus; sperm duct very short, conical and ringed; bursa copulatrix large and wrinkled, distinctly narrowing apically; spermatheca as in Fig. 29G. Measurements (n = 3): TL 7.51–9.55; PL 1.10–1.30; EL 5.71–5.90; PW 3.21–3.30; EW 7.7–9.01; GD 3.01–3.11. Ratios: PL/PW 0.39–0.41; EL/EW 0.75–0.81; EL/PL 4.30–4.40; EL/ GD 1.80–1.91; TL/EW 1.00–1.08.

Types Neda taengana: New Guinea (N.E.), Taenga, 1200 m, Upper Jimmi V., July 14 1955, coll. J.L. Gressitt (BPBM, holotype female).

Distribution of Australoneda taengana

Specimens examined PAPUA NEW GUINEA: Akivitana River, 1550m, 10.i.1965 (1, BPBM); SE Northern Dist., 1500 m, 26.x.1974, in sticky trap Upper Manki. L.A. Bulolo, M. District, 4.viii.1972, F.R. Wylie & P. Shanahan (1, ANIC); Baiyer River, 1100 m, 26.xii.78– 25.i.79, J. Sedlacek (1, BMNH); E. Highland District, Okasa near Okapa, 19.i.1965, M.E. Bacchus (1, BMNH); Okasa Pine Forest, 12.v.1967, B. Gray (1, BMNH).

Distribution Known from southern part of Papua New Guinea.

Cheilomenes Chevrolat, 1836 (Figs 30–32)

Cheilomenes Chevrolat in Dejean, 1836: 435. Type species, designated by Crotch 1874: 179, Coccinella lunata Fabricius, 1775. Mulsant 1850: 443. Chilomenes Crotch 1874: 179. Unjustified emendation. Menochilus Timberlake, 1943: 40. Replacement name for Cheilomenes sensu Mulsant 1850.

References: Sasaji 1971: 284; Iablokoff-Khnzorian 1979: 58; 1982: 144; 1986: 61; Pope 1989: 646. Diagnostic combination

Length 5–7 mm; dorsum weakly convex, glabrous. Elytral colour pattern highly variable (Fig. 32). Anterior clypeal border arcuate between lateral projections (Fig. 13C). Antenna 11segmented; distinctly shorter than head capsule width, with scape expanded (Fig. 13U) and compact three-segmented club (Fig. 13W). Terminal maxillary palpomere securiform. Prothoracic hypomeron without a fovea near the anterior angles. Prosternal process with admedian carinae extending forward to the middle of the prosternum. Anterior margin of the mesoventrite deeply emarginate medially. Elytral margin without a marginal bead; epipleuron without or with a shallow fovea. Tibial spur formula 0–2–2. Tarsal claws with a large subquadrate basal tooth. Abdominal postcoxal line not recurved, incomplete laterally; oblique postcoxal line very short. Penis consists of a single sclerite with a very long and thin apical part (Fig. 30G). Infundibulum absent; spermatheca small with long and strongly convoluted sperm duct (Fig. 30E).

Description Body elongate-oval or hemispherical; dorsum glabrous. Head. Mostly exposed with eyes clearly visible from above; their inner orbits more or less parallel. Frons broad, more than twice the width of the eye. Antenna 0.7 times as broad as the head capsule, 11-segmented; scape strongly expanded externally; antennal club compact, three-segmented, the penultimate

26

Ladybird Beetles of the Australo-Pacific Region

antennomere distinctly longer than the terminal segment, the terminal antennomere subquadrate, apically rounded. Clypeus short, at least weakly arcuate between two lateral projections. Mandible bifid apically, molar part with a basal tooth. Terminal maxillary palpomere distinctly securiform. Prothorax. Pronotum convex; anterior margin emarginate medially; anterior angles rounded or blunt. Prothoracic hypomeron without delimited foveae. Prosternal process ~0.3–0.4 times the coxal diameter, with the carinae extending anteriorly to the middle of the prosternum. Pterothorax. Scutellum ~1/12 times as broad as the elytra at shoulders. Elytra at base much broader than the pronotum, lateral margins distinctly and entirely explanate without a marginal bead; elytral epipleuron without or with shallow foveae. Mesoventrite deeply emarginate anteriorly, with a complete raised ridge; mesal surface with a deep fossa receiving the prosternal process. Metaventral postcoxal lines joined medially, complete. Tarsal claws in both sexes with a large subquadrate basal tooth; mid and hind tibiae with two apical spurs. Abdomen with six ventrites in both sexes, ventrite 1 as long as ventrite 2. Abdominal postcoxal lines separate medially, not recurved; oblique postcoxal line present (Fig. 32G). Male terminalia. Penis consists of a single, very long and apically curly sclerite. Female terminalia. Spermatheca with the cornu weakly curved; sperm duct long and very complicated; infundibulum absent. Coxites with short apical styli.

Biology

Khnzorian 1982). Ślipiński (2007) described the larva of C. sexmaculata from Australia.

Distribution Cheilomenes is mostly distributed in the Afrotropical region, with few species extending to the Middle-East and further to India, China, Japan and the Pacific Islands.

Remarks Cheilomenes is distinguished by the arcuate anterior margin of the clypeus, short antenna with scape carinate externally, and the elytral margins that are devoid of a thickened external border or a gutter along their edge. There has been disagreement about the generic limits in this genus. Mader (1954) recognised a generic group Cydoniinae, based on the relatively short antenna that was as long as the width of the frons, with a ‘spindle-forming’ club. He recognised three valid genera: Elpis Mulsant (without hypomeral but with epipleural concavities), Cheilomenes Chevrolat (without hypomeral or epipleural concavities), and Cydonia Mulsant (with hypomeral cavities but without epipleural conavities). Iablokoff-Khnzorian (1979, 1982) recognised Cheilomenes and Menochilus as valid but later synonymised Menochilus Timberlake with Elpis Mulsant and included it as a subgenus under expanded concept of Cheilomenes Chevrolat (see Iablokoff-Khnzorian 1986). This was criticised by Fürsch (1989), who argued that there were no substantial differences between these taxa and all should be united under the oldest name Cheilomenes. This convention is followed here.

All species of Cheilomenes are apparently aphidophagous, but only C. sexmaculata has been studied in detail (Iablokoff-

Key to Cheilomenes species from the Australo-Pacific Region 1. – 2(1). –

Body broadly oval, elytral colour pattern as in Fig. 30.................................................................................................... C. polynesiae Body elongate-oval, elytral colour pattern different (Figs 31, 32)...................................................................................................... 2 Length less than 4 mm; elytra with four round spots (Fig. 31)........................................................................................ C. samoensis Length more than 5 mm; elytra with different colour pattern (Fig. 32)........................................................................C. sexmaculata

Cheilomenes polynesiae Crotch, 1874 (Fig. 30)

Chilomenes polynesiae Crotch, 1874: 181. Type location: Fiji. Gordon 1987: 23 (lectotype). Cheilomenes polynesiae: Iablokoff-Khnzorian 1986: 68.

Diagnosis Cheilomenes polynesiae is easily distinguished from C. sexmaculata by its broader and larger body and very distinctive colour pattern.

Description Colour pattern (Figs 30A–D). Frons and head appendages yellow in males, in female frons yellow with extensive black marking extending to the labrum. Pronotum black with large yellow antero-lateral areas. Scutellum black. Elytra yellow, with black margins and sutural area connected by transverse fascia in the anterior third; additional longitudinal lateral fascia usually limited to the anterior half, but sometimes extending

posteriorly and complete, forming a regular pattern of four yellow spots dorsally (Fig. 30C). Prosternum black, hypomeron yellow. Pterothorax black except for the paler mesepimeron. Elytral epipleuron mostly yellow; legs black. Abdomen dark brown. Morphology. Length 5.7–6.0 mm. Frons flat, distinctly punctured, punctures 1.0–1.3 diameters apart, intervals with reticulate microsculpture. Antenna with scape strongly expanded asymmetrically into a lobe on the upper side. Pronotum with punctures on disc and margins similar in size to those of the head, but more widely separated, intervals finely reticulate and weakly shiny. Scutellum triangular, pointed apically. Elytral punctures smaller than those on the pronotum, usually separated by 1–2 diameters, intervals smooth or finely reticulate. Elytral epipleura broad, descending externally with shallow foveae. Abdomen with apical margin of ventrite 5 straight in males, arcuate in females; ventrite 6 arcuate in both sexes. Male terminalia (Figs 30G–J). Female terminalia (Figs 30E,F). Measurements (n = 3): TL 5.70–6.10; PL 0.8–1.10; EL 4.5–4.8; PW 3.02–3.21; EW 5.30–5.70; GD 2.31–2.6. Ratios:

VII. Review of the Australo-Pacific Coccinellini

27

Distribution of Cheilomenes polynesiae

PL/PW 0.3–0.4; EL/EW 0.80–0.91; EL/PL 4.10–4.40; EL/GD 2.11–2.40; TL/EW 1.05–1.10.

Types C. polynesiae: Type/ Type polynesiae Fiji (UCCC, lectotype).

Specimens examined AUSTRALIA: Norfolk Island: 300 ft, 16.iii.1939, I. McComish (1, BMNH). Philip Island: 29.07S 167.57E, Red Terraces, 20–24.xi.1984, T. A. Weir (2, ANIC); 29.07S 167.57E, Red Terraces, 20–24.xi.1984, I. D. Naumann (1, ANIC); 29.07S 167.57E, Upper Dykes, 26.iii-2.iv.1984, D. C. F. Rentz (1, ANIC); 29.07S 167.57E, Lower Long Valley, Malaise trap/ethanol, 20–24.xi.1984, I. D. Naumann (3, ANIC); 29.07S 167.57E, Lower Long Valley, 20–24.xi.1984, T. A. Weir (1, ANIC); 29.07S 167.57E, Upper long Valley, 20–24.xi.1984, L. Hill (4, ANIC); Philip Island, 21.xi.1984 (1, BMNH).

Distribution Fiji, and Norfolk and Philip Islands of Australia.

Description Colour pattern (Figs 31A–C). Frons and head appendages yellow. Pronotum yellow with large, almost subquadrite black area medially. Scutellum black. Elytra black, each elytron with one reddish medial spot and one yellow apical spot. Prosternum black, hypomeron yellow. Pterothorax black, except for a pale mesepimeron. Elytral epipleuron black, except for anteriorinternal yellow part. Anterior and middle legs brown with small dark areas, hind legs largely black. Abdomen dark. Morphology. Length 3.7–4.1 mm. Terminal maxillary palpomere strongly securiform. Elytra as long as broad or slightly shorter, deeply punctuated. Elytral epipleuron 0.2–0.3 times as broad as the metathorax. Male terminalia. As in Figs 31D–G. Female terminalia. As in Figs 31H,I. Measurements (n = 3): TL 3.70–4.10 (3.93 ± 0.21); PL 0.8– 1.00 (0.90 ± 0.10); EL 2.90–3.10 (3.03–0.12); PW 2.00–2.10 (2.07 ± 0.06); EW 3.00–3.30 (3.13 ± 0.115); GD 1.60–1.70 (1.63 ± 0.06). Ratios: PL/PW 0.40–0.48 (0.43 ± 0.03); EL/EW 0.94–1.00 (0.97 ± 0.03); EL/PL 3.10–3.63 (3.39 ± 0.27); EL/ GD 1.81–1.94 (1.88 ± 0.07); TL/EW 1.23–1.29 (1.26 ± 0.03).

Types C. samoensis: Samoa, Upolu (BMNH, holotype).

Cheilomenes samoensis Arrow, 1927 (Fig. 31)

Cheilomenes samoensis Arrow, 1927: 57. Type locality: Samoa, Upolu Island. Korschefsky 1932: 304; Iablokoff-Khnzorian 1986: 66. Cheilomenes samoensis var. tutuilensis Arrow, 1927: 58. Type locality: Tutuila Island. Korschefsky 1932: 304.

Diagnosis This highly distinctive species can be easily diagnosed by its small size, elytra with two reddish spots located in the middle of the elytra, and two yellow spots apically (Fig. 31A).

Specimens examined SAMOA: Upolu I., Mulivai, i.1978, N. L. H. Krausss (2, ANIC); South Upolu, 14.00s 171.45E, 11.iv.2003, R. Bashford (1, ANIC); Alafua, Upolu Is., 31.iii.1919 (1, NZAC); Apia Upolu, 23.vii, 1960 (1, NZAC); Alafua, Upolu, 20.vi.1977, Maddison (1, NZAC); Upolu, 5.iii.1975, T. Bengtsson (1, NZAC).

Distribution Known only from the Pacific islands of Samoa and American Samoa.

28

Ladybird Beetles of the Australo-Pacific Region

Distribution of Cheilomenes samoensis

Cheilomenes sexmaculata (Fabricius, 1781) (Fig. 32)

Coccinella sexmaculata Fabricius, 1781: 96. Type locality: India. Coccinella quadriplagiata Schoenherr, 1808: 195. Synonymised by Nakane and Araki 1959: 50. Cheilomenes duodecimpunctata Fauvel, 1867: 209. Type locality: New Caledonia, Lifu Island. Iablokoff-Khnzorian 1986: 68; Nattier et al. 2015: 322. Syn. nov. Cheilomenes sexmaculata: Dejean, 1836: 435; Mulsant 1850:444; Crotch 1871: 8; 1874: 180. Cheilomenes quadriplagiata: Dejean, 1836: 435; Mulsant 1850: 447; Crotch 1871: 8; 1874: 181. Cheilomenes sexmaculata var. flavofasciata Mulsant, 1850: 446. Orcus mollipes Oliff, 1895: 30. Type locality: Lord Howe Island. Synonymised by Pope 1989: 646. Cheilomenes sexmaculata var. australis Weise 1908: 12. Menochilus sexmaculatus: Timberlake 1943: 40; Sasaji 1971: 285; Pope 1989: 646; Nattier et al. 2015: 308.

Diagnosis This common species occurs in several colour forms (Fig. 32) but each is distinct from C. polynesiae, known only from Fiji, and Norfolk Islands, which is broader, usually larger and has a characteristic colour pattern (Fig. 30).

Description Colour pattern (Figs 32A–D). Pattern variable from almost entirely black to orange with transverse fasciate markings on the pronotum and the elytra. Head yellow in males, yellow with a dark central spot in females; ventral side of thorax and abdomen dark, abdominal margins, prothoracic hypomera and inner margins of the elytral epipleura pale, legs dark. Morphology. Length 4.0–6.0 mm. Frons flat, distinctly punctured, punctures 1.0–1.5 diameters apart, intervals with very fine, reticulate microsculpture. Antenna with scape expanded asymmetrically into a lobe on the upper side. Pronotum with punctures on disc and margins similar in size to those of the head, but more widely separated, intervals finely microreticulate and shiny. Scutellum triangular, pointed apically. Elytral punctures slightly larger than those of the pronotum, usually separated 1–2

diameters, intervals smooth. Elytral epipleura broad, moderately descending externally, without foveae. Abdomen with the apical margin of ventrite 5 straight or very slightly emarginate in males, arcuate in females. Male terminalia (Figs 32H–K). Female terminalia (Figs 32E,F). Measurements (n = 20): TL 4.00–6.00 (5.09 ± 0.59); PL 0.80–1.10 (0.97 ± 0.08); EL 3.00–4.90 (4.12 ± 0.53); PW 2.00– 3.00 (2.55 ± 0.26); EW 3.20–4.80 (4.17 ± 0.44); GD 1.50–2.65 (2.04 ± 0.27). Ratios: PL/PW 0.33–0.45 (0.38 ± 0.03); EL/EW 0.86–1.09 (0.99 ± 0.05); EL/PL 3.00–4.80 (4.24 ± 0.42); EL/ GD 1.67–2.38 (2.03 ± 0.17); TL/EW 1.12–1.34 (1.22 ± 0.08).

Specimens examined INDONESIA: Jayapura Sentani, Cyclops Mts., 300–500m, 8.viii.1992 & 31.x.1992, A. Riedel (1, NHMS); Manowari Ransiki, Mayuby, 26–30.ix.1990, 300m, A. Riedel (3,NHMS); Paniai Mulia (s.) Wuyuneeri, 1900–2200, 6–7.vii.1994, A. Riedel (1,NHMS); Boana, 1000m, 14.iv-1.v.1965, H. Pyka (1. NHMS); Anggi, Tetaho Kosmena, 1400–1750m, 26–28. iii.1993, A. Riedel (1. NHMS); 5km west Fakfak, 8.vii.1976, Schule & Stuben (1, NHMS); Nabire nach Mapia, Unipo, 24.vii.1996, Schule & Stuben (2, NHMS). Ambon: Waai, 2.iii.1966 & 9.iv.1964 a.m. R. Wegner (BPBM; CAS). PAPUA NEW GUINEA: Sandaun Pr., Telefomin, trail to Eliptamin, 1700–1900m, 16–17.v.1998, A. Riedel (3, NHMS); Timbari, 10.xi.1965, H. Pyka (2, NHMS); Southern Highl. Prov., Tari-koroba, Hake, 1700–2000m, 14.v.1998, A. Riedel (1, NHMS); Wau, 1500m, viii.1983, M. Mastaller (1, NHMS); Wau, Mt, Kaindi, 4.v.1971, R.E. Parrot (1, CNC); Wau, 4000ft, 28.vi-3.vii.1974 (1.CNC); (NE) Wau, Morobe distr., Mt. Missim, 1120m, 13.ii.1963, J. Sedlacek(1, BPBM); Wau, ii.1974 (1, CAS); Morobe Prov., Garaina, 700m, 21.iii.1998, A. Riedel (2, NHMS); Morobe District, Goroka Highlands,8. xi.1969, J. E. Tobler (4, CAS); Morobe District, Busu river, 16km E. of Lae, 3.xi.1969, J. E. Tobler (2, CAS); Munga, 25km SE Chuaye, 8.ii.1973, K. W. Stroeder (2, CNC); Mt. Hagen, 1700m, 1973, G. Baker (1, CNC); Bulolo River valley near Kaaisinik, 12.i.1978, R. Harmsen & J. Harmsen (1,CNC); Upper Manki L. A., Bulolu, M Dist., 4.viii.1972, F. R. Wylie & P. Shanahan (2, ANIC); Popondetta, Popondetta Mission, 60m,

VII. Review of the Australo-Pacific Coccinellini

29

Distribution of Cheilomenes sexmaculata

18.x.1963, P. Shanahan (1, BPBM); Pasi nr. Vanimo, 9.vi.1993, J. M. Grimshaw (1, NAQS); Mt. Lamington Dis, Northern Division, vii.1927, C. T. MacNamara (9, AM). AUSTRALIA: Northern Territory: Darwin, 1.iii.1974, N. Forester (1, ANIC). Western Australia: Perth, vii.2003, R. Horbury (16, ANIC); Bunbury (1, ANIC). Christmas Island: South Point Settlement, 30.ix.1964, T. Campbell (ANIC).

Distribution This species is widely distributed, known from China, India and Japan to New Caledonia and other Pacific Islands, and northern and western Australia. It is impossible to establish the original geographic range of this species, but based on its food preferences for aphids and the historical record, its occurrence in Australia is probably from unintentional introduction (Pope 1989).

Remarks Following the argument of Nattier et al. (2015) there is little doubt that the species described by Fauvel (1867) as Cheilomenes duodecimpunctata is identical to C. sexmaculata, commonly occurring on Lifou Island, New Caledonia. The true identity of a female specimen from ‘Ost-Celebes, Tamboegoa’ illustrated by Iablokoff-Khnzorian (1986: 68) as C. duodecimpunctata remains unknown.

Cleobora Mulsant, 1850 (Fig. 33)

Cleodora Mulsant, 1850: 160. Type species by monotypy, Cleodora mellyi Mulsant, 1850 (junior homonym). Cleobora Mulsant, 1850: 1025 (replacement name).

References: Iablokoff-Khnzorian 1982: 301; Pope 1989: 655; Ślipiński 2007: 153. Diagnostic combination Length 5–8 mm; dorsum weakly convex, glabrous. Elytral colour pattern constant, uniformly yellowish or orange with

black transverse markings (Fig. 33A). Anterior clypeal border straight. Antenna 11-segmented; slightly longer than the head capsule width, weakly club-shaped. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron without a fovea near the anterior angles. Prosternal process without admedian carinae. Anterior margin of mesoventrite deeply emarginate medially. Elytral margin with a complete bead; epipleuron not foveate. Tibial spur formula 0–2–2. Tarsal claws with a large subquadrate basal tooth. Abdominal postcoxal line not recurved, incomplete laterally; oblique postcoxal line absent. Penis consists of a single sclerite. Infundibulum absent; spermatheca long and convoluted.

Description Body elongate-oval, moderately convex, distinctly narrowing apically; dorsum glabrous. Head (Fig. 33C). Eye partially exposed, with inner orbits converging posteriorly; frons about twice as wide as the eye width. Antenna as long as or longer than the head capsule, 11-segmented; scape not expanded; antennal club loose, threesegmented; the penultimate antennomere about as long as the terminal antennomere which is distinctly elongate, apically rounded. Anterior clypeal margin straight. Mandible bifid apically, molar part with a basal tooth; prostheca distinct. Prothorax. Pronotum weakly convex; pronotal base without a marginal bead; lateral margins broadly explanate without a bead. Prothoracic hypomeron without foveae. Prosternal process extending beyond the coxae, smooth, without carinae. Pterothorax. Scutellum triangular, pointed apically. Elytra at the base much broader than the pronotum, the lateral margins very narrow but entirely visible from above, surface smooth or micro-punctate. Elytral epipleuron complete to the apex of elytra, strongly descending externally, epipleural foveae absent. Mesoventrite emarginate anteriorly, with a complete raised border; mesal surface with a deep fossa for receiving the prosternal process. Meso-metaventral process 0.5–0.6 times the mesocoxal diameter. Metaventral postcoxal lines joined medially, complete and weakly descending. Mid and hind tibiae with two apical spurs.

30

Ladybird Beetles of the Australo-Pacific Region

Abdomen with six ventrites in both sexes. Abdominal postcoxal lines separate medially, not recurved and incomplete laterally, posteriorly distinctly separated from the hind margin of the ventrite; oblique postcoxal line absent. Male terminalia. Penis guide symmetrical; penis stout, consists of a single sclerite; basal capsule T-shaped. Female terminalia. Coxites with terminal styli; infundibulum tube-like, enclosing the sperm duct; sperm duct simple, uniform in diameter. Spermatheca long and with several bends; basal ramus and nodulus moderately developed.

Biology Ślipiński (2007) described the larva of C. mellyi. The exact food preferences of C. mellyi are unknown, but various sources (cited by Pope 1989) indicate that psyllids are probably the primary prey of this species, the diet being supplemented by aphids and larval chrysomelids when available. Bashford (1999) recorded C. mellyi as an effective predator of eggs of the chrysomelid leaf beetle Chrysophtharta bimaculata (Olivier), a major defoliator of young eucalypts in Tasmania.

Distribution This monotypic genus is endemic and widespread in southern and central Australia; introduced to New Zealand.

Remarks Cleobora mellyi is a distinctive ladybird, readily distinguished by its yellowish elytra with dark, transverse markings, strongly developed tibial spurs, prothoracic hypomeron without foveae, and its smooth, non-carinate prosternal process.

Cleobora mellyi (Mulsant, 1850) (Fig. 33)

Cleodora mellyi Mulsant, 1850: 160. Type locality: Tasmania. Cleobora mellyi: Mulsant 1850: 1025 (by implication); Mulsant 1866: 123; Pope 1989: 656 (lectotype); Ślipiński 2007: 153–155. Halyzia mellyi: Masters 1888: 77.656.

Distribution of Cleobora mellyi

Description Colour pattern (Figs 33A–C). Head yellow or orange, with black transverse markings on clypeus; labrum yellow; pronotum and elytra orange or yellow, with dense black colour pattern as in Figs 33A–C, with transverse elytral bands varying in thickness. Prothoracic hypomera yellow, except for the posterior third of lateral margins, hind margins and postcoxal extensions; prosternum, mesoventrite and metaventrite black; mesepimeron pale yellow; abdominal segments mostly black with a sublateral pale area on each side of the first four ventrites. A very pale, unusually coloured specimen has been photographed in Western Australia (Fig. 3A). Morphology. Length 5.5–9.0 mm. Frons flat, frontal punctures small, 1–2 diameters apart, intervals with reticulate microsculpture. Pronotal punctures smaller than those of the head, usually separated by about 1 diameter; intervals with reticulate microsculpture. Scutellum ~1/10 of the elytral width at the humeri, punctate. Elytra with external margins having a complete but very fine bead; punctures of the disc of two sizes; intervals between the punctures smooth and shiny. Posterior margin of ventrite 5 broadly truncate in males, broadly arcuate in females. Male terminalia. As in Figs 33F–I. Female terminalia. As in Figs 33D,E,J. Measurements (n = 5): TL 5.50–9.00; PL 1.00–1.52; EL 5.81–7.40; PW 3.22–4.11; EW 5.51–6.82; GD 2.51–3.20. Ratios: PL/PW 0.31–0.37; EL/EW 1.05–1.09; EL/PL 4.92– 5.80; EL/GD 1.94–2.32; TL/EW 1.21–1.30.

Types C. mellyi: Museum Paris Australie Verreaux 1 - 46/ 1.46/ 58/ Cleobora mellyi Muls. auct. det. (MNHN, lectotype).

Specimens examined AUSTRALIA: New South Wales: Barrington Tops; Mt Irvine; Jenolan Caves; Sydney; Alpine Way; Braidwood; Clyde Mountains; Old Adaminaby; Tuross; Mt Kosciuszko. Australian Capital Territory: Canberra, Uriarra Crossing;

VII. Review of the Australo-Pacific Coccinellini

Blundells Creek. Victoria: Dartmouth Dam; Harrietville; Bogong High Plains; Mt Buller; Humevale; Healesville; Melton; Warburton; Melbourne; Singleton Reserve, Mt Dandenong; Ferntree Gully; Monbulk; Emerald; Lysterfield; Warragul; Geelong; Otway Ranges; Moonlight Head; Wilsons Promontory. Tasmania: King Island; Table Cape; Burnie; Elliot; Winnaleah; Devonport; Underwood; Launceston; Great Lake; Lake St Clair; Queenstown; Strahan; Hobart; Florentine Valley; Mt Field National Park; Mt Wellington. South Australia: Adelaide; Mt Lofty; Kangaroo Island; Mt Gambier. Western Australia: Perth; Bunbury; Busselton; Katanning; Margaret River; Hopetoun; Karridale; Boranna; Deepdene; Augusta; Mt Barker; Porongorup; Nornalup; King George Sound.

Distribution The southern ladybird is distributed in the southern and eastern parts of Australia, extending and west to Perth. In 1977, it was introduced to New Zealand from Tasmania, to control Paropsis charybdis Stål, the eucalypt tortoise beetle. Subsequent introductions to several localities in New Zealand were successful and it has apparently become established in Northland and the Bay of Plenty (Martin 2016b).

Coccinella Linnaeus, 1758 (Figs 34–36)

Coccinella Linnaeus, 1758: 364. Type species designated by Latreille 1810: 432, Coccinella septempunctata Linnaeus, 1758.

References: Iablokoff-Khnzorian 1982: 341; Gordon 1985: 783; Poorani 2002a: 325. Diagnostic combination Length 4–8 mm; dorsum convex, glabrous. Elytral colour variable, often red or orange with black markings. Anterior clypeal border straight. Antenna 11-segmented; shorter than the head capsule width, with a distinct three-segmented club. Terminal maxillary palpomere securiform. Prothoracic hypomeron without a fovea near the anterior angles. Prosternal process with admedian carinae. Anterior margin of the mesoventrite straight. Elytral margin with complete bead; epipleuron without foveae. Tibial spur formula 0–2–2. Tarsal claws with a large subquadrate basal tooth. Abdominal postcoxal line not recurved, incomplete laterally; an oblique postcoxal line present. Penis consists of a single sclerite. Infundibulum heavily sclerotised; spermatheca with a strongly developed ramus and nodulus.

Description Body broadly oval; moderately to strongly convex; dorsum glabrous. Head (Fig. 34B). Eyes largely exposed with the inner orbit almost parallel. Frons usually black with two pale spots, at least twice as broad as the eye width. Anterior clypeal border truncate. Antenna short and compact, ~0.5–0.6 times the width

31

of the head capsule, 11-segmented; scape not expanded, pedicel much shorter than the scape; club three-segmented, compact, terminal antennomere broad, subquadrate. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere strongly securiform. Prothorax. Pronotum moderately convex, slightly narrower than the elytra at the base. Prothoracic hypomeron without foveae. Prosternal process narrow, with two convergent or parallel carinae. Pterothorax. Scutellum quite small, triangular. Elytra yellow or red with maculation or not; lateral margin narrowly explanate; epipleuron narrow, nearly flat, without foveae. Mesoventral process ~0.3 times as broad as the diameter of the mid-coxae. Mesoventrite anteriorly straight and with a complete raised ridge; mesal surface without clear articulation with the prosternal process. Metaventrite with discrimen almost reaching the the anterior margin; postcoxal lines joined medially, arcuate, complete. Middle and hind tibia each with two apical spurs; tarsal claws with each with a large, basal subquadrate tooth. Abdomen with six ventrites in both sexes; intercoxal process rounded apically, ~0.2 times as broad as the abdomen; posterior margin of ventrite 5 truncate in both sexes. Abdominal postcoxal line descending, approaching the posterior margin of ventrite 1; an oblique postcoxal line present (Fig. 36J). Male terminalia. Tegmen stout with a large phallobase, penis guide broad at the base, tapering anteriorly; parameres slim, usually shorter than the penis guide. Penis short, consisting of a single sclerite; outer arm of the penis capsule long; inner arm strongly curved, sometimes indistinct. Female terminalia. Coxites broad, with distinct styli. Sperm duct short, uniform in diameter; infundibulum heavily sclerotised, usually flared at one or at both ends. Spermatheca curved to C-shaped, with a distinct nodulus and ramus.

Biology All species of Coccinella are primarily aphid feeders, and some species have a very broad range of recorded host plants (Gordon 1985). They are also found on flowers and can consume pollen and nectar.

Distribution Coccinella is a moderately large genus, with ~50 species distributed mostly in the Holarctic region, extending to Africa and the Pacific. Only C. leonina from New Zealand is endemic to the Australo-Pacific region.

Remarks Coccinella is very similar to Adalia, sharing the features of the straight, non-emarginate anterior margin of the mesoventrite, and the female genital tract with the tube-like and sclerotised infundibulum. Coccinella differs from Adalia in having the postcoxal line of the first abdominal ventrite not recurved and an associated oblique postcoxal line.

Key to species of the genus Coccinella from Australo-Pacific region 1. – 2(1). –

Elytra with two, usually zig-zag transverse bands (Fig. 35A)..................................................................................... C. transversalis Elytra with isolated spots or patches (Fig. 34A).................................................................................................................................. 2 Elytra yellow, with 11 black spots (Fig. 36A)......................................................................................................C. undecimpunctata Elytra black, with 16 yellow patches (Fig. 34A)..................................................................................................................C. leonina

32

Ladybird Beetles of the Australo-Pacific Region

Coccinella leonina Fabricius, 1775 (Fig. 34)

Coccinella leonina Fabricius, 1775: 87. Type locality: New Zealand. Goeze 1777: 243; Gmelin 1790: 1665; Mulsant 1850: 128, 1866: 100; Crotch 1874: 118; Korschefsky 1932: 508; Watt 1986. Coccinella tasmanii White, 1846: 23. Type locality: New Zealand. Synonymised by Korschefsky 1932: 509. Coccinella coriacea Broun, 1893: 1319. Type locality: New Zealand: Mount Cook, near the Hermitage. Synonymised by Broun 1893: 1319.

Diagnosis This species is endemic to New Zealand, where it overlaps with the introduced C. undecimpunctata. Coccinella leonina can be easily distinguished from all species of Coccinella by its general black colouration, with yellowish or orange spots on the elytra.

Description Colour pattern (Figs 34A–C). Frons black, with one pale spot on either side close to the eyes. Pronotum black with subquadrate yellowish-cream patch on either side. Elytra black, combined with 16 orange patches arranged as in Fig. 34A, sometimes joined transversely. Scutellum and ventral side black, the prothoracic hypomeron yellow anteriorly; elytral epipleuron yellow, except for an external black border. Legs black. Morphology. Length 4.5–5.3 mm. Frons between the eyes ~2.5 times as broad as the eye width, with punctures about as large as the eye facets. Pronotal punctures similar in size to those of the head, interspaces shiny; lateral borders weakly arcuate. Prosternum with distinct carinae, slightly convergent anteriorly. Scutellum small. Elytra slightly longer than broad, broadest at about the middle. Punctures on the elytra more deeply impressed and appearing larger than those on the pronotum, mostly separated by 1 diameter or less. Abdomen with posterior margin of ventrite 5 truncate in males, and slightly arcuate in females; ventrite 6 with a broadly transverse apical fovea in males and simple in females. Male terminalia (Figs 34D,E,I,J). Female terminalia (Figs 34F–H). Measurements (n = 5): TL 4.40–5.30 (4.74 ± 0.40); PL 0.80–1.00 (0.92 ± 0.08); EL 3.50–4.00 (3.76 ± 0.23); PW 2.20– 2.40 (2.32 ± 0.08); EW 3.20–3.90 (3.50 ± 0.27); GD 1.70–2.00 (1.88 ± 0.13). Ratios: PL/PW 0.35–0.43 (0.40 ± 0.03); EL/EW 1.03–1.11 (1.08 ± 0.03); EL/PL 3.89–4.50 (4.10 ± 0.24); EL/ GD 1.85–2.22 (2.01 ± 0.15); TL/EW 1.28–1.47 (1.36 ± 0.07).

Specimens examined NEW ZEALAND: Banks Pen., 12.8 km NW Akaroa, Barry’s Bay, 16.i.1960, C.W. O’Brien (1, BPBM); Banks Pen., Kaituna 20–60m, 9.xii.1962, J. Mather (1, BPBM); Cant. Prov., 6 km N Tai Tapu, 9.i.1960, C.W. O’Brien (1, BPBM); S. Alps, Cass, 549m, 15.x.1961, K.A.J. Wise (1, BPBM); North I: Makino (1, BPBM); Westland: Jackson Bay, 21.i.1960, R. E. Leech (1, BPBM); S. Isl. KA Okarahia Stream, 6 km SW Oaru, 25.i.1997, Schuh & Lang (1, NHMW); Weitakerei Range, H. Suter S. G. (1, ZMB); Matakataki R. Murchison area, S. Isl., 17.xii.1988, W. W. Middlekauff (2, ANIC); Mavora R., 5km above N. Mavora Lake S. Isl., 7.xii.1989, W. W. Middlekauff (1, ANIC); Nelson, 5.xi.1927, J. W. Evans (2, ANIC); Ohakune

Distribution of Coccinella leonina

King Country, N. IS.,31.xii.1916, A. E. Brooks (1, ANIC); South Island, Nelson, 26.iv.1969, J. E. Tobler (15, CAS); Mt. John 3900 Tekapo canterburg, 11.i.1970, J. C. Watt (1, NZAC); Alderman Is Ruamahuanui I., 17.xi.1972, G. W. Ramsay (1, NZAC); CO Lower Manor Burn Dam, 8.iii.1979, J.C. Watt (NZAC); Tunakino V. nr Roi, 8.x.1971, R.J.B. Power (1, NZAC); ND Poor Knights Is., Tawhiti Rahi Shag Bay, 6–12. ix.1980 (1, NZAC);

Distribution Endemic to New Zealand.

Remarks Watt (1982) recorded C. leonina from the Poor Knights Islands (New Zealand) and made an interesting observation on the occurrence of this species: ‘This common native ladybird does not occur on the mainland in Northland, Auckland, or Coromandel although it occurs on several northern islands. It appears to be a general feeder on aphids, and occurs frequently in pastures and adventive crops, as well as native habitats, in the southern North Island and in the South Island. As it is a common, conspicuous beetle where it occurs, it is unlikely that it has been overlooked in northern areas of the mainland. Its restriction to islands in the north cannot be explained by either climate alone or habitat modification. It seems most likely that it extended to the far north in pre-European times, although it was better adapted to the climate farther south. The European ladybird Coccinella undecimpunctata Linnaeus, 1758 was introduced to New Zealand in the first half of the nineteenth century. This species, which is also a general predator on aphids, is climatically versatile. Apparently, Coccinella leonina was unable to compete successfully with C. undecimpunctata in the north, although it has held its own further south. This hypothesis is supported by the absence of the European species on northern islands where C. leonina occurs, and vice versa.’

VII. Review of the Australo-Pacific Coccinellini

Coccinella transversalis Fabricius, 1781 (Fig. 35)

Coccinella transversalis Fabricius, 1781: 97. Type locality: Coromandel. Mulsant 1850: 1022; Timberlake 1943: 14; IablokoffKhnzorian 1979: 68, 1982: 391; Pope 1989: 652 (lectotype); Ślipiński 2007: 158; Nattier et al. 2015: 304. Coccinella repanda Thunberg, 1781: 18. Type locality: Cape of Good Hope. Mulsant 1850: 124; Crotch 1871: 3, 1874: 117; Froggatt 1902: 899; Weise 1902: 489; 1908: 11, 1923:132; Korschefsky 1932: 483; Bielawski 1962: 204. Synonymised by Mulsant 1850: 126. Coccinella contempta Boisduval, 1835: 592. Type locality: Australia. Synonymised by Mulsant 1850: 1022.

Diagnosis This is a very distinctive species, with the elytra bearing black, transverse zig-zag bands on a yellowish or orange background.

Description Colour pattern (Figs 35A–C). Head black with triangular yellow area on either side near inner eye borders. Head appendages usually brown to black. Pronotum black with two large rectangular yellow areas anteriorly (Fig. 35B). Scutellum black. Background colour of elytra yellow, with transverse black markings, lateral borders yellow and suture black. Ventral side mostly black with the mesepimeron, metepimeron and posterior area of the metanepisternum pale, anterior part of the prothoracic hypomeron and the elytral epipleuron yellow. Legs and abdomen black. Morphology. Length 4.0–7.0 mm. Frons 3.0–3.5 times wider than the eye, weakly convex, finely punctured with interspaces densely reticulate and weakly shiny. Pronotum with the anterior border straight medially. Prosternal process very narrow, not extending beyond the procoxae. Elytra slightly broader than the pronotum at the base, broadest at the middle; external border with a narrow margin. Scutellum at the base ~1/12–1/13 times as broad as the elytra at the shoulders. Abdominal ventrite 5 truncate in both sexes; posterior margin of ventrite 6 almost truncate in males and strongly arcuate in females.

Distribution of Coccinella transversalis

33

Male terminalia (Figs 35D–I). Penis short, ~0.6 times as long as the abdomen; tegmen (strut not included) 0.4 times as long as the abdominal disc; strut slight shorter than the tegmen; phallobase comparably broad and almost as long as the parameres. In ventral view, the penis guide is broad at the base and tapers evenly to the apex, slightly extending the parameres. In lateral view, the penis guide is almost straight. Female terminalia. Spermatheca C-shaped, with a distinct nodulus and ramus. Sperm duct slim and short; infundibulum flared at the end leading to the spermatheca (Fig. 35J). Measurements (n = 12): TL 4.20–7.00 (5.27 ± 0.79); PL 0.70–1.20 (1.00 ± 0.13); EL 3.30–5.20 (4.16 ± 0.61); PW 2.10– 2.90 (2.53 ± 0.29); EW 3.10–4.60 (3.95 ± 0.50); GD 1.70–2.60 (2.19 ± 0.31). Ratios: PL/PW 0.33–0.45 (0.39 ± 0.04); EL/EW 0.91–1.13 (1.05 ± 0.06); EL/PL 3.50–5.14 (4.20 ± 0.49); EL/ GD 1.56–2.08 (1.91 ± 0.15); TL/EW 1.16–1.52 (1.33 ± 0.11).

Specimens examined INDONESIA: Nabire, nr. Mapia, 1400m, 29.vii.1996, Schule & Stuben (NHMS); Prov. Jayaweiyaya, Djuremna, 1900–2100m, 9–11.ix.1992, A. Riedel (NHMS); Maffin Bay, 14.vii.1944, E. S. Ross (CAS); Mapia, 1480m, 30.vii.1996, Schule & Stuben (NHMS). PAPUA NEW GUINEA: Mando, 27.vii.1972 (CNC); Nadzab x.1944; J. R. Helfer (CAS); Surprise Creek, Watut River, 3400ft, 1955, G. D. Woodard (CAS); Mt. Wilhelm, 14.vii.1972 (CNC); Oro Bay, xii.1943, J. R. Helfer (CAS). NEW CALEDONIA: Aoupinie 950–1000m, 8.ii.2004, M. Wanat (ANIC); Gavatch near Tiendante stream valley, 4.ii.2004, M. Wanat (ANIC); Tiea Forest, sclerophyllous forest, 30.i.2004, M. Wanat (ANIC); 280–330 m, Haute Riviere Bleue: La Tranchee-Sentier des Kaoris. Humid forest, 28.i.2004, M. Wanat (ANIC); 4km S. Polindimie, 7.iii.1978, E.I. Schlinger (CAS); Touho, Mts., above 500 ft., 8.iii.1978, E.I. Schlinger (CAS); SOLOMON ISLANDS: Bougainville, bush E. of Buin, 9.i.1970, J. E. Tobler (CAS); Guadalcanal, Honiara, 0–100m, ii. 1985, N. L. H. Krauss (CAS). AUSTRALIA: all states and territories, very common. Norfolk Island: Point Hunter Reserve, 29.xi.1984, I.D.

34

Ladybird Beetles of the Australo-Pacific Region

Naumann (ANIC); Rocky Pt. Reserve, 14.xi-2.xii.1984, L. Hill (ANIC). Philip Island: Long Valley, 20–24.xi.1984, I.D. Naumann (ANIC); Upper Long Valley, 20–24.xi.1984, I.D. Naumann (ANIC); Lower Long Valley, 20–24.xi.1984, T. A. Weir (ANIC); Rocky Valley, 20–24.xi.1984, L. Hill (ANIC); Moo-oo beach, 20–24.xi.1984, L. Hill (ANIC); Red Terraces, 20–24.xi.1984, T.A. Weir (ANIC).

Distribution This is a widespread species, with a distribution extending from India and China to Polynesia and Micronesia. It is common in New Guinea, Australia, New Caledonia, Solomon Islands and several smaller Pacific Islands.

Coccinella undecimpunctata Linnaeus, 1758 (Fig. 36)

Coccinella undecimpunctata Linnaeus, 1758: 366. Type locality: Europe. Iablokoff-Khnzorian 1979: 66, 1982: 351, Pope 1989: 651; Ślipiński 2007: 158. Coccinella novaezealandiae Colenso, 1888: 40. Type locality: Napier, New Zealand. Synonymised by Sharp 1891: 352.

Diagnosis This species is a distinctive member of Coccinella. The relatively narrowly oval body, and yellowish-orange colour of the elytra, with 11 small isolated black spots are very diagnostic.

Description Colour pattern (Figs 36A–C). Frons black, with one pale spot on either side close to the eyes. Head appendages black or dark brown. Pronotum black, with a subtriangular yellowish-cream patch on either side of the lateral-anterior border. Scutellum black. Elytra orange-yellow, with small pale patches on either side of the scutellum and 11 black markings arranged as per Fig. 36A; external borders and suture yellow. Ventral side black, except for the mesepimeron, the anterior area of hypomeron and the elytral epipleuron Morphology. Length 3.5–5.8 mm. Frons between the eyes ~2.5 times broader than the eye width; surface with punctures

Distribution of Coccinella undecimpunctata

slightly larger than the eye facets. Pronotum weakly convex with punctures similar in size to those of the head; lateral border weakly arcuate. Prosternum with distinct carinae, convergent anteriorly. Scutellum small. Elytra slightly longer than broad, broadest at about the middle, slightly narrowing to the apices. Punctures on the disc more deeply impressed than those on the pronotum and appearing larger, mostly separated by 1 diameter or less. Posterior border of ventrite 5 truncate in males, and slightly arcuate in females. Ventrite 6 with a broadly transverse apical fovea in males and simple in females. Male terminalia (Figs 36F–I). Female terminalia (Figs 36D, E). Measurements (n = 10): TL 3.50–5.80 (4.81 ± 0.85); PL 0.60–0.90 (0.81 ± 0.11); EL 3.00–4.90 (4.10 ± 0.65); PW 1.80– 2.60 (2.23 ± 0.25); EW 2.60–4.30 (3.55 ± 0.60); GD 1.00–2.00 (1.55 ± 0.30). Ratios: PL/PW 0.32–0.40 (0.36 ± 0.03); EL/EW 1.12–1.22 (1.16 ± 0.04); EL/PL 4.25–6.00 (5.07 ± 0.53); EL/ GD 2.13–3.00 (2.67 ± 0.26); TL/EW 1.03–1.47 (1.36 ± 0.13).

Specimens examined AUSTRALIA: New South Wales: Tallaganda SF NSW, c. 8km SSE Captains Flat, Bursaria flowers, 5.iii.1989, C. Reid (1, ANIC). Victoria: Devon city, Geelong, 28.v.1976 (4, ANIC). South Australia: 34.26855S, 140.29748E, DAC6962, feeding on aphids, Ex vineyard cover crop, 24.x.2002, G. Furness & N. Schellhorn (5, ANIC). Tasmania: Gladstone, 31.iii.1991, V. Lorimer (1, ANIC); 41.48 S,146.52 E, 9 km W of Poatina, 20.i.1983, I.D. Naumann & J.C. Cardale (1, ANIC); Liffey Valley, 17.xi.1982, S. Fearn (2, ANIC); Wimot, 12.ix.1984, S. VonSee (2, ANIC); Beechford, 1.i.1985, S. Fearn (1, ANIC); Poatina, 1.i.1988, S. Vonsee (1, ANIC). Norfolk Island: 29.03S 167.55E, Rocky Pt. Reserve, 14.xi-2.xii.1984, L. Hill (1, ANIC); 29.04S 167.58E, Point Hunter Res., 29.xi.1984, T.A. Weir (3, ANIC); 1 km NE Kingston, 26.xii.1979, J.C. Cardale (1, ANIC). NEW ZEALAND: Hinemaiaia R. Lake Taupo, N. Isl., 28.i.1984, W. W. Middlekauff (1, CAS); Taupo, N. Isl., 27– 30.i.1984, W. W. Middlekauff (2, CAS); N. Isl., Ruakaka Bch., 6.vii.1983, J. Doyen (1, CAS); South Island, Te Anau, 24–26. ii.1978, W. W. Middlekauff (1, CAS); N-Isl., CL Hotwater Beach, 14km N Tairua, 21.ii.1997, Schuh & Lang (2, NHMV);

VII. Review of the Australo-Pacific Coccinellini

South Island, headwaters, Mataura R. Cattle flat, 10.ii.1978, W. W. Middlekauff (2, CAS); South Island, 12 mi N. Gore, 3.ii.1978, W. W. Middlekauff (1, CAS); Otago, 8.iv.1959, K. P. Lamb (1, NZAC); AK Noises Is., Otata I., 27.viii.1979, C. F. Butchar (1, NZAC); Wairoa Sorge Nelson, 24.xi.1969, J. S. Dugdale (1, NZAC); Mt. Ngaurohoe, 2291m, 26.i.1980, A. K. Walker (1, NZAC); Matakataki R. Murchison area, S. Isl., 17.xii.1988, W. W. Middlekauff (10, ANIC).

Distribution This species is found in Europe, the Mediterranean region, Iran, Iraq, and north-west India. It was introduced to the Falkland Islands, New Zealand and Australia.

Coelophora Mulsant, 1850 (Figs 37–39)

Coelophora Mulsant, 1850: 390. Type species by subsequent designation of Crotch 1874: 148, Coccinella inaequalis Fabricius, 1775. Spilocaria Timberlake, 1943: 58. Type species by original designation Coelophora bisellata Mulsant, 1850. Synonymised by Iablokoff-Khnzorian 1982: 218. Lemnia (Microlemnia) Iablokoff-Khnzorian, 1982: 218, 261. Invalid taxon, not accompanied by description.

References: Chazeau 1978; Iablokoff-Khnzorian 1982: 265; Pope 1989: 662; Ślipiński 2007: 158. Diagnostic combination Length 4.0–6.6 mm, elytra moderately convex, glabrous. Elytral colour pattern extremely variable. Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly shorter than the head capsule, with a moderately compact three-segmented club. Terminal maxillary palpomere securiform. Pronotal disc evenly convex transversely; the lateral margins slightly concave within upturned and slightly thickened external borders. Prothoracic hypomeron with a fovea near the anterior angles; the prosternal process with distinct carinae extending anteriorly. Anterior margin of the mesoventrite deeply emarginate medially. Elytra with narrow raised margins; epipleuron distinctly foveate at the level of the hind coxae. Tibial spurs formula 0–2–2. Abdominal postcoxal line not recurved and almost complete laterally (Fig. 39J); an oblique postcoxal line absent. Penis consists of a basal sclerite and a hinged, much smaller, apical piece (Fig. 38K). Ovipositor, coxites with apical styli; infundibulum absent; spermatheca large and irregular, cornu long C-shaped and sometimes wrinkled; nodulus and ramus poorly developed and close to each other (Fig. 38N).

Description Body moderately convex, oval; dorsum glabrous. Head strongly withdrawn into prothorax. Eyes largely beneath the pronotum, with the inner orbits strongly divergent anteriorly. Frons narrow, 1.5–1.7 times the width of the eye. Antenna 11-segmented, 0.7 times the head width, scape slightly

35

enlarged, the last three antennomeres form a rather compact club; the terminal antennomere rounded apically, longer than the penultimate. Anterior clypeal margin straight. Terminal maxillary palpomere strongly securiform. Mandible bifid apically, molar part with a basal tooth. Prothorax. Pronotum moderately convex; anterior margin strongly emarginate medially, forming a blunt anterior angle. Prothoracic hypomeron with foveae. Prosternal process with admedian carinae extending forward to the middle of the prosternum. Pterothorax. Scutellum ~1/10–1/12 times as broad as the elytra at the shoulders. Elytra convex, slightly broader than the pronotum at the base, lateral margins not explanate. Elytral epipleuron slightly inclined, with distinct foveae at the middle. Mesoventrite strongly emarginate medially with a complete raised ridge; mesal surface with clear articulation with the prosternal process. Postcoxal line of the metaventrite joined medially, complete laterally; oblique postcoxal line absent (Fig. 39J). Apex of the middle and hind tibiae each with two spurs; tarsal claws with subquadrate basal teeth. Abdomen with six ventrites in both sexes. Postcoxal lines joined or separate medially, complete laterally, not recurved; oblique postcoxal line absent. Male terminalia. Penis consists of two sclerites (Fig. 38J); the basal piece with a moderate expansion near the middle; apical sclerite short, associated with two flat sclerotised pieces on the ventral side (Fig. 38K). Female terminalia. Coxites broad-based with short, terminal styli; infundibulum absent. Spermatheca large and irregular, cornu long, C-shaped and sometimes wrinkled; nodulus and ramus poorly developed and close to each other (Fig. 38N).

Biology The detailed biology of Coelophora is unknown, but C. inaequalis commonly feeds on introduced aphids in Australia, New Zealand and Hawaii (Pope 1989). Larvae have been described by Houston (1988) and Ślipiński (2007).

Distribution This widely distributed Asian and Australian genus extends from China to Australia. It was also introduced to New Zealand, Hawaii and Florida (Gordon 1985; Pope 1989).

Remarks Pope (1989) summarised the convoluted history of Coelophora and related taxa. In contrast to his definition of the genus, we are returning to a restricted concept of Coelophora, as advocated by Iablokoff-Khnzorian (1982) supported by an unpublished molecular phylogeny of Coccinellini (Tomaszewska et al., pers. comm.). In that phylogeny, the type species, Coleophora inaequalis, is distinct from the other Australian species classified in Coelophora by Pope (1989), which form a distinct clade with Microcaria and are here moved to that genus. Unfortunately, the morphological characters that can be used to distinguish these genera are difficult to observe and mostly limited to the male and female terminalia.

Key to species of Coelophora of the Australo-Pacific Region 1. –

Elytra entirely black, or black with basal and apical orange spots (Fig. 39)....................................................................................... 2 Elytra with complex red and black colour pattern of stripes or maculae (Fig. 38).............................................................................. 3

36

Ladybird Beetles of the Australo-Pacific Region

2(1).

Pronotum entirely black or orange anteriorly, with a black posterior area separated by a straight line at the middle (Figs 39A,B); spermatheca as in Fig. 39E; male terminalia as in Figs 39F–I..................................................................................C. vesiculata – Pronotum variable, but orange and yellow anteriorly, with a black posterior area forming a pattern similar to an M in the middle (Fig. 38A); spermatheca as in Fig. 38N; male terminalia as in Figs 38J–M............................ C. inaequalis (rare colour form) 3(1). Elytra with very fine, reticulate microsculpture between punctures, the punctures small and separated by 3–5 diameters; male ­terminalia as in Figs 38J–M...................................................................................................................................... C. inaequalis – Elytra smooth between punctures, the punctures larger and usually less than two diameters apart; male terminalia different than above.............................................................................................................................................................................C. bisellata

Coelophora bisellata Mulsant, 1850 (Fig. 37)

Coelophora bisellata Mulsant, 1850: 400. Type locality: le Bengale. Booth and Pope 1989: 348 (lectotype); Poorani 2002a: 327. Spilocaria bisellata: Timberlake 1943: 58.

Diagnosis This species resembles some colour variations of Coelophora inaequalis (Fig. 38) but can easily be distinguished by differences in the male and female genitalia. The elytral surface of C. bisellata is smooth and shiny, devoid of the reticulate microsculpture that is clearly visible in most specimens of C. inaequalis.

Description Colour pattern (Figs 37A–C). Frons and head appendages yellow, the vertex behind the eyes dark brown. Pronotum yellow anteriorly, with four black spots at the base that are usually expanded and connected, forming an irregular transverse band occupying the larger part of the pronotum. Scutellum black. Elytra yellow, with black usually connected spots forming checkered pattern, external borders and suture black. Ventral side black except for a pale mesepimeron, the yellow anterior part of prothoracic hypomera, the lateral borders of prosternum and the external borders of abdomen; elytral epipleura yellow, with black external borders. Legs usually black with yellow tarsi. Morphology. Length 4.3–5.8 mm. Head deeply inserted into the prothorax, eyes completely covered from above. Frons 1.5–2.0 times as broad as the eye width. Antenna about as long as the width of the head capsule, club very loose. Pronotum moderately convex, punctate; interspaces densely reticulate, anterior border almost straight medially, lateral borders evenly arcuate with a very fine bead. Prosternal process with carinae convergent anteriorly, ending at the middle of the prosternum. Scutellum with base margin almost as long as the lateral margin. Elytra slightly broader than the pronotum, narrowly explanate laterally with a narrow bead; disc with punctures

of various sizes, larger laterally than on the disc, separated by 1–2 diameters; interspaces smooth or micropunctured, but without dense reticulate microsculpture. Elytral epipleuron 3.0–3.2 times the width of the metanepisternum, foveate near the middle. Anterior margin of the mesoventrite strongly emarginate medially, truncate posteriorly. Abdominal postcoxal line short (Fig. 37J). Posterior margin of ventrite 5 truncate in males, slightly arcuate in females. Male terminalia (Figs 37D–H). Penis consists of two sclerites; capsule T-shaped, with curved inner arm. Tegmen with a stout phallobase and thick parameres; penis guide slightly longer than the parameres, pointed apically and curved downwards; strut as long as tegmen. Female terminalia (Fig. 37I). Spermatheca strongly curved, C-shaped; sperm duct slim. Measurements (n = 18): TL 4.30–5.80 (5.03 ± 0.38); PL 0.85–1.20 (1.01 ± 0.12); EL 2.65–4.50 (3.93 ± 0.43); PW 2.25– 3.10 (2.79 ± 0.22); EW 3.30–4.80 (4.18 ± 0.34); GD 1.80–2.70 (2.24 ± 0.26). Ratios: PL/PW 0.30–0.42 (0.36 ± 0.03); EL/EW 0.70–1.03 (0.94 ± 0.07); EL/PL 3.12–4.82 (3.92 ± 0.45); EL/ GD 1.33–2.11 (1.76 ± 0.16); TL/EW 1.13–1.30 (1.20 ± 0.05).

Types C. bisellata: Bengal /Type Col: 1911 (OMNH, lectotype).

Specimens examined INDONESIA: Anggi, Tetaho Kosmena 1400–1750m, 26–28. iii.1993, A. Riedel (1, NHMS). PAPUA NEW GUINEA: West Highland District, Mt. Hagen, 1700m, 1973, G. Baker, (1, CNC); Mando, 3M. Asaro, 18M Goroka, 27.vii.1972 (2, CNC); Wau 1200m, open grassland, 8.i.1978, J. Harmsen & R. Harmsen (1, CNC); Bulolo River valley near Kaisinik, 12.i.1978, J. Harmsen & R. Harmsen (1, CNC); Nondugl 1600m, ix-xi.1951, G. Gyldenstolpe (5, NHRS); Morobe District, Wau, 2–3.x.1969, J.E. Tobler (4, CAS); Morobe District, Goroka Highlands 8.xi.1969, J.E. Tobler (1, CAS); Morobe District, Wau. B. P. Bishop Museum Field station, 3.x.1969, J.E. Tobler (2, CAS); Kaironk Valley, Schrader Range c.100mi: WSW. of Madang, 1967, G. G. Jackson (1, ANIC); Central District, Sageri, 9.iii.1973, A. Straatman; Papua Tapini, 1000–1100m, 18.v.1961, J. L. & M. Gressitt (1, BPBM).

Distribution India, China, Malaysia, Thailand, Philippines; Indonesia, Papua New Guinea (Poorani 2002a).

Coelophora inaequalis (Fabricius, 1775) (Fig. 38)

Distribution of Coelophora bisellata

Coccinella inaequalis Fabricius, 1775: 80. Type locality: Nova Hollandia.

VII. Review of the Australo-Pacific Coccinellini

Coelophora inaequalis: Mulsant 1850: 405; 1866: 266; Crotch 1874: 153; Korschefsky 1932: 292; Timberlake 1943: 31; Hales 1976: 273; 1977: 55; Chazeau 1978: 59; Houston 1979: 45; Iablokoff-Khnzorian 1979: 61; 1983: 266; Houston and Hales 1980: 669; Pope 1989: 665 (lectotypes); Yu 2010: 067; Nattier et al. 2015: 305. Coccinella punctata Fabricius, 1775: 81. Type locality: Nova Hollandia (junior homonym). Coccinella novemmaculata Fabricius, 1781: 97. Replacement name for C. novempunctata. Synonymised by Crotch 1874:153 Coelophora novemmaculata: Mulsant 1850: 398; 1866: 264. Lemnia desolata Mulsant, 1850: 387. Type locality: Nouvelle Hollande. Mulsant 1866: 254. Synonymised by Houston 1979: 49. Coelophora desolata: Crotch 1874:150; Korschefsky 1932: 292. Coelophora vidua Mulsant, 1850: 393. Type locality: Java. Synonymised by Poorani et al. 2008: 592. Vernia artensis Montrouzier, 1861: 305. Type locality: New Caledonia, Balade and Art Island. Synonymised by Chazeau 1978: 59. Coelophora ripponi Crotch, 1874: 156. Type locality: Australia. Korschefsky 1932: 296. Synonymised by Hales 1977: 77. Coelophora mastersi Blackburn, 1892: 238. Type locality: N. Queensland. Lea 1902: 490; Korschefsky 1932: 293. Synonymised by Timberlake 1943: 31. Coelophora veranioides Blackburn, 1894: 239. Type locality: N.S.W. Lea 1902: 490; Korschefsky 1932: 297; Timberlake 1943: 33. Synonymised by Hales 1977: 55. Coccinella religiosa Lea, 1902: 488. Type locality: Behn River, N.W. Australia. Korschefsky 1932: 509. Synonymised by Hales 1977: 55.

Diagnosis Some colour forms of Coelophora inaequalis resemble C. bisellata but can easily be distinguished by differences in the male and female genitalia. Also, the elytral surface of C. bisellata is smooth and shiny, devoid of the reticulate microsculpture that is clearly visible in most specimens of C. inaequalis. Very rare melanic forms of Coelophora inaequalis from New Guinea are very similar to C. vesiculata, and they can only be distinguished by the shape of the spermatheca and the penis bearing a much shorter apical sclerite.

Distribution of Coelophora inaequalis

37

Description Colour pattern. Dorsal colour patterns are highly variable (Figs 38A–I), from orange with black markings to almost uniformly black; head and appendages usually yellow; the ventral side with mesoventrite and metaventrite and central part of the abdomen usually dark, at least part of the elytral epipleura and legs are pale. Morphology. Length 3.8–6.5 mm. Eyes largely covered by the pronotum from above. Frons 1.5–2.2 times broader than eye diameter, flat, finely punctured, punctures slightly smaller than the eye facets. Pronotum moderately convex; the anterior border arcuate medially; lateral margins strongly arcuate at the base with a very fine bead. Prosternal process with a pair of carinae extending forward to almost the middle of the prosternum. Hypomeron with a well-delimited fovea anteriorly. Scutellum strongly transverse, sparsely punctured. Elytra evenly convex, with punctures as large as the eye facets; interspaces with dense reticulate microsculpture, weakly shiny; lateral borders narrowly explanate with a complete but fine bead. Elytral epipleuron broad, ~1.5–2.0 times the width of the metanepisternum, with a shallow fovea at the middle. Anterior margin of the mesoventrite deeply emarginate, posterior margin slightly arcuate. Abdomen with the posterior margin of ventrite 5 truncate in both sexes; ventrite 6 shallowly emarginate in males and arcuate in females. Male terminalia (Figs 38J–M). Penis with a strongly recurved terminal sclerite; outer arm of the capsule in line with the main tube; inner arm short, strongly curved apically; tegmen long, with the strut slightly shorter than tegmen, phallobase short, 0.3 times as long as the parameres. Female terminalia (Figs 38N,O). Spermatheca large, strongly curved, T-shaped at base, broadened at the middle; sperm duct with thick walls; infundibulum absent. Measurements (n = 5): TL 3.80–6.50 (4.97 ± 0.68); PL 0.80–1.30 (1.00 ± 0.14); EL 3.00–4.60 (3.90 ± 0.50); PW 2.00–3.00 (2.59 ± 0.32); EW 3.20–4.80 (4.14 ± 0.46); GD 1.50–2.75 (2.15 ± 0.32). Ratios: PL/PW 0.33–0.48 (0.39 ± 0.04); EL/EW 0.83–1.00 (0.94 ± 0.04); EL/PL 3.00–4.60 (3.93 ± 0.42); EL/GD 1.55–2.10 (1.83 ± 0.15); TL/EW 1.12– 1.44 (1.20 ± 0.07).

38

Ladybird Beetles of the Australo-Pacific Region

Types L. desolata: Holotype O.U.M./ 48./ P. Ess./48 Lemnia desolata Mulst. Type Mus Hope Lemnia desolata Mulsant 1850 (OMNH, holotype); C. ripponi: Austral./ TYPE / TYPE Ripponi Cr./ Coelophora inaequalis F. Khnzorian det. (UCCC, lectotype); C. mastersi: Type/ 4426 N.Qu. T/Coelophora Mastersi, Blackb. (BMNH, lectotype); C. veranioides: Type/ 5219 N.S.W.T/ Coelophora veranioides, Blackb. (BMNH, lectotype); C. religiosa: religiosa Lea TYPE Behn R/ Coccinella religiosa Lea N.W. Australia TYPE 10375 (SAM, lectotype); C. vidua: Type/ Java/ 57.71/Vidua Muls. Java. /Named by Mulsant (BMNH, lectotype).

Specimens examined We have examined over 1500 specimens from the following countries: AUSTRALIA: Western Australia (Kimberley region); Northern Territory; Queensland; northern New South Wales; Lord Howe Island; Norfolk Island. INDONESIA: Manowari Ransiki, Mayuby; Fakfak; Sorong; Biak Is; Nabire; Numfoor Island. Kai Islands: Key-Ins. PAPUA NEW GUINEA: Finschhafen; Maffin Bay; Nadzab; Noemfor Island; New Britain; Morehead, Morehead River; Cyclops Mts; Angoram, Sepik River; Bosavi Mt. Lamington; Daru Island; Lake Murray; Green River; Laloki; New Ireland; Bougainville; SOLOMON ISLANDS: Santa Ysabel; Kolombangara I; Malaita; Guadalcanal. NEW CALEDONIA: Tiendanite; Tiea; Noumea. VANUATU: Mallicolo.

Distribution China, Malaysia, Philippines; Indonesia, Papua New Guinea, New Zealand, Australia, New Caledonia, Solomon Islands. Introduced to New Zealand, Hawaii and Florida (Gordon 1985; Pope 1989).

Coelophora vesiculata Chazeau, 1990

Distribution of Coelophora vesiculata

a very fine bead; disc with distinct punctation; interspaces with dense reticulation. Prosternal process with a pair of converging carinae, extending forward to the middle of the prosternum. Hypomeron with a distinct fovea anteriorly. Scutellum transverse, triangular, micropunctured. Elytra strongly convex with lateral borders narrowly explanate with a complete bead. Elytral punctures near the lateral margins much larger and variable in size, but always larger than those on the disc; punctures on the disc about as large as those on the pronotum, interspaces reticulate. Elytral epipleuron twice the width of the mesanepisternum with a distinct fovea at the middle. Abdomen with the posterior margin of ventrite 5 truncate in both sexes; ventrite 6 shallowly emarginate in males, rounded in females. Male terminalia (Figs 39F–I). Penis with the apical sclerite short and weakly upturned; the parameres slightly longer than the penis guide. Female terminalia. Spermatheca large and strongly C-shaped with several constrictions (Fig. 39E), sperm duct short and thick-walled. Measurements (n = 2): TL 4.70–5.30; PL 0.90–1.10; EL 3.80–4.10; PW 2.80–2.85; EW 4.40–4.65; GD 2.30–2.80. Ratios: PL/PW 0.32–0.39; EL/EW 0.86–0.88; EL/PL 3.73– 4.22; EL/GD 1.46–1.65; TL/EW 1.07–1.14.

Types C. vesiculata: Cacao, Ag. Stn., Buka, 23.10.64 TLF/ Coelophora vesiculata J. Chazeau det. 1988 (BMNH, holotype).

Specimens examined

Very similar to the melanic forms of Coelophora inaequalis and can only be distinguished from them by the shape of the spermatheca, and the penis that has a much shorter apical sclerite.

INDONESIA: Biak I., i.1945, J. R. Helfer (1, CAS). PAPUA NEW GUINEA: NE Bulolo, 750m, 5.x.1965, J. Sedlacek (1, BPBM); NE Bulolo, 700m, 25.x.1969, J. & M. Sedlacek (1, BPBM); NE Wau, 1150–1300m, 29.xii.1965, J. Sedlacek (1, BPBM); NE Wau Creek, Wau, 1200–1500m, 6.v.1963, J. Sedlacek (1, BPBM); Mimika R., 1911–229, F.R. Wollaston (1, BMNH); E. New Britain, 30km SW Kokopo, vic. Arabam, 200m, 21.ii-04.iii.2000, A. Weigel (1, ANIC).

Description

Distribution

(Fig. 39)

Coelophora vesiculata Chazeau, 1990: 157. Type locality: Papua New Guinea, Buka Island.

Diagnosis

Colour pattern (Figs 39A–D). Head and appendages orange or brown, rarely almost entirely black; pronotum orange, usually with a large black medial area not extending to the anterior margin; elytra black, with reddish or orange spots at the humeral and apical areas. These spots vary in size and rarely occupy the larger sections of the elytra, or are completely absent (Fig. 39B). Ventral side brown, with paler prothoracic hypomera, mesepimeron and lateral edges of abdominal ventrites. Morphology. Length 4.7–5.3 mm. Frons flat, sparsely setose, 1.5–2.0 times the eye width. Pronotum convex; lateral margins rounded, weakly sinuate before the anterior angles with

Endemic to the island of New Guinea.

Docimocaria Crotch, 1874 (Figs 40–42)

Docimocaria Crotch, 1874: 172. Type species by original designation, Docimocaria insignis Crotch, 1874. Antineda Iablokoff-Khnzorian, 1982: 141. Type species by original designation, Neda princeps Mulsant, 1850. Syn. nov.

References: Iablokoff-Khnzorian 1982: 136; Pope 1989: 644.

VII. Review of the Australo-Pacific Coccinellini

Diagnostic combination Length 6.0–12.2 mm; elytra strongly convex, glabrous. Elytral colour pattern variable, but orange or red, usually with black markings (Fig. 41). Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly shorter than the head capsule with a rather compact three-segmented club. Terminal maxillary palpomere strongly securiform. Pronotal disc evenly convex transversely; lateral margins slightly concave; external borders with a narrow bead. Prothoracic hypomeron with a shallow fovea near the anterior angles; prosternal process with distinct carinae extending anteriorly. Anterior margin of the mesoventrite deeply emarginate medially. Elytra with margins not explanate and without a clear bead; epipleuron distinctly foveate at the level of the hind coxae. Tibial spurformula 0–2–2. Abdominal postcoxal line not recurved, and incomplete laterally; an oblique postcoxal line absent. Penis very long and thin. Infundibulum absent; spermatheca with cornu strongly C-shaped; nodulus and ramus poorly developed.

Description Body strongly convex, almost hemispherical; dorsum glabrous. Head. Eyes partially hidden beneath the anterior margin of the pronotum, with inner orbits strongly divergent anteriorly. Frons narrow, about as broad as the eye. Antenna 11-segmented, with a three-segmented club. Anterior clypeal margin straight, between two distinct lateral projections. Terminal maxillary palpomere strongly securiform. Prothorax. Pronotum convex, the anterior angles projecting moderately. Prothoracic hypomeron with shallow foveae anteriorly. Prosternal process ~0.5 times as broad as the diameter of the procoxae, with convergent carinae extending anteriorly. Pterothorax. Scutellum 1/10–1/12 times as broad as the elytra at the shoulders. Elytra much broader than the pronotum at the base; lateral margins without complete beads. Epipleuron very broad, ~0.3 times as broad as the elytra, strongly inclined, with weakly delimited foveae. Mesoventrite emarginate medially, with anterior margin entirely carinate. Metaventral postcoxal lines joined medially, complete laterally; discrimen

39

complete. Legs slim and long; apex of the middle and hind tibiae each with 2 spurs; tarsal claws with a subquadrate basal tooth. Abdomen with six ventrites in both sexes. Abdominal postcoxal line separated medially, incomplete laterally, not recurved; oblique postcoxal line absent. Posterior border of ventrite 5 usually truncate in males, arcuate in females. Male terminalia. Penis consists of a single, slim and long sclerite; the apical part tube-like without additional sclerites; capsule T-shaped, usually the inner arm longer than the outer arm. Penis guide narrow at the base in ventral view, evenly tapering from the base to the apex; parameres thick, with long hairs. Female terminalia. Coxites slim and elongate, with short apical styli. Sperm duct slim and long, membranous, tube-like. Infundibulum absent. Spermatheca large, strongly curved and C-shaped, surface comparably smooth; ramus and nodulus indistinct.

Biology The lava of the Australian species D. princeps was described by Ślipiński (2007); it has been reared on Heperomyzus aphids on Sonchus by K.J. Houston in Brisbane (Pope 1989).

Distribution Australia, the island of New Guinea and the Philippines.

Remarks Docimocaria is very similar to Megalocaria, but can be distinguished from that genus by its usually smaller size, the elytra with very narrowly explanate lateral margins, the marginal elytral bead visible at least near the anterior corners, the oblique postcoxal line absent and the penis very long and thin and pointed apically. Docimocaria pulchra and D. aruensis can be easily confused with some colour variants of Harmonia manillana (Mulsant, 1866). However, they can be distinguished from Harmonia by having the terminal antennal segment elongate and apically rounded, the anterior margin of the mesoventrite straight and the oblique postcoxal line absent.

Key to species of the genus Docimocaria Crotch from Australo-Pacific region 1. – 2(1). –

Elytra yellow or orange, with a black suture and margins (Fig. 41)................................................................................... D. princeps Elytra with black and red or orange colour pattern (Fig. 40)............................................................................................................... 2 Elytra red, with two black transverse bands (Fig. 42A)....................................................................................................... D. pulchra Elytra black, with four red spots on elytra (Fig. 40A)........................................................................................................ D. aruensis

Docimocaria aruensis (Crotch, 1874) comb. nov. (Fig. 40)

Coelophora aruensis Crotch, 1874: 150. Type locality: Aru. Korschefsky 1932: 290; Gordon 1987: 18. Antineda aruensis: Iablokoff-Khnzorian 1982: 143.

Diagnosis Similar to D. pulchra in having a black and red elytral colour pattern, but easily distinguished by the orange or reddish spots limited to the humeral and apical areas only, and the penis with a very long apical tubular area (Fig. 40D).

Description Colour pattern (Figs 40A–C). Head and appendages yellow. Pronotum black, with orange-red patches near the anterior angles, narrowly connected medially; in some specimens the orange-yellow patches are strongly enlarged, reducing the black area to a narrow stripe along the posterior margin. Elytra black, with humeral and apical reddish patches arranged as Fig. 40A, external borders and suture black. Scutellum red medially, with black external borders, rarely completely black. Ventral side black, except for the orange prosternum, mesepimeron, mesanepisternum and the external borders of the abdomen. Prothoracic hypomeron yellow, except for the black external

Ladybird Beetles of the Australo-Pacific Region

40

Archaioneda princeps: Crotch 1874: 169; Korschefsky 1932: 271. Antineda princeps: Iablokoff-Khnzorian 1982: 142; Pope 1989: 645; Ślipiński 2007: 148.

Diagnosis Docimocaria princeps is the single species of this genus known from Australia and it is easily distinguished from all its congeners in having uniformly orange elytra without any maculation or fasciae. Distribution of Docimocaria aruensis

borders. Elytral epipleuron black, except for the pale anterior part. Anterior and middle legs entirely yellow or black on femur, hind legs black. Morphology. Length 8.7–9.5 mm. Frons at the middle 1.3– 1.5 times broader than the eye width, surface flat with punctures larger than the eye facets; interspaces reticulate. Pronotum widest before the base, lateral sides arcuate, narrower anteriorly than posteriorly, with a weak bead; disc moderately convex, densely punctate, with punctures smaller than those on the frons, 1–2 diameters apart, interspaces reticulate. Prosternal process with longitudinal carinae parallel anteriorly, stopping at one-third from the anterior border of the prosternum. Punctures on elytra slightly larger than those on the pronotum, with interspaces weakly reticulate; epipleuron broad, 3.1–3.6 times broader than the metanepisternum, with a shallow fossa for the hind femora. Anterior margin of the mesoventrite deeply emarginate, marginal carina weak but complete, posterior margin slightly arcuate. Abdominal postcoxal lines joined medially, laterally approaching the posterior margin of ventrite 1 and extending to the lateral borders; the oblique postcoxal line at least rudimentary, often clearly visible. Posterior borders of ventrite 5 evenly arcuate in males, ventrite 6 not visible externally. Male terminalia (Figs 40D–H). Female terminalia. Not examined. Measurements (n = 3): TL 8.70–9.50 (9.17 ± 0.42); PL 1.60 (1.60 ± 0.00); EL 6.80–7.80 (7.40 ± 0.53); PW 4.50–5.10 (4.87 ± 0.32); EW 8.00–9.10 (8.60 ± 0.56); GD 3.30–4.50 (4.03 ± 0.64). Ratios: PL/PW 0.31–0.36 (0.33 ± 0.02); EL/EW 0.85– 0.87 (0.86 ± 0.01); EL/PL 4.25–4.88 (4.63 ± 0.33); EL/GD 1.73–2.06 (1.85 ± 0.18); TL/EW 1.04–1.09 (1.07 ± 0.02).

Types C. aruensis: aruensis Wall/ aruensis T. Wall/ Aru (UCCC, holotype)

Specimens examined INDONESIA: Dutch New Guinea, Fak-fak (2, BMNH). New Guinea Island, coll. Sayer (1, BMNH); Neuguinea (1, MIIZ).

Distribution Islands of New Guinea and Aru.

Docimocaria princeps (Mulsant, 1850) comb. nov. (Fig. 41)

Neda princeps Mulsant, 1850: 278. Type locality: Australia, Port Essington. Callineda princeps: Crotch 1871: 6.

Description Colour pattern (Figs 41A–C). Head and appendages orangeyellow. Pronotum orange-yellow, with black lateral and basal borders, colour of the disc sometimes darker medially. Scutellum usually pale with dark borders, in some specimens completely black. Background colour of elytra orange-yellow, lateral borders broadly black, anterior borders and suture narrowly black. Ventral side background body colour yellow, except for the metaventrite and metanepisternum. In some specimens, the lateral and anterior margins of the mesoventrite are black, rather than yellow. Anterior and middle legs pale, hind legs usually dark, except for the tarsi which are occasionally entirely pale. Morphology. Length 8.5–9.0 mm. Frons at the middle slightly broader than the eye diameter, with obvious punctures much larger than the eye facets. Eyes partially visible dorsally from the pronotum. Punctures on the pronotum slightly smaller than those on the head, usually separated by > 1 diameter. Prosternal process with longitudinal carinae parallel anteriorly, stopping approximately one third from the anterior border of the prosternum. Punctures on the elytra larger than those on the pronotum, often separated by > 1 diameter. Anterior margin of the mesoventrite deeply emarginated, with raised carinae arcuate medially, the posterior margin almost straight. Abdominal disc broad, ~2/3 times as long as broad. Postcoxal line disconnected medially, laterally approaching theposterior of ventrite 1 and extending to the lateral borders. Apex of ventrite 5 weakly arcuate medially, apically emarginated; ventrite 6 hidden by ventrite 5. Male terminalia (Figs 41F–I). Penis slim and long, not recurved. Penis guide as long as the parameres; parameres evenly broadened from base to apex. Female terminalia (Figs 41D,E). Sperm duct very short. Measurements (n = 5): TL8.60–9.10 (8.84 ± 0.21); PL 1.80– 2.00 (1.88 ± 0.08); EL 6.70–7.20 (6.92 ± 0.19); PW 4.30–4.70 (4.48 ± 0.18); EW 7.60–8.30 (7.96 ± 0.27); GD 4.10–4.50 (4.26 ± 0.18). Ratios: PL/PW 0.40–0.44 (0.42 ± 0.02); EL/EW 0.85–0.88 (0.87 ± 0.01); EL/PL 3.50–3.79 (3.68 ± 0.12); EL/ GD 1.56–1.66 (1.63 ± 0.04); TL/EW 1.09–1.13 (1.11 ± 0.02)

Types N. princeps: Holotype O.U.M./ 32/ P. Ess/ Neda Princeps Mulsant 1850/ Ann. Soc. Agric. Lyon (2)2: 278/ T/ Type COL: 1291 Neda princeps Mulsant (OMNH).

Specimens examined AUSTRALIA: Northern Territory: Ferguson R. X-ing. Stuart Hwy, 21.vi.2000, M.S. Moulds(1, AM); Surprise Creek, 45 km SW by S of Borroloola, 5.xi.1975, M.S. Upton (2, ANIC); Cooper Creek, 19km E. by S. of Mt. Borradaile, 9.xi.1972, M.S. Upton (1, ANIC); Bessie Spring, 8km ESE of Cape Crawford, 26.x.1975, M. Upton (1, ANIC); Katherine Gorge, 24km NE

VII. Review of the Australo-Pacific Coccinellini

41

Diagnosis This is a very distinctive species due to the red or orange elytra bearing a broad black sutural stripe and two somewhat irregular transverse bands extending to the lateral margins (Fig. 42C).

Description

Distribution of Docimocaria princeps

of Katherine, 16.x.1972, M.S. Upton (1, ANIC); Katherine (Low level Pk), 20.viii.1973, L.P. Kelsey (2, ANIC); Darwin, 23.vi.1972, M.S. Upton (1, ANIC); Katherine, 1.xii.1963, J. Sedlacek (1, BPBM); McArthur River, 16.27S 136.05E, 48km SW by S of Borroloola, 29.x.1975, M.S. Upton (1, ANIC); 12.26S 130.56E, Holmes Jungle, Berrimah, 10km S of Darwin, 8.xi.1972, E. Britton (1, ANIC). Queensland: Coleman River, at light, 25.vi.1993, I.D. Naumann & P. Zborowski (2, ANIC); Moreton, at light, 10.xii.1992, W. Dressler & P. Zborowski (2, ANIC); 12.40S 142.40E, Batavia Downs Homestead, at light, 17–23.vi.1992, T. Weir (3, ANIC); Little River E of Croydon, at light, 16.iii.1985, K. Carnaby (1, ANIC); Bertle Ck. Xing, 12km SW Heathlands, at light, 15– 26.i.1992, T. Weir & I.D. Naumann (1, ANIC); Heathland nr. gravel pit, at light, xii.1992, W. Dressler & P. Zborowski (1, ANIC); The Boulders 5 km W of Babinda, 14–15.xi.1981, J. Balderson (1, ANIC); Wenlock R. X-ing, Cape York Pen.,13– 19.x.2001, M. DeBaar & G. Pegg (1, ANIC); 40 mile Scrub N.P., iii.1973 (1, ANIC); Archer River, rainforest, at light, 26.x.1993, P. Zborowski & D. Rentz (1, ANIC); Atherton Tableland, Millstream Nat. Pk, 18–28.xi.1998, Yee, Edwards, Calder, Halliday & Sutrisno (5, ANIC); 13.2701S 142.4154E, Mungkan Kandju Nat. Pk. Langi lagoon, 29.vi-5.vii.1998, M. Neave, T. Weir, M. Pratt & L. McKenzie (2, ANIC); Lynd R. X-ing, ESE Highbury, 23.iv.1983, J. F. Donaldson & J. F. Grimshaw (1, QDPI); 6.5 km ESE of New Laura Ranger Stn Lakefield NP., 26–29.vii.1998, A. Calder (1, ANIC); Hann River, at light, 17.xii.1993, P. Zborowski (1, ANIC). Western Australia: Kimberley Research Station, 31.iii.1964, K. T. Richards (1, ANIC).

Distribution Northern and north-eastern parts of Australia.

Colour pattern (Figs 42A–C). Head and appendages orangered. Pronotum black, with large orange-red lateral areas anteriorly narrowly and connected medially, sometimes forming a broad anterior band. Background colour of elytra black; each elytron with three large transverse red patches, sometimes the posterior two patches are connected into an extensive red apical area; the lateral borders and suture are black. Scutellum black. Ventral side black, except for the mesepimeron and mesanepisternum, external borders of the prosternum and the abdomen. Prothoracic hypomeron orange-red, except for a black posterior area. Elytra epipleuron usually orange-red, with black external borders. Legs orange-red. Morphology. Length 8.5–10.0 mm. Frons at the middle 1.2– 1.5 times broader than the eye width, punctate, with punctures much larger than the eye facets. Pronotum transverse; lateral margins arcuate, weakly sinuate behind the anterior angles, with a weak bead; disc convex, densely punctate, punctures smaller than those of the head, 1–2 diameters apart; interspaces reticulate. Prosternal process with longitudinal carinae slightly convergent anteriorly, ending at one third from the anterior border of the prosternum. Scutellum equilateral triangle, sharply pointed. Elytra with much larger punctures than the pronotum, separated by > 1 diameter. Anterior border of the mesoventrite deeply emarginated, raised carinae rounded medially, posterior margin slightly arcuate. Elytra convex, narrowly explanate laterally, without a bead except for the humeral angles. Postcoxal lines connected medially, laterally reaching the posterior margin of ventrite 1 and extending to the lateral borders (Fig. 42J); oblique line not visible. Posterior borders of ventrite 5 slightly arcuate in males and strongly arcuate in females. Ventrite 6 hidden by ventrite 5. Male terminalia (Figs 42F–I). Female terminalia (Figs 42D,E). Spermatheca large, with irregular complex shape; sperm duct long. Measurements (n = 10): TL 8.50–10.30 (9.39 ± 0.58); PL 1.60–2.20 (1.88 ± 0.17); EL 6.90–8.70 (7.59 ± 0.58); PW 4.30– 5.10 (4.71 ± 0.32); EW 8.00–9.20 (8.62 ± 0.43); GD 4.10–5.00 (4.51 ± 0.28). Ratios: PL/PW 0.37–0.43 (0.40 ± 0.02); EL/EW 0.83–0.99 (0.88 ± 0.04); EL/PL 3.68–4.58 (4.06 ± 0.27); EL/ GD 1.53–1.93 (1.68 ± 0.11); TL/EW 1.05–1.13 (1.09 ± 0.03).

Types C. pulchra: pulchra T Ceram. Wall/ Type pulchra Wall/ Type (UCCC, lectotype). C. thoracica: Key-Inseln, Tual, C. Ribbe, 1884/ Anisolemnia thoracica m/Holotypus Anisolemnia thoracica Wse (ZMB, lectotype, labelled as holotype by Iablokoff-Khnzorian).

Docimocaria pulchra (Crotch, 1874) (Fig. 42)

Coelophora pulchra Crotch, 1874: 150. Type locality: Ceram. Gordon 1987: 18 (lectotype). Caria thoracica Weise, 1892a: 283. Type locality: Key Island. Syn. nov. Anisolemnia thoracica: Korschefsky 1932: 270. Docimocaria thoracica: Iablokoff-Khnzorian 1982: 136 (lectotype).

Specimens examined INDONESIA: Cyclops Mts., 4km nord, Sentani, 600m, 8–13. ix.1990, Balke & Hendrich (1, ZMB). Manowari Ransiki, Mayuby, 300m, 26–30.ix.1990, A. Riedel (1, NHMS). Aru Islands: Aru Is. H. Elgner (ANIC); Aru Is., W. Froggatt Collection (ANIC); Aru (ANIC). Kai Islands: Ins. Kei (Planten), (ZMB; MNHN, ANIC). Seram: Ceram (MIIZ). PAPUA NEW GUINEA: Papua N. Guinea (1, ANIC).

42

Ladybird Beetles of the Australo-Pacific Region

Distribution of Docimocaria pulchra

Remarks Iablokoff-Khnzorian (1982) synonymised Coeophora pulchra with C. formosa and moved both species to Docimocaria. However, he clearly confused the identities of the species and failed to recognise that both C. pulchra and C. formosa are distinct species, with the former being a senior synonym of C. thoracica. The male specimen from Seram, which is identical to the lectotype of C. pulchra, has been dissected and the male terminalia conform to the specimens of males examined from New Guinea, Aru and Kai Islands.

Distribution The island of New Guinea; Aru, Seram and Kai Islands, Indonesia.

Harmonia Mulsant, 1850 (Figs 43–60)

Harmonia Mulsant, 1850: 74, 75. Type species by subsequent designation of Timberlake 1943: 17, Coccinella marginepunctata Schaller 1783. Leis Mulsant, 1850: 241. Type species by subsequent designation of Crotch 1874: 119, Coccinella dimidiata Fabricius, 1781. Synonymised by Iablokoff-Khnzorian 1979: 71. Ballia Mulsant, 1853: 159. Type species by subsequent designation of Crotch 1874: 126, Ballia christophori Mulsant, 1853. Synonymised by Iablokoff-Khnzorian 1979: 71. Callineda Crotch, 1871: 6. Type species by subsequent designation of Rye 1873: 329, Coccinella sedecimnotata Fabricius, 1801. Synonymised by Timberlake, 1943: 17. Stictoleis Crotch, 1874: 118. Type species by original designation, Coccinella coryphaea Guérin-Méneville, 1835. Synonymised by Iablokoff-Khnzorian 1979: 71. Ptychanatis Crotch, 1874: 122. Type species by original designation, Coccinella axyridis Pallas, 1773. Synonymised by Korschefsky 1932: 439. Rhopaloneda Timberlake, 1943: 17. Type species by original designation, Callineda decussata Crotch, 1874. Synonymised by Iablokoff-Khnzorian 1982: 458.

Diagnostic combination Length 4–11 mm. Elytra moderately to strongly convex, glabrous. Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly shorter than the head capsule with a compact three-segmented club; terminal antennomere strongly broadened and truncate apically. Terminal maxillary palpomere securiform. Pronotal disc evenly convex transversely, with narrowly upturned or slightly thickened external borders. Prothoracic hypomeron without

foveae near the anterior angles; prosternal process with distinct carinae extending anteriorly but never reaching the anterior border. Anterior margin of the mesoventrite very shallowly emarginate medially or straight. Elytra with lateral margins with complete beads; epipleuron without foveae. Femurs of mid and hind legs do not extend beyond the border of the elytra. Tibial spur formula 0–0–0. Abdominal postcoxal line not recurved, incomplete laterally with an oblique postcoxal line often connected at its end. Penis consists of a basal sclerite and an additional apical piece; tegmen stout, penis guide not extending beyond the parameres apically, strongly bent towards the parameres in lateral view. Ovipositor with coxites shaped like club handles, infundibulum present; spermatheca C-shaped with a narrowed ramus, nodulus usually absent.

Description Body outline varies from strongly convex and hemispherical, to weakly convex and elongate-oval; dorsum glabrous. Head. Frons usually broader than the eye width. Eyes partially obscured by the pronotum, with inner orbits parallel or slightly divergent. Antenna 11-segmented, shorter than the width of the head capsule; scape not expanded; pedicel distinctly narrower than scape; antennal club three-segmented, tightly articulated; terminal palpomere broad, expanding before becoming truncate apically. Clypeus short, transverse, the anterior border straight medially between two lateral projections. Mandible bifid apically, molar part with a basal tooth, prostheca distinct. Terminal maxillary palpomere securiform. Prothorax 0.6–0.8 times as broad as the elytra at the base, anterior margin of the pronotum deeply emarginate. Prothoracic hypomeron without lateral foveae; prosternal process usually with admedian carinae that sometimes extend forward to the middle of the prosternum. Pterothorax. Scutellum triangular, pointed apically. Mesoventrite with the anterior margin almost straight, at most slightly emarginated medially. Metaventral postcoxal line complete laterally. Elytral epipleuron flat or weakly inclined, without delimited foveae. Tibiae without apical spurs; tarsal claws appendiculate, with a basal tooth. Abdomen with six ventrites in both sexes. Intercoxal process rounded apically. Abdominal postcoxal line extends to the hind margin of ventrite 1, incomplete laterally; the oblique postcoxal line is very close to or connected to the end of the postcoxal line (Fig. 43L). Posterior margin of ventrite 5 usually truncate in males and arcuate in females. Male terminalia. Penis as long as or slightly shorter than the length of the abdominal disc, strongly constricted near the apex, forming a very narrow apical section connected to a membranous apical pouch; outer arm of the penis capsule comparably long; the inner arm usually strongly curved. Tegmen stout, with the strut cleft apically; the penis guide as long as or slightly shorter than the parameres, tapering to the apex, strongly curved in lateral view. Female terminalia. Spermatheca strongly curved to C-shaped; nodulus slimmer than cornu; ramus indistinct, located on the inner side of the cornu. Sperm duct simple, uniform in diameter; infundibulum heavily sclerotised, tube like, enclosing the sperm duct. Coxites shaped like club handles, with apical styli.

Distribution Harmonia is a moderately large genus, with ~30 species (Coutanceau 2008) distributed throughout the Old World, but particularly abundant in the Oriental and Australo-Pacific regions.

VII. Review of the Australo-Pacific Coccinellini

43

Key to Harmonia species from the Australo-Pacific Region Elytra with a transverse preapical fold ‘plica’ (Fig. 45C); colour pattern extremely variable.............................................H. axyridis Elytra smooth apically, without folds.................................................................................................................................................. 2 Epipleuron very broad, at least 3.5 times as broad as the metanepisternum; body large and strongly convex (Fig. 52)...... H. manillana Epipleuron much narrower, at most 2.5 times broader than the metanepisternum; body shape variable but usually much less convex than above..................................................................................................................................................................................... 3 3(2). Elytral border near the humeral angle raised and forming a distinct gutter; body elongate-oval (Fig. 56) and weakly convex........... ����������������������������������������������������������������������������������������������������������������������������������������������������������������������������H. octomaculata – Elytral border thickened, but not raised to form a gutter along the humeral angle; body usually broader, oval or almost circular and convex........................................................................................................................................................................................... 4 4(3). Elytra yellow, with 10 large and circular black spots on each elytron (Fig. 49A)........................................................... H. conformis – Elytra with different colour pattern than above................................................................................................................................... 5 5(6). Pacific species...................................................................................................................................................................................... 6 – Australian and Papuan species............................................................................................................................................................. 7 6(5). Body relatively flat and elongate (Fig. 43). Elytra without a defined colour pattern or with irregular transverse black bands (Fig. 43A)................................................................................................................................................................. H. antipodum – Body oval and convex; pronotum orange or reddish, without any markings; elytra red with three transverse black fasciae (Fig. 55A)...................................................................................................................................................... H. novaehebridensis – Body oval and convex; pronotum orange bearing a medial trapezoidal black marking; elytra with two sets of spots (Fig. 51B)....... ������������������������������������������������������������������������������������������������������������������������������������������������������������������������������������� H. fijiensis 7(5). Prosternal carinae well developed, anteriorly separated from lateral borders, converging and extending forward to almost the middle of the prosternum.............................................................................................................................................................. 8 – Prosternal carinae absent or forming lateral borders of the prosternal process and do not extend beyond the anterior margin of the prosternal process........................................................................................................................................................................ 13 8(7). Elytra yellowish or yellowish-green, with black markings.................................................................................................................. 9 – Elytral background colour black, with yellow or orange markings................................................................................................... 10 9(8). Larger species, length 8.8–10.0 mm; elytra light yellow and mostly translucent with three isolated black spots on the disc; sutural margin and lateral margins pale or dark; male genitalia as in Fig. 48............................................................................ H. boothi – Smaller species, length 5–8 mm; elytra yellow to orange, with a network of black stripes that can be reduced in size to very short humeral markings (Fig. 59B); sutural and lateral margins with broad black borders; male genitalia as in Figs 59E–H............... H. testudinaria – Smaller species, length 5–7 mm; elytra yellow, with black isolated spots (Fig. 46A); sutural and lateral margins narrowly black; male genitalia as in Figs 46E–G....................................................................................................................H. basinotata [part] 10(8). Elytral epipleuron entirely yellow or orange........................................................................................................................H. tricolor – Elytral epipleuron at least partially black.......................................................................................................................................... 11 11(10). Legs entirely black, elytra black with four orange spots (Fig. 57A)...................................................................................H. ohlmusi – Legs yellow; elytra with a different colour pattern than above......................................................................................................... 12 12(11). Elytra entirely black or each with a small circular orange humeral spot (Fig. 47A)................................................. H. bicolor [part] – Elytra with large orange basal and apical areas (Fig. 60A)................................................................................................... H. tobleri 13(7). Small species, length 4.5–5.0 mm...................................................................................................................................................... 14 – Larger species, length at least 5.5 mm............................................................................................................................................... 15 14(13). Elytron with two small round spots (Fig. 53A);..................................................................................................................... H. micra – Elytron with a reticulate black pattern on yellow background (Fig. 44A).............................................................. H. areolata [part] 15(13). Lateral margins of the elytra at least partially pale (Fig. 50).................................................................................H. decussata [part] – Elytra black or with a continuous black border................................................................................................................................. 16 16(15). Elytra entirely black, or each with a single red humeral spot not extending to the lateral border (Fig. 47B); pronotum black except for the yellow narrow anterior part; large (6.0–7.5 mm long) and strongly convex species; from Australia............. H. bicolor [part] – Elytra and pronotum differently coloured than above; Papuan species............................................................................................. 17 17(16). Frons between the eyes narrow, maximum twice the eye width (Fig. 46D); elytra usually with variously developed black reticulate colour pattern or with extensive markings on a yellow or reddish background (Fig. 50).......................................................... 18 – Frons between the eyes at least 2.5 times as wide as the eye width; elytra with small isolated spots (Fig. 46)........ H. basinotata [part] 18(17). Elytral patterns variable, from almost entirely black to a black reticulate pattern on an orange or red background. If pattern reticulate and elytral border entirely black, then sutural border broad and dorsal complete (Fig. 50); male and female terminalia as in Figs 50F–H.................................................................................................................................................H. decussata [part] – Elytra with very narrow black sutural and lateral borders, disc usually with an incomplete fine reticulate pattern (Fig. 54D); male and female terminalia as in Figs 54F–J............................................................................................................H. nigromarginata – Elytra with a black reticulate colour pattern on a yellow background (Fig. 44A); lateral and sutural margins broadly black; male and female terminalia as in Figs 44E–I............................................................................................................. H. areolata [part] 1. – 2(1). –

Harmonia antipodum (Mulsant, 1848) (Fig. 43)

Coccinella antipodum Mulsant in White, 1848: 66. Type locality: New Zealand. Mulsant 1850: 1019; 1866: 182; Crotch 1874: 119. Leis antipodum: Broun 1880: 645; Korschefsky 1931: 273. Harmonia antipodum: Iablokoff-Khnzorian 1982: 463.

Diagnosis This is the only endemic species of Harmonia known from New Zealand. It can be easily recognised by its relatively elongate and weakly convex body, and the often not clearly defined lighter colour pattern on its olive-green elytra.

44

Ladybird Beetles of the Australo-Pacific Region

Description Colour pattern (Figs 43A–D). Typical form: frons and head appendages light yellow; pronotum yellow, with two oblique black fasciae medially; scutellum olive-green; elytra olive-green, except for a darker external border and a yellowish triangular area centrally (Fig. 43A); legs and venter yellowish. Melanic form: frons yellow with a black area at the base; head appendages yellow; pronotum yellow with a large black marking medially; scutellum black; elytra yellow or orange, with seven black markings arranged in three irregular transverse rows (Figs 43B,C). Morphology. Length 5.3–5.5 mm. Frons almost twice as broad as the eye width. Eyes largely exposed, inner orbits almost parallel. Antenna 1.5 times the width of the frons; terminal antennomere moderately broadened, truncate apically. Prosternal process with short lateral carinae, straight or extending slightly forward. Elytra densely punctured, broadest in the middle. Elytral epipleura 2.1–2.3 times as broad as the metanepisternum. Metaventral postcoxal line almost straight apically. Abdominal postcoxal (Fig. 43L) line relatively short, distinctly separated from the posterior margin of ventrite 1, connected to the oblique postcoxal line. Posterior margin of ventrite 5 emarginate medially in males, and broadly arcuate medially in females. Male terminalia (Figs 43H–J). Penis with an apical pouch bearing two sclerites (Figs 43H,I). Female terminalia (Figs 43F,G). Infundibulum weakly sclerotised, sperm duct very short (Fig. 43G). Measurements (n = 5): TL 5.30–5.50 (5.35 ± 0.06); PL 0.90–1.00 (0.98 ± 0.05); EL 4.30–4.50 (4.38 ± 0.10); PW 2.50– 2.80 (2.65 ± 0.13); EW 4.00–4.40 (4.23 ± 0.17); GD 2.00–2.30 (2.20 ± 0.14). Ratios: PL/PW 0.33–0.40 (0.37 ± 0.03); EL/EW 1.00–1.08 (1.04 ± 0.03); EL/PL 4.30–5.00 (4.50 ± 0.34); EL/ GD 1.87–2.20 (2.00 ± 0.14); TL/EW 1.23–1.33(1.27 ± 0.05).

Types C. antipodum: ‘antipodum White Ann. Nat. Hist.2nd ser I. p 66 (type New Zealand.’ (BMNH, holotype).

Specimens examined NEW ZEALAND: Konoronoto, 12.ix.1908, G. V. Hudson (1, BMNH); In Astelia Rimu Tree, 25.viii.1921, G. V. Hudson (1, BMNH); L. Guyon, 93–60, Pascoe (2, BMNH); AK Titirangi, 1.iv.1998, A. Crowe (1, NZAC); BP lake Rotoehu, sweeping,

15.ii.1979, J. S. Dugdale (1, NZAC); TO Ohakune, 640m, 27.i.1982, C. F. Butcher (1, NZAC); AK 67 Park Roi Titirangi, 23.v.1980, B. G. Bennett (1, NZAC); Puresoa Forest, 4.vi.1965, D. Read (1, NZAC).

Distribution Endemic to the North Island of New Zealand.

Harmonia areolata Bielawski, 1968 (Fig. 44)

Harmonia areolata Bielawski in Bielawski and Chûjô, 1968: 123. Type locality: Ambon Island. Harmonia nigromarginata: Iablokoff-Khnzorian 1982: 487.

Diagnosis The reticulate colour pattern of Harmonia areolata resembles that of H. nigromaginata and some colour forms of H. decussata (Figs 50A,C,D), but it can be easily distinguished from both these species by its smaller and weakly convex body and largely black ventral surfaces.

Description Colour pattern (Figs 44A–C). Frons and appendages yellowish. Eyes black. Pronotum yellow with two longitudinal black stripes that do not reach the anterior border, basal margin between stripes black. Scutellum black; elytral background yellow, with a dark pattern consisting of longitudinal and transverse stripes, arranged as Figs 44A,B; borders and suture black. Prosternum yellow, prosternal process dark, hypomeron yellow. Pterothorax black or dark, brown except for pale mesepimeron. Elytral epipleuron yellow, with an entirely black external margin. Legs usually yellow with dark coxae. Morphology. Length 4.2–4.9 mm. Frons between the eyes ~1.5–1.7 times the eye width. Eyes large, in repose partially beneath the anterior margin of the pronotum, orbits slightly divergent. Antenna 1.0–1.2 times the frons width, with a compact club; terminal antennomere strongly securiform. Prosternal process broad, with short parallel lateral carinae extending forward slightly. Scutellum at the base 1/14 times as broad as the elytra at the shoulder area. Elytral disc with very fine punctures; interspaces smooth and shiny; suture with micropunctures arranged in lines and parallel. Elytral epipleuron ~2.4–2.7 times as broad as the metanepisternum. Metaventral postcoxal lines thicker at the ends. Abdominal postcoxal line reaching the posterior margin of ventrite 1, connected with a long oblique postcoxal line (Fig. 44J). Posterior margin of ventrite 5 truncate in males, slightly arcuate medially in females. Male terminalia (Figs 44F–I). Female terminalia (Fig. 44E). Measurements (n = 5): TL 4.25–4.90 (4.54 ± 0.28); PL 0.65–0.95 (0.76 ± 0.11); EL 3.40–4.00 (3.73 ± 0.22); PW 2.15– 2.40 (2.28 ± 0.09); EW 3.55–3.90 (3.71 ± 0.11); GD 1.70–2.00 (1.92 ± 0.13). Ratios: PL/PW 0.29–0.40 (0.32 ± 0.03); EL/EW 0.92–1.08 (1.00 ± 0.08); EL/PL 4.11–5.54 (5.05 ± 0.22); EL/ GD 1.85–2.00 (1.95 ± 0.09); TL/EW 1.15–1.32 (1.23 ± 0.09).

Remarks

Distribution of Harmonia antipodum

Iablokoff-Khnzorian (1982) synonymised Harmonia areolata with H. nigromarginata, but after examination of the type specimens and several specimens from New Guinea, we consider them distinct species with small but consistent external differences, and a very distinctively shaped infundibulum in the female terminalia of each species (Figs 44E, 54F).

VII. Review of the Australo-Pacific Coccinellini

45

Distribution of Harmonia areolata

Types H. areolata: Is. Ambon, Indonesia, 15.viii.1960, A. M. R. Wegner/ Alt. ca. 70m, at light/ Harmonia areolata Bielawski, 1968/ Holotype male (ELKU, holotype).

Specimens examined INDONESIA: Star Mt. Sibil Val., 1245m, 18.x-8.xi.1961, S. & L. Quate (1, BPBM); 4km ost. Fak-fak, 7.vii.1996, Schule & Stuben (1, NHMS). PAPUA NEW GUINEA: Amazon Bay area, Dogon, 2300ft., 13.ix-11.xii.1962, W. Brandt (2, ANIC); NE Wau, 1200m, 3.ii.1966, J. Sedlacek (2, BPBM); SE New Guinea: Kokoda, 400m, 20.xi.1965, J. Sedlacek (1, BPBM); NE Kokoda, 400m, light trap 15–20.xi.1965, J. Sedlacek (1, BPBM); Kokoda. l. 200ft. viii.1933, L. E. Cheesman (2, BMNH); Madang Dist., 2–11.x.1964 (1, BMNH); Finsch Haven dist. (1, SAM); Madang Dist., Finisterre Mts. Budemu c. 4000 ft. 15–24.x.1964 (2, BMNH); Finisterre Mts, 4000ft, 15.ix.1957, Munroe & Holland (1, ANIC).

Distribution The islands of Ambon and New Guinea.

Harmonia axyridis (Pallas, 1773) (Fig. 45)

Coccinella axyridis Pallas, 1773: 726. Type locality: Russia: Siberia, Yeniseysk. Leis axyridis: Mulsant 1850: 226. Ptychanatis axyridis: Crotch 1874: 124. Harmonia axyridis: Timberlake 1943: 17; Sasaji 1971: 278; Iablokoff-Khnzorian 1982: 476.

Diagnosis This adventive species is easily distinguished from other Harmonia species known from the Australo-Pacific region by its apical fold ‘plica’ on the apical section of the elytra (Figs 45A–D).

Description Colour pattern (Figs 45A–F). Several colour forms coexist in a single population (Fig. 4B). Colours vary from almost uniformly yellow, to small paired black markings on the pronotum; the most common forms have black and white pronotal markings and yellow or orange elytra bearing black spots of varying sizes, to almost entirely black, melanic forms with yellow lateral areas on the pronotum and yellow spots on the elytra. Morphology. Length 5.1–8.5 mm. Frons between the eyes 2.2–2.5 times broader than the eye. Eyes partially beneath the pronotum, with the interior orbits slightly divergent. Antenna

Distribution of Harmonia axyridis

1.5–1.7 times as long as the breadth of the frons; the antennal club compact; the terminal antennomere moderately broadened. Pronotum with the anterior margin slightly arcuate centrally; the lateral margin weakly arched. Disc very finely punctate, interspaces densely shagreened. Prosternal process with carinae extending anteriorly to at least the middle of the prosternum. Elytra with a subapical transverse plica almost always visible; epipleuron 1.2–1.5 times broader than the mesanepisternum. Scutellum ~1/13 times as broad as the elytra at the humeri. Mesoventrite with the anterior margin weakly arcuate medially. Abdominal postcoxal line (Fig. 45L) distinctly separated from the posterior margin of ventrite 1, usually connected to a weak oblique postcoxal line. Posterior margin of the 5th ventrite truncate in males, arcuate medially in females. Male genitalia (Figs 45H–K). Female genitalia (Fig. 45G). Measurements. TL 5.1–8.5; PL 1.0–1.5; EL 3.9–5.7; PW 2.4–3.4; EW 3.9–6.1; GD 1.7–2.9. Ratios: PL/PW 0.44–0.47; EL/EW 0.9–1.0; EL/PL 3.81–3.95; EL/GD 1.93–1.29; TL/EW 1.32–1.43.

Distribution Harmonia axyridis, also known as the multicoloured Asian lady beetle, or the harlequin lady beetle, is distributed in Siberian Russia, Mongolia, North and South Korea, northern China and Japan. It was introduced as a biological control agent to the USA and Europe, and quickly became established across entire continents. Further introductions or invasions established this species in South America (Chile) and Africa. It has recently become established in New Zealand. It is very common in some areas of the North Island, in particular around the Bay of Plenty, Coromandel, Waikato and Auckland (B. Boyd, pers. comm.; Fig. 4B).

Harmonia basinotata Bielawski, 1964b (Fig. 46)

Harmonia basinotata Bielawski, 1964b: 9. Type locality: Papua New Guinea, Dimifa. Iablokoff-Khnzorian 1982: 493. Harmonia uninotata Bielawski, 1964b: 11. Type locality: Papua New Guinea, Mount Otto. Syn. nov.

46

Ladybird Beetles of the Australo-Pacific Region

Diagnosis This species is very distinctive due to its relatively elongateoval body, yellowish colouration of its dorsal side, paired black irregular markings on the pronotum and two or six small isolated spots on each elytron (Figs 46A,B).

Description Colour pattern (Figs 46A–D). Frons yellow, with a black anterior margin and triangular area between the eyes; labrum dark brown or black; antenna and maxilla dark. Pronotum yellow, with black paired patches; hypomeron yellow, except for a black part at the posterior angles; prosternum black. Scutellum black. Elytra yellow, with two or six black spots on each elytron arranged as Figs 46A,B; the lateral margin and suture narrowly black. Legs black. Background colour of the ventral surfaces black; mesepimeron pale. Morphology. Length 5.5–7.1 mm. Frons between the eyes about twice as broad as the eye. Eyes in repose slightly beneath the anterior part of the pronotum. Antenna length 1.2– 1.3 times the frontal width; the terminal antennomere rounded on one side, slightly angulated on the other. Pronotum 0.8 times as broad as the elytra at the base; the anterior margin slightly arcuate medially, the lateral margin arcuate evenly. Prosternal process with short carinae on the surface, not or only slightly extending forward beyond the prosternal process. Scutellum ~1/10 times as broad as the elytra. Elytra yellow, usually with two or six black spots on each elytron, the lateral margin and suture with narrow black borders. Mesoventrite with an anterior margin weakly emarginate in the middle. Elytral epipleuron narrow, ~2.5 times as broad as the metanepisternum. Abdominal postcoxal lines not connected medially, approaching the posterior margin of ventrite 1; oblique postcoxal line (Fig. 46J) usually separate and very fine. Posterior margin of ventrite 5 truncate in males and arcuate in females. Male terminalia (Figs 46E–G). Penis (Fig. 46F) with an apical pouch bearing two sclerites; parameres slim and almost straight, with sparse setae apically. Female terminalia (Figs 46H,I). Spermatheca strongly curved; nodulus slim, ramus absent; sperm duct short; infundibulum slim, not flared; bursa copulatrix lightly sclerotised apically. Measurements (n = 4): TL 5.70–7.10 (6.24 ± 0.46); PL 0.80–1.00 (0.91 ± 0.08); EL 4.90–6.00 (5.31 ± 0.39); PW 2.60– 3.10 (2.81 ± 0.19); EW 4.50–5.40 (5.01 ± 0.35); GD 2.00–3.00 (2.60 ± 0.29). Ratios: PL/PW 0.29–0.38 (0.32 ± 0.03); EL/EW 1.00–1.13 (1.06 ± 0.05); EL/PL 5.10–6.69 (5.86 ± 0.55); EL/ GD 1.93–2.45 (2.06 ± 0.15); TL/EW 1.18–1.34 (1.25 ± 0.06).

Types H. basinotata: Dimifa, SE of Mt. Giluwe, S. Highlands, 2200 m, SE New Guinea (Papua), 10.x.1958, J.L. Gressitt (BPBM, holotype). H. uninotata: New Guinea: (NE) Mt. Otto, 2200 m, 23.vi. and 24.vi.1955, J. L. Gressitt (BPBM, holotype; MIIZ, paratype).

Specimens examined PAPUA NEW GUINEA: NE, Tambul, 2200m, 8.vi.1963, J. Sedlacek (BPBM; ANIC); NE, Lake Sirunki, 2550m, 18.vi.1963, J. Sedlacek (2, BPBM); NE, Yaibos, 1650m, 10–11. vi.1963, J. Sedlacek (1, BPBM).

Distribution of Harmonia basinotata

Remarks We could not find any differences between the female genitalia of H. uninotata and H. basinotata and therefore decided to synonymise these species. Iablokoff-Khnzorian’s (1982) interpretation of H. basinotata is probably erroneous, as in his book he illustrated a specimen of H. testudinaria as H. basinotata.

Distribution Endemic to New Guinea.

Harmonia bicolor (Blackburn, 1892) (Fig. 47)

Neda bicolor Blackburn, 1892: 238, Type locality: Australia, Queensland. Archaioneda bicolor: Korschefsky 1932: 271. Harmonia bicolor: Timberlake 1943: 18; Iablokoff-Khnzorian 1979: 72; 1982: 496; Pope 1989: 691 (lectotype); Ślipiński 2007: 163. Neda bicolor var. picturata Blackburn, 1895: 239.

Diagnosis This is the only black Harmonia species known from Australia, where it may be confused with the usually smaller Microcaria jansoni, with its uniformly orange or red pronotum. Uniformly dark forms of H. decussata can be distinguished from H. bicolor by their smaller size, and the prosternal carinae being much shorter and not extending forward beyond the base of the prosternal process.

Description Colour pattern (Figs 47A–D). Frons and head appendages yellow or brown. Pronotum largely black, except for a small yellowish area at the anterior angles. Scutellum black. Elytra uniformly black, or with one orange spot close to the shoulder area on each elytron. Ventral side yellow or brown, except for the black epipleuron. Legs usually brown. Morphology. Length 5–7 mm. Frons between the eyes ~1.4– 1.6 times as broad as the eye width. Antenna slightly longer than the frons width; the terminal antennomere broadened. Pronotum ~0.6 times as broad as the elytra; the anterior margin slightly arcuate centrally, the lateral margin evenly arcuate. Prosternal process with admedian carinae extending forward to half way along the prosternum. Scutellum ~1/10 times as broad as the elytra. Elytral epipleuron narrow, about twice as broad as the metanepisternum. Mesoventrite with anterior margin slightly emarginate in the middle; posterior margin straight. Metaventral postcoxal line almost straight apically. Abdomen

VII. Review of the Australo-Pacific Coccinellini

47

Distribution of Harmonia boothi

Distribution of Harmonia bicolor

~0.7 times as long as broad, the abdominal postcoxal lines separated at the middle, approaching the posterior margin of ventrite 1, joined by a very fine oblique postcoxal line (Fig. 47K). Posterior margin of ventrite 5 emarginate centrally in males, truncate in females. Male terminalia (Figs 47G–J). Female terminalia (Figs 47E,F). Measurements (n = 6): TL 5.10–6.90 (6.34 ± 0.56); PL 0.90– 1.10 (1.01 ± 0.06); EL 4.15–5.80 (5.29 ± 0.55); PW 2.35–3.80 (3.31 ± 0.48); EW 4.35–6.10 (5.58 ± 0.60); GD 2.45–3.20 (2.97 ± 0.24). Ratios: PL/PW 0.26–0.38 (0.31 ± 0.04); EL/EW 0.90–1.04 (0.95 ± 0.04); EL/PL 4.61–5.80 (5.26 ± 0.50); EL/GD 1.68–2.00 (1.78 ± 0.10); TL/EW 1.09–1.23 (1.14 ± 0.05).

Types N. bicolor: Type/ 4428 N. Qu. T/ Neda bicolor, Blackb. (BMNH, lectotype).

Specimens examined AUSTRALIA: Queensland: Little Mitchel River nr Yalkula, K. J. Houston (4, ANIC); Cardstone, 23.i.1965, J.G. & J.A.G. Brooks (1, ANIC); Mt. Etna, 25.i.1936 (1, ANIC); Daintree, 11.vi.1971, E.F. Riek (1, ANIC); nr Cairns, 14.vii.1969, J.E. Tobler (2, CAS); Southedge via Mareeba, 27.viii.1993, J. Watson (1, QDPI); Clump Pt., x.1952, J. Sedlacek (1, CAS).

stripes beginning at the posterior margin but not reaching the anterior margin, the base between stripes black. Elytra light yellow, semi-transparent, with black short humeral fascia and two round black markings on each elytron (Fig. 48A), the external borders yellow or slightly brown; suture narrowly black. Scutellum light yellow, with lateral borders black. Venter pale. Morphology. Length 8.7–10.2 mm. Eyes almost completely exposed. Frons between the eyes ~1.7–1.9 times as broad as the eye width. Terminal antennomere moderately broadened, flat. Pronotum weakly convex, ~0.6 times as broad as the elytra, lateral border arcuate. Prosternal process with carinae slightly convergent anteriorly, extending to the middle of the prosternum. Scutellum ~1/14 times as broad as the elytra. Elytra epipleuron ~1/8 times as broad as elytra. Anterior margin of mesoventrite straight. Abdominal postcoxal lines separated medially, approaching the posterior margin of ventrite 1; oblique postcoxal line not connected to abdominal postcoxal line. Posterior margin of ventrite 5 broadly emarginate in males and almost truncate in females. Male terminalia (Figs 48E–I). Female terminalia (Fig. 48D). Measurements (n = 4): TL 8.70–10.20 (9.48 ± 0.60); PL 1.50–1.80 (1.62 ± 0.11); EL 7.00–8.50 (7.71 ± 0.52); PW 3.80– 4.70 (4.38 ± 0.28); EW 7.20–8.50 (7.92 ± 0.43); GD 3.40–4.10 (3.87 ± 0.25). Ratios: PL/PW 0.33–0.40 (0.37 ± 0.02); EL/EW 0.88–1.01 (0.97 ± O.04); EL/PL 4.12–5.07 (4.78 ± 0.30); EL/ GD 1.88–2.07 (2.00 ± 0.06); TL/EW 1.11–1.25 (1.20 ± 0.04).

Specimens examined

Distribution

Holotype male: INDONESIA: INDONESIA: N.C. SERAM, Manusela N.P., Wae, Mual Plain, lowland Alluvial forest, 25.vii9.ix.1987 (BMNH). Paratypes: same data as the holotype (11, BMNH).

This species is known only from the Atherton Tablelands and adjacent coastal areas in northern Queensland.

Distribution

Harmonia boothi sp. nov. (Fig. 48) Diagnosis Harmonia boothi is similar to some pale colour forms of H. testudinaria found in New Guinea, but can easily be distinguished by its much larger body size, lightly coloured lateral margins of the elytra and its distinctive male genitalia.

Descriptions Colour pattern (Figs 48A–C). Head and appendages light yellow. Pronotum light yellow, with two longitudinal black

Known only from Seram (Maluku), Indonesia.

Remarks This species is named after Dr Roger G. Booth of the Natural History Museum of London, who identified this species as undescribed in the BMNH collections.

Harmonia conformis (Boisduval, 1835) (Fig. 49)

Coccinella conformis Boisduval, 1835: 604. Type locality: Australia. Crotch 1874:118. Leis conformis: Mulsant 1850: 261; 1866: 182; Crotch 1871:5; Korschefsky 1932: 273.

48

Ladybird Beetles of the Australo-Pacific Region

Distribution of Harmonia conformis

Harmonia conformis: Timberlake 1943: 17; Iablokoff-Khnzorian 1979: 71; 1982: 471; Pope 1989: 489 (lectotype); Ślipiński 2007: 163. Leis conformis var. occidentalis Lea, 1902: 289; Korschefsky 1932: 273.

Diagnosis Harmonia conformis is the most common and widely distributed species of Coccinellini in Australia. It can be easily distinguished by its relatively constant elytral colour pattern, consisting of 10 round black spots on each elytron on a yellowish or orange background (Fig. 49A).

Description Colour pattern (Figs 49A–C). Clypeus and most of frons yellow, vertex and lateral margins usually black, head appendages brown. Pronotum dark yellow, with four black markings, the lateral and anterior borders transparent. Scutellum black. Elytra dark yellow, with 10 black markings on each elytron arranged in four rows as 3–3–3–1, with two adjacent marking across the suture on each elytron collectively forming a scuttelar and a subapical spots; the lateral borders and suture narrowly black. Ventral side usually black, except for the pronotal hypomeron, elytral epipleuron, mesepimeron and metepimeron, the extreme external margin of elytral epipleuron, and the lateral abdominal margins. Legs usually with at least part of the femur black. Morphology. Length 5.1–8.2 mm. Frons between the eyes about twice as broad as the eye width. Terminal antennomere slightly broadened. Pronotum moderately convex, lateral margin evenly arcuate, anterior margin slightly arcuate. Surface of the prosternal process with admedian carinae extending forward beyond the prosternal process to the middle of the prosternum. Elytra with lateral margins slightly explanate; epipleuron ~2.5 times as broad as the metanepisternum. Mesoventrite with a straight anterior border. Metaventral postcoxal line distinctly descending. Abdomen ~0.7 times as long as broad; abdominal postcoxal lines separated at the middle, approaching the posterior margin of ventrite 1 (Fig. 49K); the oblique postcoxal line usually not connected to the abdominal postcoxal line.

Posterior margin of ventrite 5 deeply emarginate in males, truncate in females. Male terminalia (Figs 49F–J). Female terminalia (Figs 49D,E). Measurements (n = 10): TL 5.10–8.20 (6.78 ± 1.02); PL 0.90–1.50 (1.20 ± 0.18); EL 4.20–6.70 (5.51 ± 0.85); PW 2.40– 3.80 (3.07 ± 0.46); EW 3.90–6.50 (5.37 ± 0.82); GD 2.20–3.20 (2.76 ± 0.36). Ratios: PL/PW 0.32–0.46 (0.39 ± 0.05); EL/EW 0.93–1.10 (1.03 ± 0.05); EL/PL 3.83–5.70 (4.63 ± 0.66); EL/ GD 1.88–2.09 (1.99 ± 0.07); TL/EW 1.18–1.32 (1.26 ± 0.05).

Specimens examined AUSTRALIA: very common and widely distributed species. Over 2000 specimens from Queensland, New South Wales, Australian Capital Territory, Tasmania, South Australia and southern part of Western Australia have been examined (see also Pope 1989). NEW ZEALAND: AK Epsom Gov. House, 21.iv.1990, P. A. Maddison (1, NZAC); NN Golden Bay Rangihaeata, on Lophomytus 15.v.2001, N. Martin (1, NZAC); AK Handerson, on Acacia longifolia 10.xi.1976, J.S. Duodale (1, NZAC); ND. Painia, xii.1973, R.A. Cumba (1, NZAC); Auckland, xii.1974, N.A. Martin (1, NZAC).

Distribution Widely distributed in Australia, introduced and established in New Zealand (Martin 2016c) and Hawaii.

Harmonia decussata (Crotch, 1874) (Fig. 50)

Callineda decussata Crotch, 1874: 161; Type locality: not fixed. Korschefsky 1932: 282; Gordon 1987: 20 (lectotype). Harmonia flavomarginata Bielawski, 1968: 125. Type locality: Ambon. Synonymised by Poorani and Booth 2016: 9. Harmonia incognita Chazeau, 1990: 161. Type locality: Papua New Guinea, Port Moresby. Syn. nov. Harmonia manillana: Iablokoff-Khnzorian 1982: 486 (misidentification).

VII. Review of the Australo-Pacific Coccinellini

Diagnosis Harmonia decussata is a highly variable species and it is not always easily recognised externally. Male genitalia have to be consulted to confirm identification. Most of the reticulate forms of that species are distinguished from the similar H. nigromarginata or H. areolata in having at least part of the lateral elytral margin lightly coloured, and it differs from other similar species by having narrowly separated eyes and short prosternal carinae.

Description Colour pattern (Figs 50A–E). Head and appendages uniformly yellow or reddish. Scutellum black. Background colour of ventral surface brown. Legs yellow. Two main types of colour patterns on the pronotum and elytra exist: Typical form (Fig. 50C): pronotum yellow, with two black longitudinal stripes converging weakly anteriorly, the lateral and anterior borders transparent; elytra with a black suture and sometimes incomplete black reticulate pattern on a yellow or reddish background colour. Melanic forms (Fig. 50B): pronotum usually uniformly reddish or yellow, rarely with two indistinct black stripes; background colour of elytra black or dark brown, always with a reddish or yellow round spot apically, and one or two subquadrate reddish markings at the base. Morphology. Length 5.3–6.7 mm. Eyes partially beneath the pronotum. Frons between the eyes 1.1–1.5 times the eye width. Antenna 1.5 times as broad as the frons; antennal club compact; terminal antennomere strongly broadened. Pronotum with the anterior margin strongly arcuate medially; lateral margin arched weakly, slightly bent upwards; anterior angles of the pronotum broadly rounded. Prosternal process surface with parallel carinae at most slightly exceeding beyond the process. Scutellum ~1/12 times as broad as the elytra. Elytral epipleuron 2.0–2.5 times as broad as the metanepisternum. Abdominal postcoxal line reaching the posterior margin of ventrite 1, joined with a long oblique postcoxal line. Posterior margin of ventrite 5 truncate in males, slightly arcuate medially in females. Male terminalia (Figs 50H–J).

Distribution of Harmonia decussata

49

Female terminalia (Figs 50F,G). Spermatheca C-shaped, with the cornu pointed apically, situated close to nodulus; sperm duct short; infundibulum strongly sclerotised, flared at the junction of the bursa copulatrix. Measurements (n = 10): TL 5.30–6.70 (6.30 ± 0.50); PL 0.90–1.30 (1.09 ± 0.11); EL 4.70–5.50 (5.19 ± 0.28); PW 2.90– 3.20 (3.04 ± 0.08); EW 4.90–5.50 (5.22 ± 0.21); GD 2.60–3.10 (2.86 ± 0.15). Ratios: PL/PW 0.30–0.43 (0.36 ± 0.04); EL/EW 0.89–1.08 (1.00 ± 0.06); EL/PL 4.00–6.00 (4.81 ± 0.56); EL/ GD 1.66–2.00 (1.82 ± 0.10); TL/EW 1.00–1.37 (1.21 ± 0.11).

Types C. decussata: ‘Type/ decussata Wall./Type’ (UCCC, lectotype). H. flavomarginata: ‘Is. Ambon, INDONESIA, 28.x.1960, A. M. R. Wegner /alt. ca. 70m, at light / Holotypus/ Harmonia flavomarginata sp. n. det. R. Bielawski 1968’ (MIIZ, holotype). H. incognita: ‘Bred UPNG Waigani, Ot Moresby, T.P.N.G., Feb 1970, W. Ewers/ coll. no 57/ Harmonia incognita, n.sp. Holotype’ (BMNH, holotype).

Specimens examined INDONESIA: Ambon: Waai, 9.xi.1960, A.M.R. Wegner (4, BPBM); Is. Ambon, 7.xi.1960, A. M. R. Wegner, alt. ca. 70m, at light (4, MIIZ); Amboina, F. Muir (1, BPBM); Waai, 21.iii.1966(1, CAS). SERAM: Manusela N. P., 25.vii-9.ix.1987 (1, BMNH). Kai Islands: Kei Eil, 1922, H. C. Siebers (1, DEI). WESTERN NEW GUINEA: Ifar, Cyclops Mts., 300–500m, 28–30.vi.1962, J. Sedlacek (1. BPBM). PAPUA NEW GUINEA: Wau, Mt. Kaindi, v.1971, R. E. Parrott (1, CNC); Lae, light trap, 20.ix.1969 (1, ANIC); Misima Island, Umana Camp, 500ft, 6.xi-7.xii.1963, W. W. Brandt (1, ANIC); Misima L., H K. Bartlett, (5, MNHN); Morobe Dist., Lae, 13.ix.1964 (1, BMNH); Jungle Vege. Alt.225, 20.x.1944, T. Asrons (1, CAS); Finsch Haven dist. (1, SAM). Normanby Island, Culeguleu, 23–29.xii.1989, R. de Keyzer (1, AM). SOLOMON ISLANDS: Bougainville, 1956, J. L. Gressitt (1, BPBM); Guadalcanal Is., Lavoro Pltn., C.E. Hart (3, AM); Halmahera, Ganee, Doherty (1, MIIZ).

50

Ladybird Beetles of the Australo-Pacific Region

Distribution New Guinea, Ambon, Seram, Kai and Solomon Islands.

Remarks The type locality of this species is not fixed, because the lectotype specimen selected by Gordon (1987) does not have a locality label. It was described from a series collected in ‘Ceram, Amboina, Macassar, Kai, Bouru, Batchian, Gilolo (Wallace)’, but the type series is known to include several similar species. The current interpretation of this species agrees with the lectotype and further specimens compared with the lectotype by Roger Booth (BMNH).

Harmonia fijiensis (Sicard, 1929a) comb. nov.

those on the pronotum, with polished and shiny interspaces; epipleuron 2.5 times as wide as the metanepisternum. Mesoventrite with shallow anterior emargination. Tarsal claw with a long and pointed basal tooth. Abdominal postcoxal line (Fig. 51F) approaching the posterior margin of ventrite 1, connected to the faint, long oblique postcoxal line. Male. Unknown. Female terminalia (Figs 51D,E). Infundibulum strongly sclerotised and bent; spermatheca entirely sclerotised. Measurements (n = 3): TL 5.30–6.50 (5.77 ± 0.64); PL 1.00–1.10 (1.03 ± 0.06); EL 4.20–5.40 (4.70 ± 0.62); PW 2.20– 3.00 (2.47 ± 0.46); EW 4.40–5.30 (4.70 ± 0.52); GD 2.30–3.00 (2.53 ± 0.40). Ratios: PL/PW 0.37–0.45 (0.43 ± 0.05); EL/EW 0.95–1.02 (1.00 ± 0.04); EL/PL 4.20–4.91 (4.54 ± 0.36); EL/ GD 1.80–1.96 (1.86 ± 0.08); TL/EW 1.20–1.25 (1.23 ± 0.02).

(Fig. 51)

Types

Diagnosis

S. fijiensis: Fiji Islands, Suva, 7.vii.1927, C.294, H.W. Simmonds/ Synharmonia fijiensis n. sp. Sicard, types/ type/ Synharmonia fijiensis Sicard, lectotype female J. Chazeau det. 1976 (MNHN, lectotype here designated).

Synharmonia fijiensis Sicard, 1929a: 515. Type locality: Fiji. Coccinella fijiensis: Korschefsky 1932: 508.

This species is easily distinguished from other Harmonia species by its unique colour pattern (Figs 51A–C), and having the infundibulum strongly sclerotised and horn-like (Fig. 51E).

Description Colour pattern (Figs 51A–C). Frons, head appendages and legs yellow or orange; pronotum orange, with a medial black trapezoidal mark extending from the anterior to the posterior margin; elytra orange with a black stripe along the elytral suture and two sets of spots as in Fig. 51B; elytral epipleura orange; ventral side dark with paler prothoracic hypomera, mesepimeron and metepimeron. Morphology. Length 5.3–6.5 mm. Frons about twice as broad as the eye width. Pronotum weakly convex, with regular small punctures about as large as the eye facets; interspaces shiny. Prosternal process with admedian carinae distinct and extending forward to about the middle of the prosternum. Hypomeron weakly concave but without a cavity. Scutellum 1/12 the width of the elytra at the humeri, black and pointed apically. Elytral surfaces with fine punctures of similar size and distribution as

Distribution of Harmonia fijiensis

Remarks The taxonomic placement of this species, known from the female sex only, is uncertain and requires further research. Based on the structure of the female terminalia it does not belong to Syncharmonia (currently a junior synonym of Oenopia), having a strongly sclerotised and hooked infundibulum and simple spermatheca. It is here placed in Harmonia, based on the carinate prosternal process, not emarginate anterior margin of the mesoventrite, and the lack of the tibial spurs.

Specimens examined FIJI: Fiji 214, Fiji Islands. 1910–111 (1, BMNH); Fiji. C294, 8.vii.1927, H.W. Simmonds (1, BMNH); Fiji. C294, 11.vii.1927, H.W. Simmonds (1, BMNH); Lau Is., Lakeba I., S of Wagatalaca, 28.vi.1977, light, J.S. Dugdale (1, NZAC).

Distribution Endemic to Fiji.

VII. Review of the Australo-Pacific Coccinellini

51

Harmonia manillana (Mulsant, 1866) (Fig. 52)

Caria manillana Mulsant, 1866: 176. Type locality: Philippines. Gordon 1987: 17 (lectotype). Leis manillana: Crotch 1874: 120; Korschefsky 1932: 275; Bielawski 1962: 197. Harmonia manillana: Iablokoff-Khnzorian 1982: 486. Leis manillana var. incompleta Crotch 1974: 120. Leis manillana var. mniszechii Crotch 1974: 120. Caria atrocincta Mulsant, 1866:175. Type locality: Manilla, Luzon. Synonymised by Crotch 1874: 120. Leis manillana var. atrocincta Crotch 1874: 120. Neda paulinae Mulsant, 1866: 203. Type locality: Manilla, Luzon. Synonymised by Iablokoff-Khnzorian 1982: 486. Leis paulinae: Crotch 1874: 120; Bielawski 1962:197. Leis cerasicolor Crotch, 1874: 121. Type locality: Bourou. Korschefsky 1932: 273; Gordon 1987: 14 (lectotype). Synonymised by Iablokoff-Khnzorian 1982: 486. Leis aterrima Crotch, 1874: 121. Type locality: Ceram. Gordon 1987: 14 (lectotype). Synonymised by Iablokoff-Khnzorian 1982: 486. Leis papuensis Crotch, 1874: 121. Type locality: New Guinea. Korschefsky 1932: 275; Gordon 1987: 14 (lectotype). Synonymised by Iablokoff-Khnzorian 1982: 486. Leis dunlopi Crotch, 1874: 121. Type locality: India. Korschefsky 1932: 275; Gordon 1987: 14 (lectotype). Synonymised by Poorani and Booth 2016: 3. Harmonia dunlopi: Miyatake 1965: 60; Iablokoff-Khnzorian 1982: 486; Poorani 2002a: 330.

Diagnosis Harmonia manillana is easily recognised by its comparatively large size, strongly convex body and the elytral epipleura being at least 3.2 times broader than the metanepisternum. Some colour forms of this species can be mistaken for Docimocaria, but they differ from members of that genus in having an almost straight anterior margin of the mesoventrite and no apical spurs on the mid and hind tibiae.

Description Colour pattern (Figs 52A–L). Dorsal colour pattern variable, but the elytra mostly with black spots or transverse bands on a yellowish or reddish background. Head reddish, sometimes black at the base. Pronotum colouration varies from complete yellow-reddish, to complete black except for a pale narrow anterior angle area. Scutellum always black. Morphology. Length 7.8–9.8 mm. Frons 2.5–3.0 as wide as the eye width. Antenna as long as the frons width. Pronotum much narrower than the elytra at the base; prosternal process with strong, anteriorly converging admedian carinae that extend forward to the middle of the prosternum. Mesoventrite shallowly arcuate anteriorly. Scutellum 1/12 times as broad as the elytra at the shoulder area. Metaventral postcoxal lines strongly arcuate before reaching the metanepisternum. Elytral epipleuron ~3.2–3.6 times as broad as the metanepisternum, yellow-reddish with black external border. Abdominal postcoxal line (Fig. 52S) almost reaches the posterior margin of ventrite 1, connected to the oblique postcoxal line. Posterior margin of ventrite 5 truncate in males, and arcuate medially in females. Male terminalia (Figs 52M–P). Female terminalia (Figs 52Q,R).

Distribution of Harmonia manillana

Measurements (n = 8): TL 7.80–9.80 (8.82 ± 0.71); PL 1.00–1.70 (1.28 ± 0.19); EL 6.60–8.60 (7.47 ± 0.67); PW 3.90– 4.60 (4.24 ± 0.22); EW 7.30–9.60 (8.53 ± 0.83); GD 3.90–4.75 (4.31 ± 0.30). Ratios: PL/PW 0.25–0.37 (0.30 ± 0.03); EL/EW 0.73–0.93 (0.88 ± 0.06); EL/PL 4.76–6.90 (5.93 ± 0.71); EL/ GD 1.59–1.85 (1.73 ± 0.07); TL/EW 0.95–1.08 (1.04 ± 0.04).

Remarks Harmonia flavomaculata was treated as a synonym of H. manillana by Iablokoff-Khnzorian (1982), but is here recognised as a synonym of Harmonia decussata, in agreement with Poorani and Booth (2016).

Types L. manillana: Type/ Type manillana Deyr. (UCCC, lectotype). L. dunlopi: Type/Type Dunlopi India (UCCC, lectotype). L. cerasicolor: Type/ Type cerasicolor Bourou (UCCC, lectotype). L. aterrima: Type/ aterrima Ceram (UCCC, lectotype). L. papuensis: Type/ Type papuensis N. Guin (UCCC, lectotype).

Specimens examined INDONESIA: Western New Guinea: Fak-fak Dutch New Guinea (1, BMNH). Ambon: Waai, 21.iii.1966, A.M.R. Wegner (1, CAS); PAPUA NEW GUINEA: Bosavi, x.1976, H. Ohlmus (4, ANIC); Wau, Mt. Kaindi, 4.v.1971, R.E. Parrott (1, CNC); Stephansort, Astrolabe, 1884, Kimzmann (1, MNHN); Dogura, 27.x.1959, E.L. Cassidy (1, MNHN); New Guinea, 88.13 (1, BMNH); 79.14; Pt. Moresby (1, BMNH).

Distribution This is a widely distributed and morphologically variable species, known from a very broad geographic range extending from India (Arunachal Pradesh) and Malaysia to New Guinea (Poorani and Booth 2016).

Harmonia micra sp. nov. (Fig. 53) Diagnosis This species is easily recognised by its small size and elytra black with two yellow spots on each elytron.

Description Colour pattern (Figs 53A–C). Head uniformly yellow. Pronotum orange, with black area medially. Scutellum black.

52

Ladybird Beetles of the Australo-Pacific Region

Distribution of Harmonia micra

Elytra black with two orange spots on each elytron. Venter completely yellow in females, in males largely yellow, except for the metaventrite, the external border of the epipleura and the medial part of abdominal ventrites 1–3 are all dark. Legs yellow. Morphology. Length 4.4–4.8 mm. Frons ~1.5 times as broad as the eye width. Eyes partially exposed. Pronotum ~0.6 times as broad as the elytra, with the lateral margin evenly arcuate. Prosternal process flat, surface with short indistinct carinae not extending beyond the process. Scutellum ~1/10 times as broad as the elytra. Elytra strongly convex, narrowly explanate laterally. Anterior margin of the mesoventrite almost straight. Abdominal postcoxal line separated medially, connected to long oblique (Fig. 53I) postcoxal line. Posterior margin of ventrite 5 emarginate medially in males, slightly arcuate in females. Male terminalia (Figs 53D–G). Female terminalia (Fig. 53H). Measurements (n = 5): TL 4.40–4.80 (4.56 ± 0.15); PL 0.60–0.70 (0.66 ± 0.05); EL 3.70–4.20 (3.90 ± 0.19); PW 2.30– 2.50 (2.40 ± 0.10); EW 3.50–3.90 (3.74 ± 0.15); GD 2.10–2.10 (2.10 ± 0.00). Ratios: PL/PW 0.24–0.30 (0.28 ± 0.03); EL/EW 1.00–1.11 (1.04 ± 0.04); EL/PL 5.29–7.00 (5.96 ± 0.75); EL/ GD 1.76–2.00 (1.86 ± 0.09); TL/EW 1.18–1.26 (1.22 ± 0.04).

Specimens examined Holotype: SOLOMON ISLANDS: Bougainville, Bush E. of Buin, 7.i.1970, James E. Tobler (CAS). Paratypes: same data as the holotype (3, CAS; 1, ANIC).

Distribution Endemic to Bougainville, Solomon Islands.

Remarks The species name, micra, refers to a small size of the beetle.

Harmonia nigromarginata Bielawski, 1968 (Fig. 54)

Harmonia nigromarginata Bielawski in Bielawski and Chûjô, 1968: 127; Type locality: Ambon. Iablokoff-Khnzorian 1982: 487. Harmonia rhopalocera Iablokoff-Khnzorian, 1982: 475. Syn. nov.

Diagnosis Harmonia nigromarginata and typical forms of H. decussata share similar body size, a broadly oval outline, and a black dorsal colour pattern on a yellowish background. Harmonia nigromarginata can be distinguished from most of colour forms of H. decussata in having a very narrow black suture and lateral margins, in addition to the incomplete black reticulate colour pattern on the elytra.

Description Colour pattern (Figs 54A–E). Head and appendages yellow. Pronotum yellow, with two black longitudinal fasciae converging weakly anteriorly, the lateral and anterior borders transparent. Scutellum black or dark brown. Elytra yellow or brownish, with a narrow and usually incomplete black reticulate pattern; the external borders and suture are dark. Background colour of the ventral surface brown. Legs yellow. Morphology. Length 5.2–6.3 mm. Frons 1.1–1.3 as wide as the eye width; surface concave centrally. Antenna 1.3–1.5 times as long as the frons width; antennal club compact; the terminal antennomere strongly broadened, almost as large as the terminal maxillary palpomere, rounded at one side, slightly angulated on the other. Anterior pronotal margin slightly arcuate medially; lateral margin upturned; the anterior angles of the pronotum broadly rounded. Prosternal process with short parallel carinae not extending beyond the process. Scutellum at base 1/12 times as broad as the elytra. Elytra narrowly explanate laterally; epipleuron ~2.2–2.6 times as broad as the metanepisternum; Mesoventrite with the anterior margin slightly emarginate medially. Abdominal postcoxal line approaches the posterior margin of ventrite 1, connected to a long oblique postcoxal line (Fig. 54L). Ventrite 5 with a posterior margin broadly emarginate medially in males, truncate in females. Male terminalia (Figs 54H–K). Female terminalia (Figs 54F,G). Measurements (n = 6): TL 5.20–6.30 (5.78 ± 0.48); PL 0.95–1.10 (1.01 ± 0.06); EL 4.20–5.20 (4.68 ± 0.43); PW 2.60– 2.90 (2.69 ± 0.14); EW 4.30–4.90 (4.53 ± 0.26); GD 2.35–2.70 (2.46 ± 0.17). Ratios: PL/PW 0.34–0.42 (0.38 ± o.03); EL/EW 0.95–1.07 (1.03 ± 0.05); EL/PL 4.36–5.20 (4.62 ± 0.39); EL/ GD 1.79–1.96 (1.90 ± 0.08); TL/EW 1.18–1.33 (1.28 ± 0.07).

VII. Review of the Australo-Pacific Coccinellini

53

Distribution of Harmonia nigromarginata

Types H. nigromarginata: Is. Ambon, INDONESIA, 10.ix.1960, A.M.R. Wegner/ Alt. ca. 70m, at light/ Harmonia nigromarginata sp.n. det. R. Bielawski, 1968 (ELKU, holotype; MIIZ, paratypes)

Specimens examined INDONESIA: Ambon: Waai, 21.iii.1966, A. W. R. Wegner (1, CAS); Ambon, 20.viii.1960, 2.xi.1960, 10.ix.1960, A.M.R. Wegner, Alt. ca. 70m, at light (3, MIIZ, paratypes). Seram: Manusela N.P., Wae Mual Plain, Lowland Alluvial forest, 25.vii-9.ix.1987 (1, BMNH).

Distribution Known from Ambon, Seram and Sulawesi, Indonesia.

Remarks Among the specimens from the Natural History Museum in Berlin we have found a male of H. niromarginata bearing a label ‘Celebes, Bantimurang, 1882, C. Ribbe’. The locality label and the specimen markings match the locality and description given for Harmonia rhopalocera Iablokoff-Khnzorian, 1982: 475. The holotype of H. rhopalocera has unfortunately not been found but there is no doubt about the proposed synonymy.

Harmonia novaehebridensis (Korschefsky, 1943) (Fig. 55)

Coelophora novaehebridensis Korschefsky, 1943: 51. Type locality: Vanuatu: Ile Pentecôte. Harmonia novaehebridensis: Chazeau 1978: 66 (lectotype).

Diagnosis Harmonia novaehebridensis is easily distinguished from other Harmonia species by its unique elytral colour pattern (Fig. 55A). However, it can easily be confused with the sympatric Microcaria vivida (Fig. 97A), in having similar body size, outline, and colour pattern. Harmonia novaehebridensis can be distinguished from Microcaria vivida in having the terminal

antennomere strongly expanded and securiform, and the mesoventrite almost straight medially.

Description Colour pattern (Figs 55A–C). Head, pronotum and appendages light reddish to light brown. Scutellum reddish centrally with black borders. Elytra reddish with three transverse broad bands; suture reddish in male and narrowly black in female. Ventral side light coloured except for darker prosternum, meso- and metaventrite and central part of abdominal ventrites. Morphology. Length 5.4 mm. Frons between eyes ~1.5–1.6 times as broad as the eye width. Eyes partly beneath theanterior margin of pronotum, interior orbits parallel. Pronotum 0.6 times as broad as the elytra; lateral margin weakly arcuate. Prosternal process with indistinct admedian carinae. Scutellum ~1/12 times as broad as the elytra. Elytra narrowly explanate laterally; epipleuron 2.0–2.2 times as wide as the metanepisternum. Anterior margin of mesoventrite slightly emarginate in the middle. Abdominal postcoxal lines separated medially, laterally joined to the oblique postcoxal line (Fig. 55J). Posterior margin of ventrite 5 truncate in males and arcuate in females. Male terminalia (Figs 55D–G). Female terminalia (Figs 55H,I). Spermatheca strongly curved with the cornu tapered apically; nodulus slim, ramus indistinct; infundibulum slim, flared at both ends. Measurements (n = 2): TL 5.3–5.4; PL 0.7–0.9; EL 4.5–4.6; PW 2.7–2.9; EW 4.3–4.4; GD 2.3–2.4. Ratios: PL/PW 0.26– 0.31; EL/EW 1.02–1.07; EL/PL 5.00–6.57; EL/GD 1.92–1.96; TL/EW 1.23–1.23.

Types C. novaehebridensis: ‘Nouvelles-Hébrides, Ile Pentecôte, Baie Melsisi, 1935–1936, E. Aubert de la Rue’ (MNHN, lectotype).

Specimens examined VANUATU: Tagobe Efate I., agriculture station, 14.ii.1986, P. A. Maddison (2, NZAC).

Distribution Known only from Vanuatu.

54

Ladybird Beetles of the Australo-Pacific Region

Distribution of Harmonia novaehebridensis

Harmonia octomaculata (Fabricius, 1781) (Fig. 56)

Coccinella octomaculata Fabricius, 1781: 97. Type locality: not specified. Harmonia arcuata var. octomaculata: Mulsant 1850: 80; Korschefsky 1932: 441. Harmonia octomaculata: Bielawski 1957: 93, 1962: 204, 1964b: 5; Sasaji 1971: 280; Iablokoff-Khnzorian 1979: 71, 1982: 475; Pope 1989: 685; Ślipiński 2007: 163. Ptychanatis octomaculata: Kamiya 1965b: 59. Coccinella arcuata Fabricius, 1787: 55. Type locality: China. Boisduval 1835: 591; Crotch 1874: 110; Froggatt 1902: 900; Lea 1902: 488; Korschefsky 1932: 440. Harmonia arcuata: Mulsant 1850: 77, 1866: 59; Crotch 1971: 3; Timberlake 1943:18; Miyatake 1965: 60.

Diagnosis Harmonia octomaculata is easily distinguished from other species in the area studied by having a relatively flat, elongateoval body, and the lateral margins of the elytra form a distinct gutter along the humeral angle.

Description Colour pattern (Figs 56A–E). Frons and head appendages yellow or reddish. Pronotum yellow or red, usually with four closely set black spots, sometimes forming a larger black central area. Scutellum darker than background colour of the elytra. Elytra dark yellow, usually with eight black spots on each elytron, the apical spot often missing, or with spots enlarged and connected (Figs 56A–D); the lateral borders yellow and the suture black. Ventral side yellow, except for darker metaventrite mesanepisternum, metanepisternum and central part of the abdomen. Legs mostly yellow, the hind femur partially dark in most of the specimens examined. Morphology. Length 5–8 mm. Frons ~1.7–2.0 times as broad as the eye diameter. Antenna much longer than the frons

width; club compact, with the terminal antennomere strongly broadened. Pronotum ~0.9 times as broad as the elytra, with the anterior margin straight, the lateral margin slightly arcuate. Surface of the prosternum with short carinae, ending well before middle of the prosternum. Elytra as long as broad, with the external borders narrowly explanate, and forming a distinct gutter around the humeral angle. Elytral epipleuron ~1.5 times as broad as the metanepisternum. Scutellum small, 1/15 times as broad as the elytra. Anterior margin of the mesoventrite almost straight, with light emargination in the middle, the posterior margin straight. Tarsal claws long. Abdominal postcoxal line short, approaching but not reaching the posterior margin of ventrite 1, connected with a short oblique postcoxal line (Fig. 56L). Apical border of ventrite 5 truncate in males and slightly arcuate medially in females, ventrite 6 distinctly emarginate in males, and arcuate with a medial tubercle in females. Male terminalia (Figs 56F–I). Female terminalia (Figs 52J,K). Spermatheca strongly C-shaped; the infundibulum slim, flared apically; the bursa copulatrix sclerotised apically. Measurements (n = 8): TL 5.10–8.00 (6.58 ± 0.68); PL 0.90–1.35 (1.02 ± 0.12); EL 4.10–6.30 (5.50 ± 0.59); PW 2.30– 3.30 (2.92 ± 0.23); EW 3.80–5.70 (4.98 ± 0.49); GD 2.10–3.00 (2.72 ± 0.24). Ratios: PL/PW 0.30–0.41 (0.35 ± 0.03); EL/EW 1.04–1.24 (1.10 ± 0.04); EL/PL 4.56–6.56 (5.43 ± 0.58); EL/ GD 1.79–2.21 (2.02 ± 0.10); TL/EW 1.23–1.46 (1.32 ± 0.06).

Specimens examined INDONESIA: Western New Guinea: Jayawijaya. Wamena, Baiemflub, 1700m, 15.x.1993, leg. A. Riedel (2, NHMS); Merauke Senggo, trail to Abau ca. 100m, 15–17.vi.1994, A. Riedel (1, NHMS); Arfak, 8.ix.1912, P. Kibler (4, NHMS); Maffin Bay, 10.viii.1944, E. S. Ross (5, CAS); Biak I., xi.1945, H. Blakenore (1, CAS); Biak Island, 20–23.i.1945, H. H. Blackmore (1, CAS). PAPUA NEW GUINEA: Bulolo, 30.v.1971, R.E. Parrott (1, ANIC); Lae, 12.x.1970, R.E. Parrott (1, ANIC); Western

VII. Review of the Australo-Pacific Coccinellini

55

Distribution of Harmonia octomaculata

Dist., Rouku, Morehead River, 19.iii-28.v.1962, W.W. Brandt (1, ANIC); Morehead, 31.v.1995, J.M. Grimshaw (1, NAQS); NE Angoram, 10m, 13.viii.1969, J.L. Gressitt (1, BPBM); Wau, Morobe Distr., 1700–1800m, 17.xi.1961, J.L. Gressitt (1, BPBM); (NE) Wau, Morobe Distr., 1000–1300m, 20.iv.1962, J. Sedlacek (2, BPBM); NE Iongai, 10km E of Mt. Albert Edward, 1450–1800m, 8–10.xi.1965, J. Sedlacek (1, BPBM); SE Woltape, 1550–1750m, 2–3.xi.1965, J. Sedlacek (1, BPBM); SOLOMON ISLANDS: Guadalcanal, Tenaru R., i.1945, G. E. Bohart (2, CAS); Guadalcanal, Honiara, 0–100m, ii.1985, N. L. Krauss (3, CAS); Guadalcanal & Florida IS, i-iii.1945, J. R. Stuntz (3, CAS); Kolombangara Isl., Ringi Cove road to Airport, R2, 18.i.1970, J. E. Tobler (1, CAS); COOK ISLANDS: Rarotonga, Taro swamp/tip nr. airport, 9–10.jun.1988, M. Lenz (3, ANIC); NEW CALEDONIA: (S) 21.37S/ 165.49E 450m, Col d’Amieu (Top of hill), light, 9.ii.2004, M. Wanat (1, AINC); (S) 22.06S/166.39E 180m Riviere Bleue P. (N) 1km of vic. of scientists refuge, 27.i.2004, M. Wanat (1, ANIC); (N) 20.45S/164.53E, Tiendanite, 50m, Solanaceae shrubs, 4.ii.2004, M. Wanat (2, AINC); Noumea, 0–100m, xii.1984, N.L. Krauss (1, CAS); Loyalty Islands: We (Lifen), 11.x.1955, J. Rageau (1, CAS). AUSTRALIA: Queensland: Hammond Island, Torres Strait; Thursday I.; Lockerbie; Bamaga; Weipa; Iron Range; Claudie R.; Coen; Silver Plains HS.; Stewart R., Musgrave; Cooktown; Normanby R.; Laura; Kowanyama; Mornington I.; Mt Molloy; Kuranda; Redlynch; Cairns; Mareeba; Chillagoe; Danbulla Forestry Reserve; Tolga; Atherton; 27–32 km W. of Atherton; Johnstone; Mt Garnet; Forty Mile Scrub; Silkwood; 8 km E. of Cardstone; Cardstone; 32 km S. of Ravenshoe; Doomadgee Mission; Ingham; Townsville; Ayr; Lansdowne Stn.; Port Denison; Mackay; Gannet Cay; Rockhampton; Gladstone; Moura; Bundaberg; Gayndah; Cooloola; Kingaroy; Mapleton; Brisbane; Moreton Bay. Northern Territory: Coopers Ck.; Darwin; Cannon; Cahills Crossing; Berrimah; Humpty Doo; Batchelor; Stapleton; Adelaide R.; Katherine; Delamere; Batten; Jasper. Western Australia: Prince Regent Reserve; Carson Escarpment; Kimberley Res.; Wyndham; Kununurra; West Kimberley; Derby; Broome; Perth.

Distribution Indonesia, Papua New Guinea, Australia, New Caledonia, Solomon Islands, Philippines, Fiji, Tonga, Malaysia, Laos, North Vietnam, China, India, Cook Islands.

Harmonia ohlmusi sp. nov. (Fig. 57) Diagnosis This species is easily recognised by its large size and black elytra, each bearing two large orange spots (Fig. 57A).

Description Colour pattern (Figs 57A–D). Frons usually brown, sometimes with dark brown lateral and anterior borders, rarely entirely black. Labrum usually dark. Pronotum black, with two large light yellow spots laterally; lateral borders partially black. Scutellum black. Elytra entirely black, or with two orange or reddish spots on each elytron. Venter black, except for the pale prothoracic hypomeron and mesepimeron. Legs usually black. Morphology. Length 6–9 mm. Frons about twice as broad as the eye width. Antenna slightly longer than the frons width. Anterior margin of the pronotum straight medially; the lateral margin evenly arcuate. Prosternal process with carinae extending to the middle of the prosternum. Scutellum 1/13 times as broad as the elytra. Lateral border thickened, and very narrowly explanate. Anterior margin of the mesoventrite weakly emarginate medially, posterior margin straight. Elytral epipleuron ~2.5 times as broad as the metanepisternum, strongly descended. Abdominal postcoxal line approaching the posterior margin of ventrite 1, joined apically with a very fine oblique postcoxal line. Posterior margin of ventrite 5 slightly emarginate medially in males and arcuate in females. Male terminalia (Figs 57E–J). Female terminalia (Fig. 57K). Spermatheca strongly C-shaped; cornu rounded apically and broadened at the end close to the nodulus; nodulus slim; infundibulum slim, not flared.

56

Ladybird Beetles of the Australo-Pacific Region

Measurements (n = 4): TL 6.30–9.30 (7.57 ± 0.68); PL 0.80–1.30 (1.07 ± 0.14); EL 5.50–7.90 (6.41 ± 0.59); PW 3.30– 4.30 (3.66 ± 0.26); EW 5.10–7.50 (6.40 ± 0.57); GD 2.90–4.10 (3.44 ± 0.33). Ratios: PL/PW 0.24–0.35 (0.29 ± 0.04); EL/EW 0.90–1.14 (1.00 ± 0.06); EL/PL 4.69–7.89(6.08 ± 0.85); EL/ GD 1.68–2.13 (1.87 ± 0.10); TL/EW 1.07–1.33 (1.18 ± 0.07).

Specimens examined Holotype male: PAPUA NEW GUINEA: 6.05S-145.25E, Goroka, viii.1976, H. V. Ohlmus Coll. (ANIC). Paratypes: PAPUA NEW GUINEA: New Guinea NE Hagen town, 4’48’S-14’17’E, 1650m, 30.v.1966, J. L. Gressitt (BPBM); NE Wau, Edie Creek, 1891m, 8.viii.1963(1, BPBM); New Guinea: (NW) Wisselmeren Enarotadi, 1800–1900m, 22.viii,1962, J. Sedlacek (1, BPBM). Other material examined: PAPUA NEW GUINEA: NE. Wau,1200m, M.V. light trap, 4–13.vii.1965, J. & M. Sedlacek (1, BPBM); SE Koroba, 40km W of Tari, 1650m, light trap, 18.xi.1963, R. Staatman (1, BPBM); (NW) Wissel Lakes Enarotadi, 1900–2000m, 2–11.vii,1962, J. Sedlacek (1, BPBM); New Hagen town, 4.48S-144.17E, 1650m, 30.v.1966, J. L. Gressitt (1, BPBM); NE Hagen town 4’43’s,144.17’E, 1650m, 30.v.1966, J. L. Gressitt (2, BPBM); NE Wau, 2400m, 9–12.i.1962, J. H. & M. Sedlacek, G. Monteith & Native Collectors (1, BPBM); (NW) Wisselmeren, Itouda Kamo V., 1500–1700m, 18.vii.1962, J. Sedlacek (1, BPBM); NE: Morobe Dist: Wau, light trap, 27.iv.1982, R.T. Bell (1, BPBM); Upper Manki L. A. Bulolo, M. Dist. In sticky trap, 23. vi.1972, F. R. Wylie & R. Shanahan (1, ANIC); SE Popondetta Dist., Jumbora, 23.x.1963, P. Shanahan (1, BPBM); Kmu, vii.1975, H. Ohlmus Collector(2, ANIC); Morobe Distr.(E) Mt Kainai, nr Wau, 2350m, 3.ix.1973(1, BPBM); Papua New Guinea, Morobe District Aseki 1200m, 12.iv.1974, A. D. Hart (1, BPBM); Papua New Guinea, Morobe Distrct, Watut-Aseki Div. 2800m, 10.iv.1974, J. L. Gressitt (1, BPBM); Upper Manki L. A. Bulolo, M. Dist. In sticky trap, 29. ix.1972, F. R. Wylie & R. Shanahan (1, ANIC); Papua New Guinea, Morobe District Aseki 1200m, 14.iv.1974, Reni Sakomdaro (1, BPBM); (NE) Mt. Kaindi, 16km SW of Wau, 2300m, 6.x.1962, light trap, J. Sedlacek (1, BPBM); NE Mt Kaidi, 2350m, light trap, 25.xi.1971, TAWI (1, BPBM); New Guinea, Friaano, v.1975, H. Ohlmus (1, ANIC); Upper Manki L. A. Bulolo, M. Dist. In sticky trap, 28. vii.1972, F. R. Wylie & R. Shanahan (1, ANIC); NE Wau, 1200–1500m, M. V. light trap, J. & M. Sedlacek, 30.ix.1965(2, BPBM); Morabi Dist. Wau. 6.x.1960, James E. Tobler (1,CASC); (NE), Wau, Morobe Dist. Mt. Missim, 1100m, 16.ii.1963, J. Sedlacek (1, BPBM); Miwaka, viii.1975, H. Ohlmus (1, ANIC); New Guinea: Tiffalmin, ii.1970, H. Ohlmus (1, ANIC); NE Wau, Bulolo R., 900–1100m, 25.ix.1965, J. Sedlacek(1, BPBM). Baliem Tal, 1700m, iii.1992, Jiri Kolibac (1, BA-0112); NE Wau, Bulolu R., 900–1100m, 25.ix.1965, J. Sedlacek (1, BPBM); NE Hagen town, 4.48’s-144.17E, 1650m, 30.v.1966, (2, J. L. Gressitt); NE Wau, 1200m, Light trap, 26–27.ix.1964, J. Sedlacek (1, BPBM); NE Morobe District, Mindik, 1200–1600m, ix.1968, N. L. H. Krauss (1, BPBM); NE Okapa, 1900m, 24–25.vi.1967, G. A. Samuelson (1, BPBM); NE Mt. Kaindi, 2350m, light trap,25. xi.1971/ K-447 (1, BPBM); Baiyer R., N. Guinea,1100m, 26.xii.1978–25.i.1979, J. Sedlacek (1, BMNH); PAPUA: Mondo, 5000 ft, i-ii.1934, L. E. Chessman (1, BMNH); Goroka East, Highlands, New Guinea, 9.x.1956(1, BMNH); Exgorona, 17.ix.19576, N. Aroley (1, BMNH); Banz, West Highl., Gorden of R. C. Mission, on grape-fruit tree infested with Icerya sp., 20.x.1954, Szent-Ivany (1, BMNH); Tanuri Pyn., Sepik Distr., on Cacao 14.x.1957, J.J.H. Szent-Ivany (1, BMNH).

Distribution of Harmonia ohlmusi

INDONESIA: Western New Guinea: Irian Jaya: Biak-Is. Suneri, 4.x.1990, leg. A. Riedel (1, NHMS).

Distribution Endemic to the island of New Guinea.

Remarks Named after Mr Henry V. Ohlmus, amateur collector of insects in Papua New Guinea, who has collected many valuable ladybirds, including the holotype of this species, and deposited them in the Australian National Insect Collection.

Harmonia tricolor (Korschefsky, 1944) comb. nov. (Fig. 58)

Coelophora tricolor Korschefsky, 1944: 56. Type locality: Bougainville, Kieta.

Diagnosis Harmonia tricolor is very similar to H. tobleri, occurring on the same islands of the Solomon Islands, but it can be distinguished by its entirely orange elytral epipleuron and its distinctive male genitalia.

Description Colour pattern. Frons and head appendages dark yellow. Labrum yellow. Pronotum light yellow, with central part largely black, the lateral borders black. Scutellum black. Elytra red or yellow, with the shoulder area and large central area black, the external borders and suture yellow, except for an area connected with black spots. Background colour on the ventral side black, except for the yellow prothoracic hypomeron, elytral epipleuron and mesepimeron. Legs yellow. Morphology (Figs 58A–C). Length 5.7–7.8 mm. Frons about twice as broad as the eye width. Antenna slightly longer than the frons width; terminal antennomere slightly broadened. Pronotum weakly convex, the anterior margin straight medially; the lateral margin evenly arcuate. Prosternal process with carinae extending forward to beyond the middle of the prosternum, convergent anteriorly. Elytra almost as long as broad, comparably broadly explanate, the external borders with large and dense punctures. Anterior margin of mesoventrite almost straight medially, posterior margin straight. Scutellum ~1/15 times as broad as the elytra. Elytral epipleuron about three times as broad as the metanepisternum, strongly descending. Abdominal postcoxal line approaching the posterior margin of ventrite 1 and extending to the lateral margin; the oblique postcoxal line not connected to postcoxal

VII. Review of the Australo-Pacific Coccinellini

57

Distribution of Harmonia tricolor

line (Fig. 58J). Posterior margin ventrite 5 broadly emarginate medially in males and slightly arcuate in females. Male terminalia (Figs 58D–H). Female terminalia (Fig. 58I). Spermatheca strongly bent; cornu pointed apically and broadened at the end closed to the nodulus; nodulus slim, ramus absent; infundibulum slim, not flared. Measurements (n = 5): TL 5.70–7.80 (7.04 ± 0.58); PL 0.80–1.20 (1.04 ± 0.13); EL 4.50–6.50 (5.95 ± 0.58); PW 2.80– 3.50 (3.24 ± 0.22); EW 4.90–6.80 (6.12 ± 0.56); GD 2.50–3.50 (3.17 ± 0.32). Ratios: PL/PW 0.27–0.39 (0.32 ± 0.04); EL/EW 0.92–1.02 (0.97 ± 0.04); EL/PL 4.09–7.13 (5.80 ± 0.87); EL/ GD 1.71–2.07 (1.88 ± 0.11); TL/EW 1.10–1.20 (1.15 ± 0.04).

Specimens examined SOLOMON ISLANDS: ‘Southern coast of Kolombangara Is. New Georgia Group Solomon Islands, i-x.1990, Fairlamb Family’ (ANIC). Kieta District, Buka, Hanahan, 24.xii.1969 (7, CAS); Bougainville Island, Konga Village (Buin), 21.iii-2.vi.1961, W. W. Brandt (1, ANIC); New Georgia I: Munda, 0–100m, xi.1980, N. L. H. Krauss (3, BPBM); New Georgia I: Munda, 0–150m, xi.1976, N. L. H. Krauss (2, BPBM); Kolombangara Isl., Ringi Cove, Main Road near R6 Road, 17.i.1970, James E. Tobler (53, CAS; 2, SYSBM); Kolombangara Isl., Ringi Cove, Main Road near R6 Road, 16.i.1970, James E. Tobler (38, CAS); Kolombangara Isl., Ringi Cove, Road to Airport, R2, 18.i.1970, J. E. Tobler (5, CAS); Guadalcanal, Sol. Is., 1920, J. A. Kusche (1, BPBM).

Distribution Endemic to the Solomon Islands.

Harmonia testudinaria (Mulsant, 1850) (Fig. 59)

Daulis testudinaria Mulsant, 1850: 300. Type locality: Australia. Neda testudinaria: Masters 1888: 90; Lea 1902: 489. Harmonia testudinaria: Timberlake 1943: 18; Bielawski 1964b: 7; Iablokoff-Khnzorian 1979: 72, 1982: 494; Pope 1989: 687 (lectotype); Ślipiński 2007: 164.

Diagnosis In Australia, H. testudinaria is easily recognised by its light yellow or orange elytra, with a more or less complete black reticulate colour pattern (Figs 59A–C). In the Papuan region, several species share a similar colour pattern, and the distinguishing characters of H. testudinaria include having long prosternal carinae, broad black sutural and lateral margins, and distinctive male genitalia.

Description Colour pattern (Figs 59A–D). Head and appendages yellow. Pronotum yellow, with two longitudinal stripes not reaching the anterior margin (Fig. 59D); the lateral and anterior borders are transparent. Scutellum black. Elytra usually yellow, with a variously developed black reticulate pattern; pale forms with elytra uniformly yellow or orange, with black sutural and lateral margins and a short humeral stripe (Fig. 59B). Prothoracic hypomeron yellow. Prosternum yellow, with dark margins in males, and black in females. Pterothorax usually black or dark brown ventrally except for the pale mesepimeron. Elytral epipleuron yellow, with black external margins. Abdomen black or dark brown. Legs usually dark. Morphology. Length 5.3–8.3 mm. Frons between the eyes ~1.5 times as broad as the eye diameter. Antenna slightly longer than the frons width; the terminal antennomere securiform, slightly angulated. Pronotum ~0.8 times as broad as the elytra; the anterior margin slightly arcuate medially, the lateral margin evenly arcuate. Prosternum with carinae extending to the middle of the prosternum, converging anteriorly. Scutellum ~1/12 times as broad as the elytra. Elytra narrowly explanate laterally; epipleuron about twice as broad as the metanepisternum. Abdominal postcoxal line approaches the posterior margin of ventrite 1; the oblique postcoxal line fine and not connected to postcoxal line. Posterior margin of ventrite 5 strongly emarginate medially in males and broadly arcuate in females. Male terminalia (Figs 59E–H). Female terminalia (Figs 59I,J). Spermatheca strongly curved; the infundibulum slim, not flared; the bursa copulatrix lightly sclerotised apically. Measurements (n = 10): TL 5.30–8.30 (6.93 ± 0.81); PL 0.90–1.60 (1.11 ± 0.17); EL 4.30–6.80 (5.69 ± 0.62); PW

58

Ladybird Beetles of the Australo-Pacific Region

2.50–3.70 (3.18 ± 0.27); EW 4.30–6.60 (5.58 ± 0.56); GD 2.30–3.90 (2.98 ± 0.38). Ratios: PL/PW 0.28–0.46 (0.35 ± 0.04); EL/EW 0.93–1.08 (1.02 ± 0.03); EL/PL 3.94–6.22 (5.19 ± 0.46); EL/GD 1.65–2.10 (1.91 ± 0.11); TL/EW 1.10–1.38 (1.24 ± 0.07).

Types D. testudinaria: Museum Paris Nov. Hollande Gory 229–35/91/ Daulis testudinaria Muls. auct. det.’ (MNHN, lectotype).

Specimens examined INDONESIA: Western New Guinea: Kecamatan Abenaho Pass Valley, 3.31S-139.05E, 1700–2250m, Dist. Mtn. rainforest, 18–25.ii.2005, T. Lackner (1, NBCL); Jawawijaya. Wamena. Baliemflub, 1700m, 15.x.1993, A. Riedel (1,NHMS); Iba 1300m, 7–8.iv.1993, A. Riedel (1,NHMS); Jayawijawa, N. Bime 2000–2070m, 21.ix.1993, A. Riedel (1, NHMS); Anggi, Tetaho Kosmena 1400–1750m 26–28.iii.1993, A. Riedel (1, NHMS); Prov. Jayawijaya. Bommela, 1750m, 30.viii-1. ix.1992, A. Riedel, (1, NHMS); Jayawijaya, Membahan, 23.ix.1991, A. Riedel (1, NHMS); Japen Seruin, Ambeidiru, 1000m, 5.viii.1996, Schule & Stuben (1, NHMS); Manokwari Pr., Ca. 10km, NW Ransiki, kali way 1300m, 01.25’03’’S 134.01’49’’E, 03.iii.1927, A. Weigel (1, NKME). PAPUA NEW GUINEA: Cromwell Mts, 1929, O. Bryant (6, CAS); Owgarra, Meek, 1952, R. Oberthur (1, MNHN); Prov. Morobe. Aseki, 1000–1300m, 13.x.1992, A. Riedel (5, NHMS); Wau, Mt. Kaindi, 1650m, 7.x.1992, A. Riedel, (2, NHMS); Wau, Mt. Kaindi, 1550–2150m, 7–8.x.1992, A. Riedel, (1, NHMS); Upper Manki, Bulolo, M. Dist., 4.viii.1972, F.R. Wylie & R. Shanahan (2, ANIC); Biarum MP, 1800m, vi.1998 (1, ANIC); Morobe Prov., Garaina, 700m, 21.iii.1998, A. Riedel (1, NHMS); Mando, 3N, Asaro, 18m Gorka, 27.vii.1972 (1, CNC); Goroka, 25.iii.1973, K.W. Stroeder (1, CNC); New Guinea: Lae, 7.i.1970, R.E. Parrott (1, CNC); Wau, Mt. Kaindi, 4.v.1971, R.E. Parrott (1, CNC); Nondugl, 1600m, ix-xi.1951, G. Gyldenstolpe (2, NHRS); Goroka, iii.1974, H. Ohlmus (1, ANIC); Amazon Bay area, Komania, 3400ft, 11–26.xi.1962, W. W. Brandt(1, ANIC); Western Highlands, Jimi River, 4700ft, 16.vii-21.ix.1961, W.W. Brandt (1, ANIC); M’wakA, viii.1975, H. Ohlmus (5, ANIC); Wabag, 24.vii.1974, H. Ohlmus (3, ANIC); Bosavi, v.1975, H. Ohlmus (1, ANIC); Mendi, xi.1975, H. Ohlmus (1, ANIC); Kmu, 27.xii.1969, H. Ohlmus (2, ANIC); Frigano, v.1974, H. Ohlmus (3, ANIC); Lae, xii.1968, H. Ohlmus (2, ANIC); Nambayufa, xii.1976, H. Ohlmus (1, ANIC); Wau, ii.1974, Willard H. Nutting, JR. Collection (1, CAS); Biarum MP 1800m, vi.1998 (2, ANIC); Wau, 1200m, 12–23.v.1965, H. Pyka (1, NHMS); Morobe Prov., Wau, Mt Kaindi, 1400m, 8.iii.1998, A. Riedel (1, NHMS); Goroka Highlands, 8.xi.1969, J.E. Tobler (1,CAS); Morobe District, Wau, Big Wau Creek, 1200m, 20.iv.1973, T.W. Davies (1,CAS). AUSTRALIA: Queensland: Mossman; Kuranda; Cairns; Kairi Creek, Tinaroo Dam; Walkamin; Kairi; Tolga; Upper Mulgrave River; Atherton; East Barron; Herberton; Crater National Park; Atherton Tableland; Evelyn; Millaa Millaa; Nerada; South Johnstone; Mt Spec; Paluma; Burleigh; Eungella; Bundaberg; Camp Milo, Cooloola; Gympie; Montville; Caloundra; Bunya Mountains; Mt Glorious; Brisbane; Moreton Bay; North Stradbroke Island; Toowoomba; Gatton; Mount Tamborine; Canungra; Cunningham’s Gap National Park; Currumbin Creek; Lamington Natl Park. New South Wales: Murwillumbah; Mullumbimby; Rosebank; Clarence River; Dunoon; Grafton; Coffs Harbour; Dorrigo; Boambee; Bonville;

Distribution of Harmonia testudinaria

Port Macquarie; Comboyne; Barrington Tops; Narara; Gosford; Blue Mountains; Kenthurst; Cabramatta; Sydney; Helensburgh; Minnamurra Falls; Kiama; Gerroa; Durras North; Hastings. Norfolk Island. Lord Howe Island.

Distribution Widely distributed species on the island of New Guinea and in eastern Queensland and New South Wales, Australia.

Harmonia tobleri sp. nov. (Fig. 60) Diagnosis Harmonia tobleri is very similar to H. solomonensis, which co-occurs on the same islands of the archipelago, and shares a similar black and orange colour pattern on the elytra. However, Harmonia tobleri can be distinguished by the elytral epipleuron bearing extensive black areas (Fig. 60B), and by very distinctive male genitalia.

Description Colour pattern (Figs 60A–C). Head uniformly reddish. Pronotum black, except for a narrow pale antero-lateral area (Fig. 60C). Scutellum black. Elytra black, with two yellowish or reddish markings on each elytron, a large basal area and a smaller and round spot apically; both spots occasionally connected medially; epipleuron orange at base and apex, but black in medial section. Prosternum, mesoventrite, mesepimeron, the anterior part of meso-metaventral process and the external borders of abdomen yellow in males; ventral side black except for the yellow mesepimeron, partial elytral epipleuron and the narrowed external borders in females. Legs yellow in males and with dark coxae in females. Morphology. Length 6.5–8.5 mm. Frons between the eyes about twice as broad as the eye width. Pronotum ~0.6 times

VII. Review of the Australo-Pacific Coccinellini

59

Distribution of Harmonia tobleri

as broad as the elytra, the lateral margin evenly arcuate, the anterior margin slightly arcuate. Prosternal process with carinae extending to the middle of the prosternum. Scutellum at the base ~1/10 times as broad as the elytra. Elytra narrowly explanate externally with large punctures along external borders. Epipleuron ~1/6 as broad as elytra. Anterior margin of the mesoventrite almost straight. Abdominal postcoxal line (Fig. 60I) approaching the posterior margin of ventrite 1; an oblique postcoxal line distinct. Posterior margin of ventrite 5 broadly emarginate in males and almost truncate in females. Posterior margin of ventrite 6 lightly emarginate in males and broadly arcuate in females. Male terminalia (Figs 60E–H). Female terminalia (Fig. 60D). Infundibulum slim, flared apically; the bursa copulatrix lightly sclerotised. Measurements (n = 3): TL 6.50–8.50 (7.52 ± 0.77); PL 1.00–1.20 (1.10 ± 0.06); EL 5.30–7.20 (6.35 ± 0.71); PW 3.10– 4.00 (3.53 ± 0.37); EW 5.40–7.10 (6.50 ± 0.70); GD 2.90–3.70 (3.40 ± 0.30). Ratios: PL/PW 0.28–0.35 (0.31 ± 0.03); EL/EW 0.90–1.01 (0.98 ± 0.04); EL/PL 5.30–6.45 (5.76 ± 0.42); EL/ GD 1.82–1.95 (1.86 ± 0.05); TL/EW 1.07–1.20 (1.16 ± 0.05).

Specimens examined Holotype: SOLOMON ISLANDS: Kolombangara Isl., Ringi Cove, Main Road near R6 Road, 17.i.1970, James E. Tobler (CAS). Paratypes: Kolombangara Isl., Ringi Cove, Main Road near R6 Road, 17.i.1970, James E. Tobler (18, CAS; 5 ANIC; 2, SYSBM); Kolombangara Isl., Ringi Cove, Main Road near R6 Road, 16.i.1970, James E. Tobler (20, CAS); Guadalcanal: Suta, 500–1200m, 27.vi.1956, J. L. Gressitt (1, BPBM); Guadalcanal: Kolsulu, 20.v.1960, C. W. O’Brien (BPBM).

Distribution Endemic to the Solomon Islands.

Remarks Named after James E. Tobler, collector of the holotype.

Heterocaria Timberlake, 1943 (Figs 61–71)

Heterocaria Timberlake, 1943: 58. Type species: Heterocaria papuana Timberlake, 1943 by original designation. Oiocaria Iablokoff-Khnzorian, 1982: 154. Type species: Oiocaria samuelsoni Iablokoff-Khnzorian, 1982 by original designation. Syn. nov.

Diagnostic combination Length 3.2–5.3 mm. Elytra moderately to strongly convex, glabrous. Anterior clypeal border straight between lateral projections. Antenna 11-segmented; as long as the width of the head capsule, with three-segmented club; terminal antennomere longer than the penultimate, rounded apically. Terminal maxillary palpomere securiform. Pronotal disc evenly convex transversely, with narrowly upturned or slightly thickened external borders. Prothoracic hypomeron without foveae near the anterior angles; the prosternal process broad, with distinct carinae extending forward to the middle of the prosternum. Anterior margin of the mesoventrite emarginate medially. Elytra with lateral margins with complete beads; epipleuron without foveae. Mid and hind femurs do not extend beyond the border of the elytra. Tibial spur formula 0–0–0. Abdominal postcoxal line not recurved, incomplete laterally; oblique postcoxal line absent. Penis consists of two sclerites; tegmen stout, the penis guide extending beyond the parameres apically, strongly bent towards the parameres in lateral view. Infundibulum absent; spermatheca C-shaped, with slim ramus and nodulus.

Description Body broadly oval, moderately convex; dorsum glabrous. Head. Eyes partially beneath the pronotum, inner orbits parallel. Frons as broad as the eye width. Antenna 11-segmented, as long as the width of thehead capsule; scape of typical shape; antennal club weak, three-segmented, loosely articulated; terminal antennomere elongate-oval. Clypeus short, the anterior border straight medially between two lateral

60

Ladybird Beetles of the Australo-Pacific Region

projections. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere broadened, forming an equilateral triangle. Prothorax. Pronotum weakly convex, the anterior border almost straight; the lateral borders slightly arcuate. Prothoracic hypomeron concave with shallow foveae. Prosternal process broad, almost as broad as the coxal diameter; surface with longitudinal carinae extending to the middle of the prosternum, converging weakly anteriorly. Pterothorax. Mesoventrite at the middle 0.8 times as long as the mesocoxae diameter; mesal surface with a well-delimited fossa receiving the prosternal process; the anterior margin depressed medially, with a raised ridge. Postcoxal line on the metaventrite joined at the middle, incomplete laterally. Elytra at the base much broader than the pronotum; margins explanate laterally, very upturned near the shoulders; disc densely punctured. Elytral epipleuron without delimited foveae. Tibiae slim and long, without apical spurs; tarsal claws each with a subquadrate basal tooth. Abdomen with six ventrites in both sexes. Postcoxal line close to the posterior margin of the ventrite, not recurved, almost reaching the lateral border; an oblique postcoxal line absent. Male terminalia. Penis tube-like, slim and long, consisting of two sclerites; the apical sclerite short and often complex; capsule T-shaped, with the inner arm much longer than the outer arm. Tegminal phallobase short, 0.3 times as long as the parameres. Female terminalia. Spermatheca comparably small, surface smooth. Ramus and nodulus large, but lightly sclerotised. Sperm duct uniform in diameter. Infundibulum absent. Styli terminals well developed.

Distribution Known from Australia and New Guinea.

Biology The biology of this genus is unknown. The Australian species H. barronensis was reared by J.J. Houston (QDPI, Brisbane) using Hyperomyzus aphids on Sonchus. The larva of the same species was described and illustrated by Ślipiński (2007).

Remarks Heterocaria was described by Timberlake for a single new species, H. papuana, and is characterised by its extremely elongate and tubular penis. That genus was ignored by IablokoffKhnzorian (1982), who placed H. papuana in Docimocaria, apparently due to the similarity in male genitalia. IablokoffKhnzorian failed to recognise the similarities of H. papuana with the newly erected genera Oiocaria (1982) and Archegleis (1984), as well as with some species included in his expanded concept of Lemnia. Six Australian species of Archegleis were reported by Pope (1989), but we have excluded A. barronensis and A. delta from that genus. Based characters of male and female genitalia these two species are here included in Heterocaria. Both Heterocaria and Archegleis are externally very similar, and in the key to genera the distinguishing characters of the shorter abdominal postcoxal lines in Archegleis may not be easily observed and are variable. Heterocaria may still be an aggregate genus as there is a considerable variation in the structure of the male genitalia, particularly in the penis, but the scarcity of available material and the lack of molecular data for this group made it impossible for us to categorise it any further. Iablokoff-Khnzorian (1982) recorded Coelophora schultzei (Weise, 1910) from New Guinea based on a male from Baiyer River. The specimen he examined and illustrated certainly belongs to an undescribed Heterocaria, but the identification as Coelophora schultzei is incorrect. We were not able to examine the original series of C. schultzei to confirm its identity, but the original description by Weise (1910) is detailed enough to confirm the mistake by Iablokoff-Khnzorian (1982).

Key to known species of Heterocaria Elytra yellowish-green with longitudinal fasciae (Fig. 65A)....................................................................................................H. delta Elytra with different colour pattern than that above ........................................................................................................................... 2 Pronotum entirely orange, or with a black transverse stripe at the base (Fig. 68A) ........................................................................... 3 Pronotum mostly black, or with a medial black area much larger and usually extending to at least the middle of the pronotum (Figs 63B, 64A) ........................................................................................................................................................................... 5 3(2). Elytra with broad yellow lateral margins; disc with mottled black and grey pattern (Fig. 63A) .............................H. cinerea [part] – Elytra black with lateral margins black; colour pattern different than that above .............................................................................. 4 4(3). Elytra with large orange spots covering most of the surface (Fig. 62A) .................................................................................H. bella – Elytra with very small orange spots andmost of the elytral surface black (Fig. 68A) ...................................................... H. papuana 5(2). Elytra with lateral margins completely yellow, or only with the apex black ...................................................................................... 6 – Elytra with lateral margins entirely or mostly black ........................................................................................................................... 8 6(5). Elytra black, with at least the apical part bearing white or yellow isolated spots (Fig. 70A) ............................ H. samuelsoni [part] – Elytra black, with large red and yellow spots (Figs 66A–C) .............................................................................................. H. funerea – Elytral pattern different than those listed above ................................................................................................................................. 7 7(6). Elytra with yellow and black colour pattern, usually resembling a black cross on a yellow or orange background (Figs 67A,B) ���������������������������������������������������������������������������������������������������������������������������������������������������������������������������H. kaszabi [part] – Elytra mostly with a mottled pattern of dark punctures on lighter background (Fig. 63B), or rarely uniformly dark (Fig. 63C) ���������������������������������������������������������������������������������������������������������������������������������������������������������������������������H. cinerea [part] 8(6). Elytra yellow or orange, with a black suture and external margins (Fig. 67C) ........................................................H. kaszabi [part] – Elytra with different colour pattern than above .................................................................................................................................. 9 9(8). Pronotum with medial black marking extending from the base to the anterior margin (Fig. 64C) .................................................. 10 – Pronotum with medial black marking distinctly separated from the anterior margin (Fig. 69C) ..................................................... 11 10(9). Elytra with small isolated spots (Fig. 64A) ............................................................................................................ H. decemmaculata – Elytra with transverse yellow fasciae (Fig. 71A) ................................................................................................ H. transversoguttata 11(9). Elytron with eight small markings (Fig. 70A) ................................................................................................... H. samuelsoni [part] – Elytra with several fasciae often interconnected (Fig. 61A) ............................................................................................................. 12 12(11). Australian species; elytral margins entirely black (Fig. 61B) ...................................................................................... H. barronensis – New Guinean species; elytral margins near the humeri yellow (Fig. 69B) .......................................................................... H. riedeli 1. – 2(1). –

VII. Review of the Australo-Pacific Coccinellini

61

Heterocaria barronensis (Blackburn, 1895) comb. nov. (Fig. 61)

Cycloneda barronensis Blackburn, 1895: 238; Type locality: Queensland, Barron River. Lea 1902: 489. Callineda barronensis: Weise 1923: 133; Korschefsky 1932: 282. Egleis barronensis: Timberlake 1943: 27. Harmonia barronensis: Iablokoff-Khnzorian 1982: 461. Lemnia barronensis: Iablokoff-Khnzorian 1984a: 206. Archegleis barronensis: Iablokoff-Khnzorian 1984b: 118; Pope 1989: 697 (lectotype); Ślipiński 2007: 151.

Diagnosis Heterocaria barronensis is distinguished by its circular and strongly convex body, pronotum with a medial black area not reaching the anterior margin, and the elytra bearing a complex pattern of whitish or yellow fasciae on a black background (Fig. 61A).

Distribution of Heterocaria barronensis

Description Colour pattern (Figs 61A–C). Frons yellow, except for two black markings at the base in males, with black internal margins close to eyes in females. Head appendages yellow. Labrum yellow. Pronotum yellow, with two black markings, the basal marking situated medially and connected with the base, the anterior marking strongly transverse and connected with the anterior border between the lateral projections. Scutellum black. Elytra black with three yellow markings on each elytron, in some specimens the middle marking is separated into two spots, the lateral borders and suture black. Prothoracic hypomeron pale, except for the posterior half of the lateral external border black. Prosternum pale with a black area near thecoxae, the process usually dark and broad, the pterothorax usually dark except for the pale mesepimeron. Abdomen yellow, except for the central part of ventrite 1 black. Front legs usually yellow, mid and hind legs with femur black in females and usually yellow in males. Morphology. Length 4.7–5.6 mm. Frons slightly broader than the eye width. Prosternal process with long, almost parallel carinae extending forward 4/5 to the anterior margin. Elytra slightly shorter than broad, with broadest part at 1/3 of the elytra from the base. Scutellum at the base ~1/11 as broad as the elytra at the shoulder area. Elytral borders explanate laterally, with thickened margins, upturned near the shoulders. Anterior border of mesoventrite shallowly and broadly emarginate, with raised carinae connected medially. Metaventrite with a postcoxal line almost straight medially. Elytral epipleuron complete from base to apex, ~1/8 times as broad as the elytra. Abdominal ventrite 5 with the posterior margin truncate in males and slightly arcuate in females; ventrite 6 with the posterior margin slightly emarginate in males and arcuate in females. Male terminalia (Figs 61D–G). Female terminalia (Figs 61H,I). Measurements (n = 7): TL 4.70–5.65 (5.19 ± 0.31); PL 0.70–0.95(0.79 ± 0.09); EL 3.80–4.85 (4.29 ± 0.32); PW 2.60– 3.00 (2.77 ± 0.15); EW 4.20–5.00 (4.57 ± 0.29); GD 2.10–2.80 (2.45 ± 0.26). Ratios: PL/PW 0.24–0.37 (0.28 ± 0.04); EL/EW 0.90–1.00 (0.94 ± 0.04); EL/PL 4.42–6.93 (5.52 ± 0.74); EL/ GD 1.65–1.87 (1.76 ± 0.08); TL/EW 1.08–1.24 (1.14 ± 0.06).

Types C. barronensis: Type/ 7544 N. Qu. T/ Neda barronensis Blackb. (BMNH, lectotype).

Specimens examined AUSTRALIA: Queensland: 17.10S 145.93E, Danbulla SF., Entrance, 13.vi.1992, C. Reid, beating vines, rainf. trees, Acacia (4, ANIC); Speewah, viii.1969, R. E. Parrott (1, ANIC); Lake Eacham, 20.v.1952, J. Sedlacek (1, BPBM); Little Crystal Ck. Bridge, rd. to Paluma, 28.vi.1992, C. Reid, beating rainf. bushes (1, ANIC); Kuranda (1, ANIC); Kuranda, iv.1947, J.G. Brooks, (2, ANIC); Innisfail, 28.x.1934, J.G. Brooks (1, ANIC); Cairns, W. Ferguson (2, ANIC).

Distribution This rarely collected species is known from northern Queensland, primarily from the Atherton Tablelands, and a narrow stretch along the coast from Kuranda to Paluma.

Heterocaria bella sp. nov. (Fig. 62) Diagnosis Heterocaria bella is easily distinguished from other Australian and Papuan species by its entirely reddish head and pronotum, and elytra covered by regular yellow spots on a black background.

Description Colour pattern (Figs 62A–C). Frons and head appendages yellow. Pronotum yellow, except for the black basal border at the middle and the lateral margin also partially black. Scutellum black. Elytra black, each with nine large yellow spots; the lateral border and suture black. Ventral side black, except for the prosternum, prosternal hypomeron, mesepimeron and the central part of epipleuron all light yellow. Front and middle legs yellow, hind legs yellow with the femur partially black. Morphology. Length 4.2–4.5 mm. Frons flat, 1.4 times as wide as the eye. Eyes large, obscured partially beneath the anterior margin of the pronotum, the interior orbits almost parallel. Prothorax. Pronotum strongly transverse, with the lateral margins distinctly arcuate posteriorly; disc weakly convex, with very fine and sparse punctures; interspaces shiny. Prosternal process with sharp carinae converging anteriorly,

62

Ladybird Beetles of the Australo-Pacific Region

deep punctures and bearing a silver-grey or black coarse pattern on an indistinct yellowish background. The darker forms (Fig. 63C) have almost uniformly black elytra, with distinct yellow margins not seen in any other Heterocaria.

Description

Distribution of Heterocaria bella

extending almost to the anterior margin of the prosternum. Scutellum triangular, at the base ~1/10 times as broad as the elytra at the shoulder area. Elytra with lateral margins narrowly explanate, the lateral margins with a complete bead; disc distinctly punctate, dorsal punctures at least twice as large as the pronotal ones, becoming large to very coarse towards the margins. Mesoventrite broad, with the anterior margin shallowly emarginate. Abdomen with posterior margin of ventrite 5 strongly emarginate medially in males. Male terminalia (Figs 62D–G). Penis slim and very long, tube-like. Tegmen with penis guide slightly longer than the parameres, strongly bent towards the parameres apically. Female. Unknown. Measurements (n = 2): TL 4.2–4.5; PL 0.70; EL 3.50–3.80; PW 2.30–2.40; EW 3.80–3.90; GD 2.00–2.10. Ratios: PL/PW 0.29–0.30; EL/EW 0.92–0.97; EL/PL 5.00–5.43; EL/GD 1.75– 1.81; TL/EW 1.11–1.15.

Specimens examined Holotype: AUSTRALIA. Queensland: Gordon Creek, Claudie River District, QLD, 12.42 S 143.17 E, 10.xii.1986, G. Daniels & M. A. Schneider (QMB). Paratype: West Claudie River, QLD, 25.vi.1982, M. A. Schneider & G. Daniels (ANIC).

Distribution Known from a single locality near the Iron Range in north Queensland Australia.

Remarks The species name is derived from the feminine Latin adjective bella meaning beautiful, pretty.

Colour pattern (Figs 63A–D). Frons yellow, with a black medial patch in some specimens. Clypeus and labrum yellow. Head appendages yellow. Pronotum yellow, or with medial trapezoidal black area. Prothoracic hypomeron yellow. Elytra with a silvery brown pattern of irregular punctures on a lighter background, rarely uniformly brown or black with yellowish apices and margins. Scutellum black. Tibia yellow or brown; anterior and middle femur pale, hind femur usually black. Abdominal ventrite 1 black or partially black, remaining parts yellow. Morphology. Length 3.9–4.7 mm. Frons 2.0–2.2 times broader than the eye width. Pronotum smoothly arcuate anteriorly; convex and finely punctured. Scutellum triangular, 1/14 width of the elytra at the base. Elytra slightly broader than the pronotum at the base, arcuate and distinctly narrowing apically, moderately convex; disc deeply punctured; the lateral margins narrowly explanate; the epipleuron about twice the width of the metanepisternum. Prosternum with longitudinal carinae extending close to the anterior margin. Male terminalia (Figs 63E–H). Female terminalia (Figs 63I,J). Measurements (n = 15): TL 3.90–4.70 (4.18 ± 0.21); PL 0.60–0.80 (0.66 ± 0.06); EL 3.25–3.90 (3.48 ± 0.18); PW 1.95– 2.50 (2.23 ± 0.13); EW 3.05–4.00 (3.41 ± 0.22); GD 1.75–2.20 (1.90 ± 0.11). Ratios: PL/PW 0.26–0.35 (0.30 ± 0.02); EL/EW 0.97–1.10 (1.02 ± 0.04); EL/PL 4.50–5.92 (5.31 ± 0.42); EL/ GD 1.71–2.00 (1.84 ± 0.07); TL/EW 1.16–1.29 (1.23 ± 0.04).

Types L. cinerea: Papua New Guinea, Aiyura (Kainantu), sur taro, J. Gutierrez, 24–Viii-1982/ Lemnia kaszabi cinerea, n. ssp., Holotype J. Chazeau det. 1988 (MNHN, holotype).

Specimens examined INDONESIA: Western New Guinea: Prov. Jayawijaya, Wamena, Pronggoli, 2000–2400m 17–19.ix.1991, A. Riedel (1,); Jayawijaya. Emdoman-okloma. 16.ix.1992, 1660–2000m, A. Riedel (1, NHMS). PAPUA NEW GUINEA: Wau, Morobe Distr. 1100m, 29.viii.1961, J. Sedlacek (6, BPBM); Wau: Edie Creek Road: 1280m, 19.vi.1984, W.C. Gagne (3, BPBM); Wau, 1200m, 2.ii.1966, J. & M. Sedlacek (1, BPBM); Kassem Pass 1400m, 4.i.1965, J. & M. Sedlacek (1, BPBM); Wau, Big Wau Ck, 1200–1500m, xi.1965, J. & M. Sedlacek (1, BPBM); NE, E. Highlands, Purosa, 1700m, 17–25.v.1996, Gressitt & Tawi (1,

Heterocaria cinerea (Chazeau, 1990) stat. et comb. nov. (Fig. 63)

Lemnia kaszabi cinerea Chazeau, 1990: 165. Type locality: Papua New Guinea, Aiyura Valley.

Diagnosis Heterocaria cinerea is easily diagnosed by its elongate-oval body, at least laterally yellow pronotum, and the elytra with

Distribution of Heterocaria cinerea

VII. Review of the Australo-Pacific Coccinellini

63

BPBM); NE W. Highlands, Baiyer R. 1500–1600m, 14.xi.1971. N.L.H. Krsusa (1, BPBM); Wau, ii.1974 (1, CAS).

Distribution Endemic to the island of New Guinea.

Remarks Chazeau (1990) described this species as a subspecies of H. kaszabi, recognising the pronounced morphological differences in colour pattern, but close similarity in male terminalia between H. kaszabi and H. cinerea. We have examined many individuals of this species, and found small but consistent differences in the male terminalia between H. kaszabi and H. cinerea, and regard them as a separate species rather than geographic subspecies, as both forms occur sympatrically at least in the Wau and Western Highlands areas of Papua New Guinea.

Heterocaria decemmaculata sp. nov. (Fig. 64) Diagnosis Heterocaria decemmaculata can be distinguished from similar Papuan species in having black elytral margins, a pronotal central black marking extending from the base to the anterior margin, and the elytra bearing 10 small circular spots.

Description Colour pattern (Figs 64A–C). Frons and head appendages yellow. Pronotum medially black, with broad yellow areas laterally; the lateral margins black. Scutellum black. Elytra black, each with five large yellow spots arranges as in Fig. 64A, the external border and suture black. Ventral side yellow, except for the metaventrite, metanepisternum, the external borders of the elytral epipleuron and the central part of first and second abdominal ventrites black. Legs brown. Morphology. Length 5.2 mm. Frons 1.4 times broader than the eye width. Pronotum ~0.9 times as broad as the elytra at the base. Anterior border almost straight medially; the lateral margins weakly arcuate, explanate and distinctly upturned; pronotal disc finely punctured, punctures as large as the eye facets, 1–2 diameters apart; interspace weakly reticulate. Prosternal process broad, the surface with long parallel carinae reaching almost to the anterior margin. Scutellum ~1/10 times as broad as the elytra, with the basal margin as long as the lateral margin. Elytra slightly broader than long, narrowing distinctly apically. Elytron with lateral borders distinctly explanate, the margin with a complete bead and upturned near the shoulders. Elytral epipleuron ~2.5 times as broad as the metanepisternum with shallow fossae to receive the apices of the hind femora. Elytral disc with variable punctures, most twice larger than those on the pronotum, becoming larger laterally, and very coarse on the sloping explanate parts. Anterior border of the mesoventrite shallowly and broadly emarginated, with raised carinae connected medially, the posterior border almost straight. Abdominal postcoxal line (Fig. 64I) does not extend to the posterior margin, complete laterally. Ventrite 5 with truncate posterior margin, ventrite 6 with the posterior margin slightly emarginate medially. Male terminalia. Penis strongly bent apically, with long sclerite (Fig. 64D). Tegmen slim and long, the penis guide

Distribution of Heterocaria decemmaculata

slighter longer than the parameres; the middle paramere with a large projection strongly arcuate and crenulated between the projection and the apex (Fig. 64G). Female. Unknown. Measurements. TL 5.25; PL 0.90; EL 4.35; PW 2.95; EW 4.55; GD 2.50. Ratios: PL/PW 0.31; EL/EW 0.96; EL/PL 4.83; EL/GD 1.74; TL/EW 1.15.

Specimens examined Holotype: INDONESIA. Western New Guinea: ‘Aimau Dutch, New Guinea, 16.x.1939, R. G. Wind’ (CAS, male).

Distribution Known only from Western New Guinea, Indonesia.

Remarks The species name is formed from the Latin numeral decem, referring to the number of elytral spots.

Heterocaria delta (Weise, 1898), comb. nov. (Fig. 65)

Egleis delta Weise, 1898: 225. Type locality: New South Wales. Korschefsky 1932: 576; Timberlake 1943: 26. Archegleis crotchi Iablokoff-Khnzorian, 1984b: 116. Type locality: Australia. Synonymised by Pope 1989: 698. Archegleis delta: Pope 1989: 698 (lectotype); Ślipiński 2007: 151.

Diagnosis The elytral colour pattern of Heterocaria delta is unique in this genus, but it strongly resembles some species of Archegleis. However, it can be distinguished from Archegleis, in addition to the specific characters of the genitalia, by a black suture and a smooth and shiny pronotal disc between fine punctures.

Description Colour pattern (Figs 65A–C). Frons yellow, with a small black patch on either side of vertex in males, and narrowly black anteriorly and laterally above the antennal insertion in females. Head appendages yellow. Pronotum yellow, with a black base and a central marking shaped like a ‘Y’, the lateral margin largely black. Scutellum black. Elytra greenish yellow, with black stripes arranged as Fig. 65A, the external border and suture black. Ventral side background colour black except for the yellow prothoracic hypomeron, the central elytral epipleuron, and the central part of the abdomen. Front and mid legs largely pale, while hind legs largely black.

64

Ladybird Beetles of the Australo-Pacific Region

Morphology. Length 6.1–6.7 mm. Frons 1.5 times as broad as the eye width. Eyes partially obscured beneath the pronotum. Antenna about twice as long as the frons width, club weakly three-segmented, loosely articulated, the terminal antennomere elongate-oval. Prosternal process broad, the surface with long and almost parallel carinae extending close to the anterior margin. Elytra with the lateral borders explanate with a thickened margin upturned near the shoulder. Scutellum 1/12 as broad as the elytra. Anterior border of the mesoventrite shallowly and broadly emarginate, with raised carinae connected medially, the posterior border slightly arcuate. Metaventrite with a postcoxal line almost straight medially. Elytral epipleuron 2.3–2.5 times as broad as the metanepisternum. Abdominal postcoxal line close to the posterior margin of ventrites and complete laterally. Ventrite 5 with the posterior margin truncate in males and slightly arcuate in females; ventrite 6 with the posterior margin weakly emarginate in males and arcuate in females. Male terminalia (Figs 65D–G). Female terminalia. Not examined. Measurements (n = 3): TL 6.10–6.70 (6.37 ± 0.31); PL 0.90–1.10 (0.97 ± 0.12); EL 5.20–5.60 (5.40 ± 0.20); PW 3.00– 3.40 (3.20 ± 0.20); EW 5.50–5.90 (5.63 ± 0.23); GD 2.95–3.80 (3.25 ± 0.48). Ratios: PL/PW 0.28–0.32 (0.30 ± 0.02); EL/EW 0.95–0.98 (0.96 ± 0.02); EL/PL 5.09–6.00 (5.62 ± 0.47); EL/ GD 1.37–1.90 (1.69 ± 0.28); TL/EW 1.11–1.15 (1.13 ± 0.02).

Types E. delta: N.S. Wales Stauding/ Egleis delta m/ Holotypus Khnz 76 Egleis delta Ws (ZMB, lectotype). A. crotchi: Type/ Type Edwardsi Reich (UCCC, holotype).

Specimens examined AUSTRALIA: Queensland: Mt. Tamborine (2, ANIC); D’Aguilar National Park, 7.viii.2004, M. Wanat (ANIC). New South Wales: Tooloom, 29.x.1961, I.F.B. Common & M.S. Upton (1, ANIC); Red Cedar FR, Wild Cattle Creek, 9.iv.1993, C. Reid (1, ANIC); Orara River, H. Davidson (ANIC).

Distribution Australia, known from a few localities in southern Queensland and northern New South Wales.

Heterocaria funerea (Crotch, 1874) comb. nov. (Fig. 66)

Halyzia funerea Crotch, 1874: 132: Type locality: New Guinea, Dorey [= Manokwari] – Korschefsky 1932: 571. Lemnia (s. str.) funerea: Iablokoff-Khnzorian 1982: 258; Gordon 1987: 15. Coelophora extensa Weise, 1902: 499. Type locality: New Guinea. Syn. nov. Coelophora extensa var. mysolensis Weise, 1902: 499. Type locality: Mysol. Lemnia (s. str.) extensa: Iablokoff-Khnzorian 1982: 254.

Diagnosis Heterocaria funerea can easily be recognised by its unique orange and red spots on black elytra.

Description Body weakly convex, elytra broadly explanate. Colour pattern (Figs 66A–C). Pronotal and elytral colour pattern variable. Head and head appendages yellow. Prothoracic hypomeron, prosternum and abdomen yellow. Scutellum black. Pterothorax dark, except for the pale mesanepisternum and mesepimeron. Elytral epipleura usually pale, with external borders black. Legs usually yellow but often hind legs partially black. Morphology. Length 5.4–7.0 mm. Eyes partially beneath the pronotum, the interior orbits almost parallel. Antennae long, almost as long as the head capsule. Pronotum weakly convex; prothoracic hypomeron without a well-delimited fovea. Prosternal process extending well beyond the anterior coxae; surface with longitudinal carinae converging faintly anteriorly. Elytra with the lateral border broadly explanate, upturned close to the shoulder. Anterior margin of the mesoventrite emarginate medially. Abdominal postcoxal line approaching the posterior margin but not extending to the lateral margin (Fig. 66J). Posterior margin of ventrite 5 truncate in males and slightly arcuate in females. Male terminalia (Figs 66D–G). Penis consists of a single sclerite, strongly curved, with membranous tissue apically. Female terminalia (Figs 66H,I). Spermatheca C-shaped, with an indistinct nodulus; coxites broadened at the middle, pointed apically, with small styli. Measurements (n = 9): TL 5.40–7.20 (6.34 ± 0.64); PL 0.90–1.10 (1.03 ± 0.07); EL 4.50–6.10 (5.26 ± 0.54); PW 3.20– 4.00 (3.54 ± 0.30); EW 4.70–6.80 (5.76 ± 0.71); GD 2.65–3.70 (3.06 ± 0.42). Ratios: PL/PW 0.28–0.32 (0.29 ± 0.02); EL/EW 0.87–0.96 (0.92 ± 0.03); EL/PL 4.64–5.55 (5.09 ± 0.28); EL/ GD 1.64–1.81 (1.73 ± 0.07); TL/EW 1.06–1.16 (1.10 ± 0.04).

Types H. funerea: TYPE funereal Cr/ TYPE (UCCC, holotype). L. extensa: New Guinea Aegna/ Coelophora extensa m/ 1078/ (ZMB, lectotype). L. extensa var. mysolensis: Misool Insel/ Type/ extensa Ws v. mysolensis Ws/ Typen (ZMB, lectotype and paralectotype; here designated).

Specimens examined

Distribution of Heterocaria delta

INDONESIA: Western New Guinea: Nabire S Geelvink Bay, 10–40m, 1–4.ix.1962, J. Sedlacek (1, BPBM); Nabire, 5–50m, 25.viii-2.ix.1962, J. Sedlacek (1, BPBM); Japen I.,

VII. Review of the Australo-Pacific Coccinellini

Distribution of Heterocaria funerea

SSE Sumerbaba, Dawai R. Jungle, 20.x.1962, H. Holtmann (1, BPBM); Irian Jaya: Japen Serui N. Ambeidiru, 1000m, 5.viii.1996, Schule & Stuben (2, Stuttg); Cyclops Mts., 600m, 4km nordi Sentani, 8–13.ix.1990, Balke & Hendrich (3, ZMB); Sorong 11.v.1949, Sten Bergman (1, Riksmuseum li2705) N. Guin, Dorey, 370. (1, MIIZ). PAPUA NEW GUINEA: NE Feramin, 1450m, 21.viii.1963, R. Straatman (2, BPBM); NE Angoram, 10m, 13.viii.1969, J. L. Gressitt (1, BPBM); Bulolo, 30.v.1971, R.E. Parrott (1, CNC); NE Morobe District, Ulap, 800–1100m, ix.1968, N.L.H. Krauss (1, BPBM); Lae, 30.i.1971, R.E. Parrott (1, CNC); 16 miles Port Morobe, 22.xii.1973, D. Markay (1, AM); Klunga, 13.v.1992, J.F. Grimshaw (1, NAQS); Daru, 09.04S- 143.13E, 23–29.xi.1989, J.W. Turner (1, NAQS); Daru, ex. Sugarcrane PNG 485, 23–29.xi.1989, J.F. Grimshaw (1, NAQS); Pasi nr. Vanimo, 9.vi.1993, J.F. Grimshaw (1, NAQS); Kiunga, ex. Guava JFG 1264, 13.v.1992, J.F. Grimshaw (1, NAQS); Mt. Lamington Dist., Northern Division, Papua, 1928, C.T. McNamara (1, SAM); Ponpondetta Dist, Sangara, 22.iii.1965, E.S. Brown (1, BMNH); Madang Dist., Finisterre Mts, 5500 ft, 30.x.-15.xi.1964, C. Morro (1, BMNH); Mt. Tafa, 8500ft, ii.1934, L.E. Chessman (1, BMNH).

Distribution Endemic to the island of New Guinea.

Heterocaria kaszabi (Iablokoff-Khnzorian, 1982) comb. nov. (Fig. 67)

Lemnia (Lemnia) kaszabi Iablokoff-Khnzorian, 1982: 223. Type locality: New Guinea, Wau. Lemnia kaszabi kaszabi: Chazeau 1990: 164.

Diagnosis The typical form of H. kaszabi has a black cross-like elytral pattern on a yellow or orange background, and is easily distinguished from other species. However, several less conspicuous colour variants have been examined and these should be identified using the diagnostic features of the male genitalia.

(Fig. 67A); elytra rarely almost entirely black, with small spots at humeri and apices, or only the suture and lateral areas dark, the remaining part of elytron yellow. Meso- and metathorax black, except for the yellow mesepimeron. Legs usually yellow, but hind femur always black. Morphology. Length 3.8–4.9 mm. Frons 1.4–1.6 times wider than the eye width. Pronotum weakly convex medially, weakly concave laterally; disc finely punctured. Pronotum widest just above the base, with the lateral margins more strongly curved anteriorly. Elytra slightly broader than the pronotum, strongly narrowed apically; disc coarsely punctured, with interspaces bearing micropunctures and often reticulate microsculpture. Elytral epipleuron without foveae. Abdominal postcoxal line (Fig. 67J) approaching the posterior margin of ventrite 1, extending to the lateral border. Posterior margin of ventrite 5 truncate in males, arcuate and with a very faint suture medially in females. Male terminalia (Figs 67F–H). Female terminalia (Fig. 67I). Measurements (n = 13): TL 3.80–4.85 (4.34 ± 0.31); PL 0.65–0.90 (0.76 ± 0.09); EL 3.15–4.10 (3.50 ± 0.25); PW 2.00– 2.40 (2.21 ± 0.13); EW 3.00–4.00 (3.40 ± 0.28); GD 1.65–2.20 (1.83 ± 0.14). Ratios: PL/PW 0.29–0.41 (0.35 ± 0.04); EL/EW 0.92–1.12 (1.03 ± 0.05); EL/PL 3.76–5.86 (4.64 ± 0.61); EL/ GD 1.78–2.08(1.92 ± 0.09); TL/EW 1.21–1.40 (1.28 ± 0.06).

Types L. kaszabi: Wau, McAdam Park, 18–21.4.1965, J. Balogh et J. Szent-Ivany leg (TMB, holotype).

Specimens examined INDONESIA: Western New Guinea: Baliem-tai. Jiwika 1700–2300m, 1–2.ix.1990, A. Riedel (2, NHMS); Baliem, Dist Wamena, 1600m, 31.viii.1990, A. Riedel (1, NHMS); Paniai Mulia (s.), Wuyuneeri, 1900–2200m, 6–7.vii.1994, A. Riedel (1, NHMS); Mokwam Warmare, 300–1400m, 19.iv.1993, A. Riedel (1, NHMS); Prov. Manokwari. Membey, 800–1200m, 31.viii.1991, A. Riedel (1, NHMS); Japen Serui n. Ambedirm, 1000m, 5.viii.1996, Schules/Stuben (1, NHMS). PAPUA NEW GUINEA: Upper Manki L.A. Bulolo M. Dist, in sticky trap, F. R. Wylie & P. Shanahan, 1.ix.1972 (2, ANIC); Morobe Prov., 1400–1800m, 20.iii.1998, A. Riedel (1, NHMS); E W. Highlands, Baiyer R. 1500–1600m, 14.xi.1971. N.L.H. Krsusa (1, BPBM); (NE) Wau, Morobe Distr. 1100m, 2.ix.1961, J. Sedlacek (5, BPBM); (NW) Wisselmeran Enarotadi, 1850m, 4.viii.1962; Koitaki 1500ft, x-xi.1928, Pemberton (1, ANIC).

Distribution Widely distributed in New Guinea.

Description Colour pattern (Figs 67A–E). Frons yellow or brown, sometimes with black spots at the base. Antennal club dark. Pronotum with transparent anterior and lateral margins, the disc yellow, with black markings medially, epipleuron yellow. Scutellum black. Elytra with variously developed black crosslike markings, with two or three yellow areas on each elytron

65

Distribution of Heterocaria kaszabi

66

Ladybird Beetles of the Australo-Pacific Region

Heterocaria papuana Timberlake, 1943 (Fig. 68)

Heterocaria papuana Timberlake, 1943: 58. Type locality: New Guinea, Laloki. Docimocaria papuana: Iablokoff-Khnzorian 1982: 141.

Central Dist. Koitaki Est. 28.iv.1957, in ornamental garden, J.J. Szent-Ivany (1, BMNH); same collector, Central Dist., Eriama Est., 29.iii.1956, comp with holotype, RGB (1, BMNH).

Distribution Endemic to New Guinea.

Diagnosis Heterocaria papuana is easily distinguished from other species in having an entirely orange pronotum and black elytra, each with nine small isolated spots, including three larger ones along the elytral bases.

Description Colour pattern (Figs 68A–C). Frons and head appendages yellow to orange. Pronotum orange, with the basal border narrowly black medially; the anterior border margin narrowly transparent. Scutellum black. Elytron black with nine small yellow spots. Prothorax orange on the ventral side; pterothorax black, except for the pale mesepimeron. Front leg orange, middle and hind legs largely black. Morphology. Length 5.3–5.5 mm. Frons 1.7–1.9 times as broad as the eye width. Eyes partially beneath the anterior margin of the pronotum, the ocular canthus large and almost rounded, the interior orbits slightly divergent anteriorly. Antennae long, length 1.5 times the frons width. Prosternal process broad, truncate apically, the surface with distinct long carinae extending to almost the anterior margin. Mesoventrite as broad as the mid-coxa, the anterior margin weakly emarginate at the middle, the posterior margin slightly arcuate medially. Metaventrite with a postcoxal line complete laterally, connected and straight medially. Abdominal postcoxal line (Fig. 68F) reaching the posterior margin, extended and complete laterally. Posterior margin of ventrite 5 weakly arcuate centrally in females. Male terminalia. Not examined. Female terminalia (Figs 68D,E). Measurements (n = 5): TL 5.30–5.50; PL 0.70–0.70; EL 4.50–4.50; PW 2.80–2.90; EW 4.80–4.90; GD 2.50–2.85. Ratios: PL/PW 0.24–0.25; EL/EW 0.92–0.94; EL/PL 6.43; EL/ GD 1.58–1.80; TL/EW 1.10–1.12.

Heterocaria riedeli sp. nov. (Fig. 69) Diagnosis Heterocaria riedeli is very similar to the Australian species H. barronensis, but it can be distinguished by its slightly larger size, different elytral colour pattern, and distinctive male genitalia.

Description

H. papuana: Papua, vii.1909, Laloki, F. Muir, Heterocaria papuana Timb. Type (BPBM, holotype).

Colour pattern (Figs 69A–C). Frons and head appendages yellow. Pronotum yellow anteriorly, with a large medial black area (Fig. 69A), not extending to the anterior margin. Scutellum black. Elytra black, each with a yellow rectangle beginning at the elytral base, and two spots laterally, one near the middle and another close to the apex; the external borders and suture black; the epipleuron usually pale anteriorly, posterior half black. Prosternum mostly pale, except for the black prosternal process; the hypomeron pale. Mesoventrite and metaventrite black; the mesanepisternum and mesepimeron pale. Legs yellow. Morphology. Length 5.1–6.3 mm. Frons 1.4–1.5 times as broad as the eye width. Pronotum weakly convex, the lateral margin upturned and thickened. Prosternal process setose; the prosternal carinae extending forward to almost the anterior margin. Elytra slightly broader than the pronotum at the base, with distinct lateral margins. Abdominal postcoxal line (Fig. 69I) approaching the posterior margin of the ventrite, extending close to the lateral margin. Posterior margin of ventrite 5 emarginate medially in males and arcuate in females. Male terminalia (Figs 69D–G). Female terminalia (Fig. 69H). Measurements (n = 17): TL 5.10–6.30 (5.61 ± 0.29); PL 0.85–1.10 (1.00 ± 0.08); EL 4.20–4.80 (4.41 ± 0.20); PW 2.30– 3.20 (2.82 ± 0.23); EW 4.10–4.90 (4.46 ± 0.28); GD 2.20–2.65 (2.38 ± 0.14). Ratios: PL/PW 0.31–0.48 (0.36 ± 0.04); EL/EW 0.94–1.05 (0.99 ± 0.03); EL/PL 3.82–5.11 (4.44 ± 0.37); EL/ GD 1.74–1.96 (1.86 ± 0.06); TL/EW 1.16–1.40 (1.26 ± 0.08).

Specimens examined

Specimens examined

Types

PAPUA NEW GUINEA: Koitaki 1500ft, x-xi.1928, Permberton (1, ANIC); Port Moresby, 29.xii.1973, D. Mackay (1, AM);

Distribution of Heterocaria papuana

Holotype: INDONESIA: Western New Guinea: Irian Jaya: Baliem Distr., Kangime, 1500–1600m, 3.ix.1990, A. Riedel (NHMS). Paratypes: Irian Jaya: Jayawijaya, Bime 1600–1900m, 22.ix.1993, A. Riedel (2, NHMS); Irian Jaya: Jayawijaya, Taramlu, 1500–1700m, 6.ix.1993, A. Riedel (4, NHMS); Irian Jaya: Jayawijaya, Bime, 1600–2000m, 10.ix.1993, A. Riedel (4, NHMS); Irian Jaya: Jayawi-Galbok b. Nalca, 1700–1800m, 3.x.1993, A. Riedel (4, NHMS); Irian Jaya: Jayawijaya, Bime 1600–1900m, 11.ix.1993, A. Riedel (3, NHMS); Irian Jaya: Pro. Jayawijaya, Wamena, AnggurukTanggeam, 1500–1800m, 28.ix.1991, A. Riedel (2, NHMS); Irian Jaya: Baliem Distr., Kangime, 1500–1600m, 3.ix.1990, A. Riedel (2, NHMS); Irian Jaya: Jayawijaya, Wamena, Pronggoli, 2000–2004, 17–19.ix.1991, A. Riedel (1, NHMS); Irian Jaya: Jayawijaya, Okloma,1650–1800m, 30.ix-1.x.1993,

VII. Review of the Australo-Pacific Coccinellini

Distribution of Heterocaria riedeli

A. Riedel (1, NHMS); Irian Jaya: Jayawijaya, Anggruk, 1200–1500m, 23.ix.1992, A. Riedel (1, NHMS); Irian Jaya: Jayawijaya, Membahan, 23.ix.1991, A. Riedel (1, NHMS); Irian Jaya: Baliem Dist., Kangime, 1500–1600m, 3.ix.1990 (1, NHMS); Irian Jaya: Jayawijaya, Membahan, 23.ix.1991, A. Riedel (1, NHMS); New Guinea: NETH., 8.xi.1958, J. L. Gressitt (1, BPBM). PAPUA NEW GUINEA: NE Feramin, 1450m, 26.viii.1963, R. Straatman (1, BPBM); New Guinea: NE Feramin, 1450m, 27.viii.1963, R. Straatman (1, BPBM).

Distribution Endemic to New Guinea.

Remarks This species is named after Dr Alexander Riedel, Staatliches Museum für Naturkunde Karlsruhe, Germany, who collected this and many other ladybirds during his explorations of New Guinea.

Heterocaria samuelsoni (Iablokoff-Khnzorian, 1982) comb. nov. (Fig. 70)

Oiocaria samuelsoni Iablokoff-Khnzorian, 1982: 155. Type locality: New Guinea, Wau.

Diagnosis Heterocaria samuelsoni can be distinguished by its largely yellow pronotum with a small black basal area in the middle, mostly black elytra, each with a transverse humeral fascia, extending shortly along the inner scutellar margin and suture, and five spots usually dispersed on the disc or partially connected laterally to a broad yellow margin.

Description Colour pattern (Figs 70A–C). Frons and head appendages yellow. Pronotum yellow, with a central black marking not extending to the anterior margin; the lateral margins partially darkened. Scutellum black. Elytra background colour black, with yellow humeral fascia shortly extending from the scutellum along the suture and along the humeral angle, and usually four to five spots partially connected to the lateral margin; the elytral edge usually yellow with darkened edge; in some specimens, dorsal spots are absent or the marginal yellow stripe is not visible and the elytral margins are entirely black. Ventral side of the prothorax yellow-orange; the pterothorax dark except for the pale mesepimeron. Elytral epipleuron is usually yellow, with the interior border narrowly black. Abdominal disc yellow, except for ventrite 1 largely black. Legs yellow, sometimes the femora are partially black.

67

Morphology. Length 3.8–4.6 mm. Frons 1.5 times as broad as the eye diameter. Eyes partially beneath pronotum, the ocular canthus large and almost rounded. Pronotum anterior border strongly emarginate laterally and straight medially. Prosternal process with long carinae extending to almost to the anterior margin. Elytron with lateral border narrowly explanate; scutellum strongly transverse. Elytral epipleuron without foveae. Mesoventrite with the anterior margin broadly emarginate medially, the posterior margin straight. Abdominal postcoxal line (Fig. 70I) extending to the posterior margin of the ventrite, almost complete laterally. Male terminalia (Figs 70D–G). Female terminalia (Fig. 70H). Measurements (n = 5): TL 3.85–4.65 (4.26 ± 0.28); PL 0.60–0.75 (0.68 ± 0.06); EL 3.25–3.85 (3.56 ± 0.24); PW 2.35– 2.55 (2.46 ± 0.09); EW 3.50–4.10 (3.85 ± 0.23); GD 1.80–2.10 (2.00 ± 0.14). Ratios: PL/PW 0.25–0.29 (0.28 ± 0.02); EL/EW 0.88–0.96 (0.92 ± 0.03); EL/PL 4.86–5.54 (5.25 ± 0.26); EL/ GD 1.71–1.83 (1.78 ± 0.05); TL/EW 1.04–1.15 (1.11 ± 0.04).

Types ‘New Guinea: NE Wau, 1200m, 23.10.1973/ on Myrtaceae: Eugenia alata/ J.L. Gressitt coll’r Wau Ecology Inst. (Bishop)’ (BPBM, Neotype female, here designated)

Specimens examined PAPUA NEW GUINEA: NE Wau, 1200m, Coll’d on Castanopais sp, 3.viii.1973, Kewisef (1, BPBM) NE, Wau, 1200m, 31. vii.1973, N. L. H. Krauss (1, BPBM); NE Wau, 1200m, Castanopsis sp., 31.vii.1973, Tawi Bukam (1, BPBM); New Guinea: NE Morobe District, Ulap, 800–1100m, ix.1968, N. L. H. Krauss (1, BPBM); Mt. Lamington N. E. Papua, 1300–1500ft, C. T. McNamara (2, SAM); Mt. Lamington Dist., Northern Division, Papua, ix.1929, C. T. McNamara (3, AM)

Remarks Iablokoff-Khnzorian (1982) stated there were four specimens from Wau referred to as ‘4 types’ without designating the holotype and the paratypes. He never returned these specimens to the Bishop Museum in Honolulu (J. Boone, pers. comm.). Judging from the vague statement in the original publication, these specimens were probably lost before the book was published. To fix the identity of this species, we have selected a female specimen from the type locality that matches the drawings and original description by Iablokoff-Khnzorian (1982) and designated it as the neotype of Oiocaria samuelsoni (BPBM).

Distribution Endemic to New Guinea.

Distribution of Heterocaria samuelsoni

68

Ladybird Beetles of the Australo-Pacific Region

Heterocaria transversoguttata sp. nov.

Distribution

(Fig. 71)

Endemic to Papua New Guinea.

Diagnosis

Remarks

In addition to the very distinctive male genitalia, Heterocaria transversoguttata is distinguished from other species by the black pronotum bearing an oval yellow area laterally, and the black elytra with large yellow spots connected into transverse fasciae.

The species name refers to the transverse, fasciate pattern on the elytra.

Description

Heteroneda Crotch, 1871: 6. Type species, by subsequent designation of Crotch 1874: 162, Coccinella reticulata Fabricius, 1801 (= H. billardieri Crotch, 1874). Iablokoff-Khnzorian 1982: 538.

Colour pattern (Figs 71D–H). Frons and head appendages yellow. Labrum yellow. Pronotum black, with two yellow oval areas laterally; lateral borders black. Scutellum black. Elytra black, each with five yellow spots forming transverse fasciae (Fig. 71A); the external border and suture black. Ventral side yellow, except for the metaventrite, metanepisternum, external borders of the elytral epipleuron and the central part of abdominal ventrites are black. Legs mostly brown. Morphology. Length 5 mm. Frons 1.7 times wider than the eye diameter. Eyes partially beneath the pronotum. Antennae about twice as long as the frons width. Pronotum ~0.9 times as broad as the elytra at the base. Anterior border almost straight medially; disc with fine and regular punctures about as large as the eye facets. Prosternal process with long parallel carinae extending close to the anterior margin of the prosternum. Elytra broadest in the anterior third, distinctly narrowing posteriorly; lateral borders explanate, lateral margin thickened, weakly upturned near the shoulder; disc with a mixture of larger and smaller punctures, the larger punctures at least twice as large as the pronotal ones and becoming coarser towards the lateral margins; interspaces reticulate but shiny; epipleuron ~1.7 times as broad as metanepisterum. Scutellum 1/11 times as broad as the elytra, with a basal margin as long as the lateral margin. Anterior border of the mesoventrite shallowly and broadly emarginate, with raised carinae connected medially, the posterior margin almost straight. Abdominal postcoxal line approaching the posterior margin, complete laterally. Ventrite 5 with the posterior margin truncate, ventrite 6 with the posterior margin slightly emarginate medially. Male terminalia (Figs 71D–H). Female. Unknown. Measurements (Holotype): TL 5.00; PL 0.90; EL 4.10; PW 2.80; EW 4.40; GD 2.35. Ratios: PL/PW 0.32; EL/EW 0.93; EL/PL 4.56; EL/GD 1.74; TL/EW 1.14.

Specimens examined Holotype: PAPUA NEW GUINEA: Lae km4, Jul 1975, H. Ohlmus (ANIC).

Distribution of Heterocaria transversoguttata

Heteroneda Crotch, 1871 (Figs 72–74)

Diagnostic combination Length 4–6 mm. Elytra moderately to strongly convex, glabrous. Anterior clypeal border straight between the lateral projections. Antenna 11-segmented; slightly shorter than the head capsule, with a weak three-segmented club; terminal antennomere longer than the penultimate, rounded apically. Terminal maxillary palpomere securiform. Pronotal disc evenly convex transversely, with narrowly upturned or slightly thickened external borders. Prothoracic hypomeron without foveae near the anterior angles; prosternal process with distinct carinae extending forward to the middle of the prosternum. Anterior margin of the mesoventrite very deeply emarginate medially. Elytra with lateral margins with complete beads; epipleuron without foveae. Mid and hind femurs do not extend beyond the border of the elytra. Tibial spur formula 0–2–2. Abdominal postcoxal line not recurved, incomplete laterally; an oblique postcoxal line absent. Penis consists of a single basal sclerite and an additional, largely membranous, apical pouch; tegmen stout, penis guide does not extend beyond the parameres apically, strongly bent towards the parameres in lateral view. Ovipositor with coxites of a clubhandle type, infundibulum absent; spermatheca C-shaped with a narrowed ramus usually indistinct, nodulus slim.

Description Head strongly withdrawn into prothorax. Eyes largely beneath the anterior margin of the pronotum, the interior orbits strongly divergent anteriorly. Frons narrow, about as broad as the eye diameter. Antenna 11-segmented, as broad as the head capsule, scape slightly enlarged interiorly, club slightly serrate, the terminal antennomere is rounded apically, longer than the penultimate. Anterior clypeal margin straight between two distinct lateral projections. Terminal maxillary palpomere weakly securiform. Mandible bifid apically, the molar part with a basal tooth. Prothorax. Pronotum weakly convex, the lateral border area broadly upturned; the anterior angle and posterior angle blunt. Prothoracic hypomeron concave, without distinct foveae. Prosternum in front of the procoxae as long as the prosternal process; the prosternal process 0.3 times as broad as the procoxae, the surface with carinae extending to the middle of the prosternum, slightly convergent anteriorly. Pterothorax. Mesoventrite as long as broad, ~0.6 times as broad as the mid-coxal diameter; the anterior margin strongly emarginated medially, with a complete raised ridge; mesal surface with clear articulation with the prosternal process; posterior margin truncate. Postcoxal line on the metaventrite joined medially, complete laterally; discrimen complete. Elytra much broader than the pronotum at the base, moderately convex, finely punctuated, narrowly explanate laterally. Elytral

VII. Review of the Australo-Pacific Coccinellini

epipleuron moderately broad, slightly inclined, without foveae. Apex of mid and hind tibiae with two spurs apically; tarsal claws with subquadrate basal teeth. Abdomen with six ventrites in both sexes. Postcoxal lines connected medially, complete laterally, not recurved; an oblique postcoxal line absent. Posterior margin of ventrite 5 truncate in both sexes. Male terminalia. Penis consists of a single sclerite, and a largely membranous apical pouch bearing multiple sclerotisations. Female terminalia. Spermatheca strongly curved, surface smooth, ramus indistinct, the nodulus narrower than the cornu in diameter. Sperm duct short; infundibulum absent; coxites small.

Biology Heteroneda billardieri is regarded as a generalist aphid feeder, but it has also been found to feed on planthoppers and psyllids in Malaysia and the Philippines (Barcos et al. 2014).

69

Distribution In addition to the three Papuan species treated here, this genus includes the widely distributed H. billardieri whose distribution extends from India, Singapore and Myanmar to Indonesia and the Philippines.

Remarks In most cases, the very similar species of Harmonia, with their reticulate elytral patterns, can be distinguished from Heteroneda by the development of the elytral cells. In Heteroneda, the first, most lateral cell is shorter than first dorsal cell, while in Harmonia both cells are approximately the same length, or the most lateral one is slightly longer than first dorsal cell (Figs 44C, 73C). Otherwise both genera are very distinct in having different antennae, mesoventrites and genitalia in both sexes.

Key to species of the genus Heteroneda from the Australo-Pacific Region 1. – 2(1). – 3(1). –

Elytra orange or yellow, with black reticulate pattern (Figs 73, 74) ...................................................................................................2 Elytra black with orange or yellow spots ............................................................................................................................................3 Larger species (length 6.0–7.6 mm); elytron with four cells along the lateral margin (Fig. 74A) [typical form] ........ H. principalis [part] Smaller species (length 5.2–5.7 mm); elytron with three cells along the lateral margin (Fig. 73A) ................................H. mimetica Smaller species (length 3.8–6.3 mm); elytron with humeral and apical orange spot, or only with apical spot reaching the apex of the elytron (Fig. 72A) ...................................................................................................................................................... H. grata Larger species (length 6.0–7.6 mm) elytron rarely completely black, usually black with two to four orange or red spots along the margins (Fig. 74C) or with a single apical spot distinctly separated from the apex (Fig. 74D) [rare forms] ................................ ��������������������������������������������������������������������������������������������������������������������������������������������������������������������� H. principalis [part]

Heteroneda grata (Iablokoff-Khnzorian, 1982) comb. nov. (Fig. 72)

Lemnia grata Iablokoff-Khnzorian, 1982: 245. Type locality: New Guinea, Wau.

Diagnosis Heteroneda grata is easily distinguished from other species in the genus by its black elytra bearing reddish spots. A rare, melanic forms of H. principalis (Fig. 74D) are quite similar to H. grata, but can be distinguished by an apical reddish spot not extending to the apicosutural angle of the elytra.

Description Colour pattern (Figs 72A–D). Head and head appendages orange yellow. Pronotum usually yellow, with two small triangular black patches at the base that are often enlarged to a broad trapezoidal medial area. Scutellum black. Elytron black, with apical or apical and humeral reddish patches. Prothoracic hypomeron and prosternum yellow, the pterothorax black except for the yellow mesanepisternum and mesepimeron, elytral epipleura usually black, with anterior and posterior parts yellow-reddish. Abdomen black, with external borders yellow. Anterior legs largely brown, middle and hind legs largely black. Morphology. Length 3.8–6.3 mm. Eyes facetted relatively coarsely, partially beneath the anterior margin of the pronotum. Antenna long, ~0.8 times as broad as the head capsule. Pronotum weakly convex, the lateral margin arcuate. Prosternum with parallel carinae almost reaching the anterior border. Anterior margin of the mesoventrite strongly emarginate medially, the

posterior margin slightly arcuate. Abdominal postcoxal line (Fig. 72K) approaching the posterior margin of ventrite 1, incomplete laterally. Posterior margin of ventrite 5 truncate in both sexes. Male terminalia (Figs 72E–H). Female terminalia (Figs 72I,J). Measurements (n = 12): TL 3.8–6.30 (5.39 ± 0.49); PL 0.65–1.95 (0.96 ± 0.20); EL 2.90–4.95 (4.34 ± 0.37); PW 2.00– 3.00 (2.67 ± 0.17); EW 3.80–5.00 (4.40 ± 0.33); GD 2.00–2.70 (2.39 ± 0.18). Ratios: PL/PW 0.24–0.67 (0.34 ± 0.03); EL/EW 3.22–6.86 (4.85 ± 0.68); TL/EW 1.45–2.00 (1.85 ± 0.13); EL/ GD 1.00–1.38(1.16 ± 0.03).

Types L. grata: Papua, New Guinea Morobe distr. Wau, 1150.20.6.1974; A. D. HART Coll. R. W. E. I (BPBM, holotype).

Species examined INDONESIA: Western New Guinea: Mapia 1480m. 30 vii, 1996 leg. Schule & Stuben (1, NHMS); Nabire n. Mapia, km177, Ugida, 1400m. 29 vii, 1996 leg. Schule & Stuben (1, NHMS); 10km SW Nabire, 30. ix.1990 leg. M. Balke. (14, ZMB); Japen Seruin, Amberdiru 5.viii.1996 1000m, Schule & Stuben (11, NHMS); Neth. Waris, S. Of Hollandia, h 50–500m, 16– 23.viii.1958. T. C. Maa (1, BPBM); Japen, SSE Sumberbaba, Dawai R. X. 1962. N. Wilson Collector (1, BPBM); Numfoor I., ii-1945, H.H. Blakemore (2, CAS). PAPUA NEW GUINEA: NE Wau, Morobe Distr., 1200m, 9 ix, 1972, J. L. Gressitt (1, BPBM); Wau, 4000 ft., 24–30.vi.1974, H. F. Howden (1,CAS); NE Wau, 1200–1450m, 18.vi.1968, N. L. H. Krauss (1, BPBM); (NE) Wau, Morobe Distr., 1300m,

70

Ladybird Beetles of the Australo-Pacific Region

Distribution of Heteroneda grata

Distribution of Heteroneda mimetica

27.vii.1961, J. Sedlacek (1, BPBM); NE W. Highlands, Baiyer R. 1200m, 1.viii.1972, J. L. Gressitt (1, BPBM); Mt. Lamington Dis, Northern Division, Papua, vii.1927, C. T. MacNamara (8, AM).

2.95 (2.78 ± 0.10); EW 4.30–5.00 (4.70 ± 0.20); GD 2.20–2.60 (2.44 ± 0.12). Ratios: PL/PW 0.34–0.41 (0.36 ± 0.02); EL/EW 0.87–1.02 (0.92 ± 0.04); EL/PL 3.73–4.56 (4.32 ± 0.23); EL/ GD 1.64–1.96 (1.78 ± 0.09); TL/EW 1.08–1.26 (1.15 ± 0.06).

Distribution Known only from New Guinea.

Heteroneda mimetica Chazeau, 1990 (Fig. 73)

Heteroneda mimetica Chazeau, 1990: 162. Type locality: Papua New Guinea, Port Moresby.

Diagnosis This species is very similar to some sympatric Harmonia species, including Harmonia decussata, H. nigromarginata and H. testudinaria, but it differs in having a deeply emarginate anterior margin of the mesoventrite, and an elongate and rounded apical antennomere. It can be distinguished from Heteroneda billardieri by the pronotum not having a median black stripe, always present in Heteroneda mimetica.

Description Colour pattern. Frons and head appendages yellow. Pronotum yellow, with two longitudinal stripes usually not reaching the anterior border. Scutellum black. Elytra yellow or orange, with longitudinal and transverse stripes as in Figs 73A–C, the external borders and suture black. Ventral side completely yellow, except for the black external borders of the elytral epipleura. Morphology. Length 5.2–5.7 mm. Frons flat, without distinct punctures, 1.1–1.3 times as broad as the eye width. Prothorax distinctly narrower than the elytra at the bases; the anterior margin emarginate; the anterior angles projecting; the lateral margins arcuate; the disc with shallowly impressed punctures, smaller than the eye facets; interspaces shiny. Surface of prosternal process with parallel longitudinal carinae, extending to the middle of the prosternum. Scutellum 1/12 times the width of elytra at humeri. Elytral margins narrowly explanate; epipleuron ~2.5 times as broad as the metanepisternum. Abdominal postcoxal lines separated medially, incomplete laterally (Fig. 73J). Posterior margin of ventrite 5 truncate and hairy in males, slightly arcuate in females. Male terminalia (Figs 73D–G). Female terminalia (Figs 73H,I). Spermatheca slim, strongly curved and C-shaped, the nodulus and ramus distinct, the sperm duct very short and slightly broadened. Measurements (n = 11): TL 5.20–5.70 (5.40 ± 0.16); PL 0.90–1.10 (1.00 ± 0.06); EL 4.10–4.60 (4.33 ± 0.18); PW 2.60–

Types H. mimetica: ‘Port Moresby, Papua Home Garden, 28–I-1956, coll. W. Brunhorst/ Heteroneda mimetica n. sp., Holotype’ (BMNH, holotype).

Specimens examined INDONESIA: Western New Guinea: Jayapura Sentani, Cyclop Mts., 19–21.9.1990, 300m, Leg. A. Riedel (1, NHMS); Ayam 60km Neot Agats, 18–27.xii.1976, J. C. Wright (1, BPBM); Iran Jaya: i-iii. 1944, M. Marquis (1, CAS). KAI ISLANDS: Key Inseln (1, MNHN); Key, Kraatz (1, DEI); Little Kei (H. Kuhn) (4, MIIZ). PAPUA NEW GUINEA: NE Bulolo, 700m 6.xi.1969, J. Sedlacek (1, BPBM); N. Bulolo, 700m, 31.xii.1969 (2, BPBM); Northern District Bulolo, 700m, 29.xii.1970. B. B. Lowery (1, ANIC); Busa River, E. of Lae, N. G., 24.viii.1957, Holland-Munroe (1, CNC); NE Wau, 1200m, 9.vii.1965, J. & M. Sedlacek (1, BPBM); NE Wau, Morobe Distr., 900m, 26.viii.1963, H. Clissold (1, BPBM); NE Wau, 1200m, 13– 15.ix.1965 (1, BPBM); NE Wau, 1200–1300m, 15.viii.1965 (1, BPBM); NE Wau, Morobe Distr., 1200m, 1–15.v.1962, J. Sedlacek (1, BPBM); NE Wau, 1200m, 23.ii.1966, J. Sedlacek (1, BPBM); Port Moresby, (Mt. Lawes, 1300ft), 5.iii-12.v.1963, W. W. Brandt (2, ANIC); Port Moresby, (Mt. Lawes, 1300ft), 3–12.v.1963, W. W. Brandt (6, ANIC); Morobe District, Wau. B. P. Bishop Museum Field Station 1.x.1969, James E. Tobler (1, CAS); Brown RIV. 30mi. N. Pt Moresby, 14–15.vii.1974, H. Howden (1, ANIC); SE Weam, 9m 20.v-6.vii.1964 (1. BPBM); NE Bulolo R., 480m, 23.iv.1969(1, BPBM); Daru Papua, Mountain of Fly R., vii.1941 (1, CAS); Brown Riv., 30mi. N. Pt Moresby, 14–15. vii.1974, H. Howden (1, CAS)

Distribution Western New Guinea and Kai Islands, Indonesia; Papua New Guinea.

Heteroneda principalis (Weise, 1895) comb. nov. (Fig. 74)

Callineda principalis Weise, 1895: 134. Type locality: Kei Island. Korschefsky 1932: 283. Lemnia (Microlemnia) gressitti Iablokoff-Khnzorian, 1982: 264. Type locality: New Guinea. Syn. nov.

VII. Review of the Australo-Pacific Coccinellini

71

Diagnosis The typical reticulate form of Heteroneda principalis can be distinguished from similar species and genera in having four elytral cells along the external margin, and the first dorsal cell along the suture is very long and expanded posteriorly (Fig. 74A). The melanic forms (Figs 74C,D) are distinguished from Heteroneda grata by their larger size, and the apical macula not extending to the apicosutural angle.

Description Colour pattern (Figs 74A–F). Head and appendages yellow. Pronotum yellow, with two longitudinal stripes, the area between the stripes usually darker than the lateral areas. Scutellum yellowish. Elytra yellow or orange, with a black suture and external margins and black reticulate colour pattern (Figs 74A–D). In some specimens, elytra are mostly black with reddish spots limited to the humeri and apices (Fig. 74C), or missing altogether (Fig. 74D). Ventral side yellowish, except for the external borders of elytral epipleura black. Legs yellow. Morphology. Length 6.0–7.6 mm. Frons 1.2–1.4 times broader than the eye. Eyes partially obscured beneath the anterior margin of the pronotum. Pronotum weakly convex, explanate laterally. Anterior margin slightly arcuate medially. Apex of prosternal process triangular and extending behind the procoxae, the surface with carinae extending beyond the middle of the prosternum. Prothoracic hypomeron without well-defined foveae. Elytra slightly broader than the pronotum. Scutellum with a basal margin as long as the lateral margin. Anterior margin of the mesoventrite with a strong medial notch, the posterior margin slightly arcuate. Elytral epipleuron about twice as wide as the metanepisternum, with a shallow concavity. Abdominal postcoxal line (Fig. 74M) incomplete laterally; the posterior margin of ventrite 5 truncate in both sexes. Male terminalia (Figs 74G–K). Penis strongly curved at base with additional sclerite at the apex. Female terminalia (Fig. 74L). Spermatheca small; cornu strongly C-shaped; sperm duct slim, originating in the middle of the bursa. Measurements (n = 30): TL 6.10–7.60 (6.65 ± 0.38); PL 0.95–1.50 (1.15 ± 0.14); EL 4.90–6.00 (5.44 ± 0.28); PW 3.20– 3.90 (3.53 ± 0.18); EW 5.10–6.50 (5.78 ± 0.30); GD 2.70–3.70 (3.05 ± 0.20). Ratios: PL/PW 0.25–0.42 (0.33 ± 0.04; EL/EW 0.85–1.00 (0.94 ± 0.04); EL/PL 4.00–5.50 (4.80 ± 0.47); EL/ GD 1.57–1.94 (1.79 ± 0.09); TL/EW 1.06–1.27 (1.15 ± 0.05).

Types C. principalis: ‘Key-Ins’ (ZMB, Neotype, here designated). L. gressitti: New Guinea NW. Wau. 13.8.1974, 1200m. J.L. Gressitt (BPBM, holotype).

Specimens examined INDONESIA: Western New Guinea: Star Mountains, ABMISIBII, 1950–2200M, 4.38S-140.33E, 29.i-9.ii.2005, T. Lackner (1, NBCL); G. den. Hoed Ned. NW. Guinea, R. T. Simon, 4.viii.1958 (1, NCBL). KAI ISLANDS: Ins. Kei, (Planten) (MNHN). PAPUA NEW GUINEA: Madang Dist., Finisterre Mts., Damanti 3.500 ft, 2–11.x.1964, M.E. Bacchus (2, BMNH); Wau: Golden Ridges, iv.1958, E. Berg (1, BMNH); Weam, viii.1976, H. Ohlmus (3, ANIC); NE Wau, Morobe Dist., 1200m, 6.xi.1966, J. & M. Sedlacek (1, BPBM); Morobe Dist., Wau, M. V. Light trap, 9.ix.1972, G. G. E. Scudder (1, BPBM);

Distribution of Heteroneda principalis

Wau, 4000 ft, 24–30.vi.1974, H.F. Howden (2, BMNH); Bosavi, x.1975, H. Ohlmus (7, ANIC); NE Mt. Kaindi, 16km SW of Wau, 1500–1800m, viii.1963, H. Clissold & P. J. Shanahan (2, ANIC); Goroka, Mt. Hagen, 1966, P. Voss (1, NHMB).

Distribution This species is found on the island of New Guinea, and the Kai Islands, Indonesia.

Remarks The generic placement of this variable species is not well established, as the male genitalia, in particular the broad penis guide and the penis with its sclerotised apical element, departs from the typical Heteroneda. The fovea on the prothoracic hypomeron is also relatively weakly defined. We were not able to locate the type material of Callineda principalis and to stabilise the nomenclature and identity of this variable species we have designated the Neotype from two specimens bearing lanbel ‘Key-Ins’ and identified as Callineda principalis in the Weise collection in ZMB Berlin. Iablokoff-Khnzorian (1982) has ignored Weise’s species and described it again under the name Lemnia (Microlemnia) gressitti.

Hippodamia Chevrolat, 1836 (Fig. 75)

Hippodamia Chevrolat in Dejean, 1836: 432. Type species, designated by Crotch 1874: 94, Coccinella tredecimpunctata Linnaeus, 1758. Adonia Mulsant, 1846: 39. Type species, by monotypy, Coccinella mutabilis Scriba, 1791.

References: Iablokoff-Khnzorian 1982: 308; Gordon 1985: 706. Diagnostic combination Length 4.2–5.2 mm, elongate-oval, elytra moderately convex, glabrous. Anterior clypeal border straight between the lateral projections. Antenna 11-segmented; slightly shorter than the head capsule with moderately compact three-segmented club. Terminal maxillary palpomere strongly securiform. Pronotal disc evenly convex transversely, to narrowly upturned external borders; pronotal base with an entire margin. Prothoracic hypomeron without a fovea near the anterior angles; the prosternal process very narrow, without distinct carinae. Anterior margin of the mesoventrite straight and faintly elevated. Elytral margin with narrow raised margins; epipleuron not foveate. Tibial spur formula 0–2–2; tarsal claws split apically. Abdominal postcoxal line recurved and complete; an oblique postcoxal line absent.

72

Ladybird Beetles of the Australo-Pacific Region

Description

Diagnosis

Body elongate-oval, moderately convex, glabrous; apices of femora visible dorsally. Head exposed dorsally, antenna as long as the head capsule, 11-segmented; scape normal, pedicel distinctly narrower than the scape; antennal club compact, three-segmented, the penultimate antennomere distinctly shorter than the terminal segment, the terminal antennomere subquadrate, apically truncate; clypeus short, transverse, the anterior clypeal margin straight. Labrum entirely exposed. Mandible bifid apically, the molar part with a basal tooth. Prothorax. Base of the pronotum with a bead. Intercoxal process of the prosternum narrow, usually weakly convex, without carinae. Prothoracic hypomeron without delimited foveae. Pterothorax. Elytra at the base much broader than the pronotum, the lateral margin very narrow but entirely visible from above. Mesoventrite straight anteriorly without a raised border; the mesal surface without clear articulation with the prosternal process. Meso-metaventral process narrow, 0.6 times as wide as the coxal diameter. Elytral epipleuron incomplete apically. Apex of the middle and hind tibiae each with two spurs. Tarsal claw is cleft. Abdomen with six ventrites in both sexes; ventrite 1 as long as, or slightly longer than, ventrite 2. Abdominal postcoxal line meeting at the middle, recurved (Fig. 75K); the oblique postcoxal line is absent. Male terminalia (Figs 75E–G). Penis stout, consisting of a bent and complex single sclerite (Fig. 75E). Female terminalia (Figs 75I,J). Spermatheca distinctly curved, ramus and nodulus present, infundibulum absent, coxal plate elongate, the stylus distinct.

Hippodamia variegata is externally most similar to Coccinella undecimpunctata, but can be immediately distinguished by its apically cleft tarsal claws, a narrow and smooth prosternal process, and the completely recurved abdominal postcoxal lines.

Biology Members of the genus Hippodamia are general aphid predators (Savoiskaya 1983; Gordon 1985). Franzmann (2002) lists 12 aphid species and a psyllid as food sources of the single introduced Australian representative, H. variegata. Ślipiński (2007) illustrated the larva of H. variegata.

Distribution Hippodamia is essentially a Holarctic genus of ~35 species, widely distributed in the Palaearctic Region, North America and Mexico (Iablokoff-Khnzorian 1982; Gordon 1985). One species, H. variegata, has been introduced to many countries.

Description Colour pattern. Labrum, clypeus and frons in males are anteriorly yellow, black posteriorly; in females the black area variously extends medially reaching the anterior margin of the clypeus, usually leaving only yellow areas adjacent to eyes. Pronotum black with the anterior and lateral margins narrowly yellow; the disc with a medial yellow stripe and two smaller spots. Scutellum black. Elytron yellow or orange, with a black scutellar spot and four extra smaller or larger spots (Fig. 75A); ventral surface almost all black except for the yellow mesepimeron and metepimeron; legs mostly black, procoxae paler. Morphology. Length 3.7–5.2 mm. Body elongate-oval and weakly convex. Frons flat and sparsely punctate, ~2.7–3.0 times as broad as the eye. Pronotum weakly convex, regularly punctate; the interspaces mostly smooth and shiny. Scutellum triangular and small, ~1/14–1/15 the width of the elytra at the humeri. Elytral surfaces punctured with reticulate, weakly shiny interspaces. Epipleuron very narrow, 1.1–1.5 times as broad as the metanepisternum. Basal tarsomere of front and mid tarsi expanded in males. Male terminalia (Figs 75E–H). Female terminalia (Figs 75I,J). Measurements (n = 5): TL 3.70–5.15; PL 0.70–1.05; EL 3.00–4.05; PW 1.60–2.30; EW 2.30–3.32; GD 0.8–1.5. Ratios: PL/PW 0.42–0.51; EL/EW 1.20–1.34; EL/PL 3.95–4.20; EL/ GD 3.3–3.7; TL/EW 1.51–1.70.

Specimens examined AUSTRALIA: Queensland: Gatton, 9.i.2001, B.A. Frantzmann (ANIC). Western Australia: Perth, Brigadon, 1.vii.2003, R. Horbury (ANIC); Australian Capital Territory: Campbell, 3.xii.2002, M. Carver (ANIC); Watson, 3.xi.2002, K. Pullen (ANIC); Belconnen markets, 24.x.2002, A. Ślipiński (ANIC); Acton, ANU campus, 6.xi.2002, A. Ślipiński (ANIC); Fyshwick Markets, 8.xi.2002, A. Calder (ANIC).

Remarks Adonia Mulsant is often regarded as a separate genus or subgenus within Hippodamia (Iablokoff-Khnzorian 1982; Gordon 1985). It differs from other Hippodamia by having a margin on the base of the pronotum, dilated tarsi in males, and a recessed infundibulum in females.

Hippodamia variegata (Goeze, 1777) (Fig. 75)

Coccinella variegata Goeze, 1777: 247. Type locality: Paris. Adonia variegata: Mulsant 1846: 39. Hippodamia (Adonia) variegata: Iablokoff-Khnzorian 1982: 326; Ślipiński 2007: 164.

Distribution of Hippodamia variegata

VII. Review of the Australo-Pacific Coccinellini

Distribution This is a widely distributed Palaearctic species extending from Europe and northern Africa and to Korean Peninsula. It has been introduced, or made its way naturally, to southern Africa (South Africa, Kenya), Asia (India, China), North America and South America (Chile). In Australia, it was first found in Queensland (Franzmann 2002), but it quickly colonised new areas and now it occurs throughout the continent, including in Western Australia, South Australia and Tasmania.

Hysia Mulsant, 1850 (Fig. 76)

Hysia Mulsant, 1850: 36, 47. Type species by monotypy, Coccinella endomycina Boisduval, 1835.

References: Iablokoff-Khnzorian 1982: 453. Diagnostic combination

Length 5.2–7.0 mm, elongate-oval, elytra weakly convex, glabrous. Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly longer than the head capsule with moderately compact three-segmented club; the terminal antennomere truncate. Terminal maxillary palpomere securiform. Pronotal disc flat; pronotal base without a marginal bead. Prothoracic hypomeron without foveae near the anterior angles; prosternal process short, without distinct carinae. Anterior margin of the mesoventrite straight. Elytral epipleuron narrow, not foveate. Tibial apices visible dorsally; tibial spurs formula 0–2–2; tarsal claws with a basal tooth. Abdominal postcoxal line recurved and incomplete; an oblique postcoxal line absent. Penis consists of a single sclerite. Infundibulum sclerotised, tubular. Spermatheca C-shaped, with a poorly developed ramus and nodulus.

Description Medium size, weakly convex and strongly elongated, glabrous; the femora extend beyond the elytra and are visible from above; the pygidium often partially visible dorsally. Head transverse, almost entirely visible. Clypeus straight between lateral projections. Eyes large, completely exposed, the interior margin slightly convergent. Antenna 11-segmented, longer than the width of the head capsule; the antennal insertion very close to the inner eye margin, distinctly covered from above; the scape twice as long as the pedicel, slightly arcuate; the antennal club loosely articulated, the terminal antennomere subquadrate and truncate apically. Terminal maxillary palp securiform. Prothorax. Pronotum weakly convex; the anterior margin straight or arcuate; the lateral border evenly arcuate, narrowly explanate. Prosternal process broad, not exceeding beyond the posterior margin of the coxae; the surfaces flat, without carinae. Prothoracic hypomeron without delimited foveae. Pterothorax. Mesoventrite small, with the anterior border straight or faintly arcuate, the mesal surface without a distinct fossa to receive the prosternal process. Mesepimeron strongly elongate. Metaventrite with an incomplete discrimen; the postcoxal line complete laterally. Scutellum very small. Elytra weakly convex, slightly broader than the pronotum at the base; the lateral borders narrowly explanate. Elytral epipleuron flat, narrow and incomplete to the apex, without delimited foveae. Legs long, femorae partially exposed dorsally; apex of the middle and hind tibiae each with two spurs; tarsal claws long and with a small basal tooth.

73

Abdomen with six ventrites in both sexes. Abdominal process pointed apically. Abdominal postcoxal line recurved and incomplete laterally (Fig. 76J); the oblique postcoxal line absent. The terminal tergite, or ‘pygidium’, usually partially visible dorsally. Male terminalia (Fig. 76E). Penis guide slender and distinctly narrowing apically; the penis stout, consisting of a single sclerite with an apical pouch. Female terminalia (Fig. 76I). Spermatheca lightly sclerotised, with cornu curved apically; the ramus and nodulus indistinct; the spermathecal accessory gland adjacent to the sperm duct. Sperm duct very short; infundibulum present, flared at both ends.

Biology Unknown.

Remarks Hysia is a very distinctive genus due to its general appearance, recurved abdominal postcoxal lines and long legs, with most of the femora visible externally. It is monotypic.

Hysia endomycina (Boisduval, 1835) (Fig. 76)

Coccinella endomycina Boisduval, 1835: 603. Type locality: New Guinea. Hysia endomycina: Mulsant 1850: 36, 47; 1866: 33; Crotch 1874: 99; Korschefsky 1932: 438; Iablokoff-Khnzorian 1982: 453.

Description Colour pattern (Figs 76A–C). Clypeus and labrum yellow; frons dark with long white hairs; antennae slightly longer than the head capsule, largely yellow, with a dark club. Maxillary palps yellow. Pronotum black, with two yellow subquadrate sports in the anterolateral corners. Prosternum usually black, with a brown anterior margin; prothoracic hypomeron black, but yellowish anteriorly; scutellum black. Elytra orange, with three black transverse fasciae of variable sizes, with the third one located at the elytral apices; epipleura yellow. Pterothorax complete black ventrally. Legs slim and long, usually black. Morphology. Length 5.3–7.0 mm. Eyes completely exposed, finely facetted, convex. Frons 1.4–1.6 times broader than the eye diameter, weakly concave; punctured and setose; interspaces densely reticulate and weakly shiny. Pronotum widest posteriorly; the lateral margins arcuate and upturned. Pronotal disc densely punctured, punctures larger than the eye facets, 1.5–2.5 diameters apart. Scutellum triangular, width ~1/12 of the elytra at the humeri. Elytra slightly broader than the pronotum; the border of the lateral margins narrowly explanate, margins slightly thickened. Disc with irregular punctures; the interspaces reticulate and wrinkled. Abdominal postcoxal line (Fig. 76J) extending to the middle of the ventrite, recurved and incomplete laterally. Posterior margin of ventrite 5 truncate in males and slightly arcuate medially in females. Male terminalia (Figs 76D–G). Female terminalia (Figs 76H,I). Measurements (n = 21): TL 5.30–7.00 (6.28 ± 0.53); PL 1.00–1.50 (1.20 ± 0.13); EL 3.70–5.00 (4.57 ± 0.36); PW 2.15– 2.65 (2.45 ± 0.15); EW 3.10–4.00 (3.63 ± 0.23); GD 1.75–2.10 (1.98 ± 0.11). Ratios: PL/PW 0.39–0.61 (0.49 ± 0.05); EL/EW 1.10–1.37 (1.26 ± 0.07); EL/PL 3.13–4.55 (3.85 ± 0.43); EL/ GD 2.10–2.50 (2.30 ± 0.12); TL/EW 1.51–1.92 (1.73 ± 0.10).

74

Ladybird Beetles of the Australo-Pacific Region

Distribution of Hysia endomycina

Specimens examined

Diagnostic combination

PAPUA NEW GUINEA: E. Highlands Dist., Groka, 29.v.1971, R. L. Rivers (1, ANIC); Kondiu Mai, 1964, P. Cl. Voss (8, NHMS); Kainantu, 27.v.1979, W. G. Ullrich (1, MHNG); NE: E. Sepik Prov., Wom, 10m, 24.xi.1979, W. C. Gagne (1, BPBM); SE: S Highland Dist., Pimaga to Lake Kutubu, Grasses by Lake, 1000m, 6.ii.1978, J. L. Gressitt (6, BPBM); Norekori, 15km S Kainantu, 9.v.1973 (1, ANIC); Kondiu Mai, 1964, P. Cl. Voss (1, NHMS); Morobe Dist., Gusap, Markham Valley, 90 m, W of Lae, 27–30.i.1965, M.E. Bacchus (BMNH). INDONESIA: SERAM: Ceram: Umg. Wahai, 12.2., x.1989, Schodl (1, NHMW).

Length 3.7–6.2 mm; dorsum flat or weakly convex, glabrous. Elytral colour pattern yellow and black (Figs 77, 78). Antennae distinctly longer than the head width, with a loose club. Anterior clypeal margin arcuate, with lateral projections reduced or absent. Mandible bifid apically, with microdenticulation along the incisor edge, the molar part with a basal tooth. Terminal maxillary palpomere strongly securiform. Prothoracic hypomeron without a fovea. Prosternal process short, with long carinae extending forward to almost the anterior margin of the prosternum. Anterior border of the mesoventrite straight. Elytral margins upturned, but without a marginal bead; epipleuron without foveae. Tibial formula 0–0–0. Tarsal claws appendiculate. Abdominal postcoxal line not recurved; oblique postcoxal line absent. Penis consists of a single sclerite. Infundibulum present; spermatheca not sclerotised.

Distribution The island of New Guinea, and Maluku and Sulawesi, Indonesia.

Illeis Mulsant, 1850 (Figs 77–80)

Psyllobora (Illeis) Mulsant, 1850: 1026. Type species by monotypy, Psyllobora galbula Mulsant, 1850. Leptothea Weise, 1898: 227. Type species by original designation, Psyllobora galbula Mulsant, 1850. Synonymised by Timberlake 1943: 42. Illeis (Leptothea): Iablokoff-Khnzorian 1979: 63. Illeis Mulsant: Crotch 1871: 4; Korschefsky 1932: 558; Timberlake 1943: 42; Iablokoff-Khnzorian 1979: 63; 1982: 282. Egleis Mulsant, 1850: 167. Type species, designated by Kovář 2007: 598, Coccinella cincta Fabricius, 1792 (junior homonym). Synonymised by Timberlake 1943: 42. Kiiro Kitano, 2014: 490 (replacement name for Egleis Mulsant).

References: Bielawski 1961a; Iablokoff-Khnzorian 1982: 282; 1984a: 216; Pope 1989: 639.

Description Body broadly oval, moderately convex, dorsum glabrous. Head. Eyes large, coarsely facetted, usually partially covered by the anterior margin of the pronotum. Frons narrow, usually as broad as the eye width. Antennae much longer than the head width; club loosely articulated, with the apical segment elongate-oval (Fig. 77H). Clypeus short, transverse, the anterior clypeal margin arcuate, without distinct lateral projections. Mandible bifid apically, with denticulation along the incisor edge (Fig. 11M), prostheca multi-layered, the molar part with a basal tooth and brush of setae. Terminal maxillary palpomere very strongly securiform (Fig. 77I). Prothorax. Pronotum weaky convex or flat; the anterior border almost straight, without a depression; the lateral margins arcuate; the hind border without a margin. Prosternum short. Prosternal process does not exceed the posterior margin of the coxae, the surface with longitudinal carinae, slightly convergent

VII. Review of the Australo-Pacific Coccinellini

anteriorly. Prothoracic hypomeron without delimited foveae. Pterothorax. Mesoventrite with a straight anterior margin. Postcoxal lines of the metaventrite joined medially, complete laterally. Elytra much broader than the pronotum at the base, finely punctured; lateral margins distinct and narrowly explanate, without a marginal bead. Elytral epipleuron complete to the apex, flat or faintly raised, epipleural foveae absent. Scutellum triangular, ~1/12 times as broad as the elytra at the base. Tibiae without apical spurs; tarsal claws in both sexes with a large subquadrate basal tooth. Apices of hind femurs reaching far beyond the outer margin of the elytron. Abdomen with six ventrites in both sexes. Abdominal process rounded apically; abdominal postcoxal line separate medially, not recurved, incomplete laterally (Fig. 77J); the oblique postcoxal line absent. Posterior margin of ventrite 5 shallowly emarginate in males, strongly arcuate in females. Male terminalia. Penis stout, consisting of a single sclerite; tegmen with parameres slightly divergent in ventral view; the penis guide bent apically. Female terminalia. Spermatheca small, curved and

75

distinctly enlarged apically, the ramus and nodulus weakly developed. The sperm duct is slim and short; the apical part of bursa slightly sclerotised; infundibulum present. Coxites shaped like a club handle, styli strongly reduced and barely visible.

Biology Illeis species are exclusively mycophagous. Adults and larvae of I. galbula in Australia were found feeding exclusively on spores and hyphae produced by species of Oidium (Erysiphaceae) infecting various plants, including cucurbits, crepe myrtle (Lagerstromia sp.) and Lonicera fragrantissima (see Anderson 1981, 1982).

Distribution Iablokoff-Khnzorian (1982) listed 13 species of Illeis distributed in the Oriental and Australo-Pacific regions, with the majority of species known from continental Asia.

Key to the Australo-Pacific species of the genus Illeis 1. – 2(1). – 3(2). –

Elytra with a black and yellow colour pattern (Figs 78A–D) ............................................................................................... I. galbula Elytra yellow without spots ................................................................................................................................................................. 2 Pronotum with three separate black spots near the base (Fig. 79A); eyes separated by less than the eye width .................I. gressitti Pronotum with a solid black area at the base, sometimes extending anteriorly; eyes separated by at least 1.3 times the eye width (Fig. 77A) ..................................................................................................................................................................................... 3 Pronotum with lateral margins always dark posteriorly; frons about twice the eye width; male genitalia as in Fig. 77A ....... I. flava Pronotum with lateral margins entirely yellow; frons ~1.5 times the eye width; male genitalia as in Fig. 80D .....................I. wyliei

Illeis flava Pope, 1989 (Fig. 77)

Illeis flava Pope, 1989: 640. Type locality: Australia: New South Wales, Cascade.

Diagnosis

Male terminalia (Figs 77D–G). Female terminalia. Not examined. Measurements (n = 5): TL 4.40–5.00 (4.82 ± 0.25); PL 0.80–1.00 (0.90 ± 0.07); EL 3.50–4.10(3.90 ± 0.23); PW 2.10– 2.50 (2.26 ± 0.17); EW 3.60–4.10 (3.76 ± 0.23); GD 1.90–2.10 (2.00 ± 0.07). Ratios: PL/PW 0.38–0.43 (0.40 ± 0.02); EL/EW 0.97–1.11(1.04 ± 0.06); EL/GD 1.84–2.05 (1.95 ± 0.08); TL/ EW 1.22–1.36 (1.28 ± 0.06).

Illeis flava is easily distinguished from I. galbula by its uniform elytral colouration and from the Papuan I. wyliei by its broader frons and distinctive male genitalia.

Types

Description

Illeis flava: Dorrigo, N. S. Wales/P-Type Illeis flava sp.n./R. D. Pope det. 1989 (ANIC, SAM, BMNH, paratypes).

Colour pattern (Figs 77A–C). Head and sides of pronotum white-cream, middle part of the pronotum with a black trapezoidal marking, sometimes extending to the lateral margins, lateral margins always dark in the posterior half. Scutellum black. Elytra yellow. Ventral side brownish-black, except for the pale prothoracic hypomeron and elytral epipleura. Legs brown or black. Morphology. Length 4.4–5.0 mm. Frons about twice as wide as the eye, frontal punctures smaller than the eye facets, intervals densely reticulate. Pronotum with anterior angles almost as narrowly rounded as the posterior angles; surface of disc and margins usually smooth. Prosternal process with longitudinal carinae convergent anteriorly, not extending to the anterior margin of the prosternum, sometimes indistinct. Scutellum triangular, sparsely punctured. Elytral surface smooth. Apical border of abdominal ventrite 5 sternite bisinuate in males, broadly emarginate in females; ventrite 6 with median emargination in males and truncate in females.

Distribution of Illeis flava

76

Ladybird Beetles of the Australo-Pacific Region

Specimens examined

Description

AUSTRALIA: Queensland: Margin (1, ANIC); 17.06S 145.36E, 16 1.5 km W of Mt Haig, G.B. Monteith (1, QM); 17.15S 145.38 E, Lake Barrine, Atherton Tableland, 11.xii.1985, J. Balderson (1, ANIC); Palmerston Nat. Pk., 1000′, TullyCairns Powerline, Rainforest, at light, 6.xi.1966, E. Britton (1, ANIC). New South Wales: Starr’s Ck., Lansdowne S.F.N of Taree, 9. i.1987, G. Williams (1, ANIC); Dorrigo NP., Never Never picknick site, 6.iv.1993, C. Reid (1, ANIC).

Colour pattern (Figs 78A–D). Head yellow; pronotum yellow, with a triangular black spot; prothoracic hypomera yellow; prosternum brown. Scutellum black. Elytra pale yellow to orange, with an extensive black pattern as in Figs 78A–D; epipleura yellow, with a darker area corresponding to black spots on the dorsal side. Mesoventrite, metaventrite and abdomen black; legs brown or black. Morphology. Length 3.7–5.2 mm. Head between the eyes 1.5 times as broad as the eye, with fine, reticulate microsculpture and minute punctures. Pronotum with anterior angles much more strongly rounded than the posterior angles; surface smooth, minutely and sparsely punctured, with a group of larger punctures in front of the scutellum. Scutellum triangular, finely punctured. Elytra slightly longer than wide, almost completely smooth. Prosternal intercoxal carinae convergent anteriorly, extending almost to the anterior margin of the prosternum. Abdomen with an apical border of ventrite 5 broadly emarginate in males, straight in females; the apical margin of ventrite 6 deeply emarginate medially in males with median V-shaped notch in females. Male terminalia (Figs 78E–G). Female terminalia (Figs 78H,I). Measurements (n = 13): TL 3.75–5.20 (4.19 ± 0.39); PL 0.60–1.10 (0.83 ± 0.12); EL 2.80–3.90 (3.31 ± 0.30); PW 1.60–2.30 (1.91 ± 0.21); EW 2.70–4.00; (3.27 ± 0.31); GD 1.25–1.90 (1.55 ± 0.20). Ratios: PL/PW 0.34–0.57 (0.44 ± 0.06); EL/EW 0.91–1.12 (1.01 ± 0.05); EL/PL 3.29–5.67 (4.02 ± 0.54); EL/GD 1.91–2.72 (2.16 ± 0.22); TL/EW 1.18–1.48 (1.29 ± 0.07).

Distribution Known only from a few scattered localities in Queensland and northern New South Wales, Australia.

Illeis galbula (Mulsant, 1850) (Fig. 78)

Psyllobora (Illeis) galbula Mulsant, 1850: 166. Type locality: Australia. Mulsant 1850: 1026; 1866: 127. Thea galbula: Crotch 1874: 135. Halyzia galbula: Masters 1888: 89; Lea 1902: 490. Leptothea galbula: Weise 1898: 227; Korschefsky 1932: 570; Bielawski 1961a: 354, 1961b: 411; Richards 1980: 26. Illeis galbula: Crotch 1871: 4; Timberlake 1943: 43; Ślipiński 2007: 169. Illeis (Leptothea) galbula: Iablokoff-Khnzorian 1979: 63, 1982: 287; Pope 1989: 640; Leptothea csikii Weise, 1902: 497. Type locality: New Guinea. Korschefsky 1932: 570; Bielawski 1961b: 407. Synonymised by Iablokoff-Khnzorian 1979: 63 (lectotype). Leptothea moseri Weise, 1902: 497. Type locality: Key Islands. Korschefsky 1932: 570; Bielawski 1961b: 412. Synonymised by Iablokoff-Khnzorian 1979: 63.

Diagnosis Illeis galbula is easily distinguished from other species of the genus by its black and yellow colour pattern.

Distribution of Illeis galbula

Types I galbula: Museum Paris Australie Verreaux 2–47/ 825/ 60/2.47/Psyllobora galbula Muls. auct. det. (MNHN, lectotype). I. csikii: Nova-Guinea: Sattelberg (TMB, holotype). L. moseri: ‘Leptothea moseri m. Key-ins (ZMB, lectotype).

VII. Review of the Australo-Pacific Coccinellini

77

Specimens examined INDONESIA: Western New Guinea: Prov. Jayawijaya, Wamena, Jiwaka, 11.iv.1991, 1700–2000m, A. Riedel (1, NHMS); Prov. Baliem Tal Jiwika 1700–2300m, 1–2.ix.1990, A. Riedel (1, NHMS); Baliem Tal, 1700m, iii.1992, Jiri Kolibac (3, NHMB); (NW) Hollandia (Kota Baru), sweeping, 25–28. vi.1962, N.Wilson (1. BPBM); Japen Isl., Serui. Mantembu, 100–500m, 16.xii.2000, A. Riedel (1, NHMS). KAI ISLANDS: Key Ins. (3, MNHN). PAPUA NEW GUINEA: (NE) Wau, Morobe Distr. 1300m, 21.viii.1961, J. Sedlacek (1, BPBM); New Guinea: (NE) Umi R. Highlands, Hwy., 88mi. w. Lae, 30.iii.1973, K.W. Stroede (1. ANIC); Dogura, Papua, vii.1930, R. L. Cassidy (1, CAS); Aseki, ii.1972, H. Ohlmus (1, ANIC); SE Rouna, 300–500m, xi.1968, N. L. H. Krauss (3, BPBM); Guawa R. 147.10E-6.45S, 11.iv.1966, P. Shanahan (1, BPBM); Umboi Island, 1km N. Awikom, 600m, light trap, 21–28.ii.1967, G. A. Samuelson (1, BPBM). AUSTRALIA: Queensland: Eclectus, Iron Range; Magnetic Island, Arcadia; Mt. Tozer; Cooktown; Ayr; Mt Molloy; Kuranda; Barron Falls; Cairns; Elizabeth Gorge; Atherton; Ravenshoe; Millaa-Millaa; Mt Garnet; Townsville; Ayr; Mackay; Eungella; Yeppoon; Rockhampton; Bundaberg; Kenilworth State Forest; Bunya Mts; Brisbane; Mt Glorious; Brisbane; Moreton Bay; Ormiston; Gatton; Mt Tamborine. New South Wales: Eltham; Lismore; Richmond Range; Dorrigo; Port Macquarie; Comboyne; Tubrabucca; Wingham; Maitland; Abernethy; Huntley; Ourimbah; Narara; Gosford; Erina; Richmond; Sydney; Picton; Helensburgh; Otford; Mt Keira; Wollongong; Minnamurra Falls; Kangaroo Valley; Tomakin. Australian Capital Territory: Weston; McKellar; Waniassa; Gordon. Victoria: Millgrove; Melbourne; Warburton. NEW ZEALAND: CL little Barrier I., Walpawa TK, 27.iii.1997, D. M. Gleeson (1, NZAP); AK Kohimarama, on rose bush, 18.iv.1996, L. H. Clunie (1, NZAP); AK Avondale, on cucumber, ii.1993, G. Hall (1, NZAP); AK Bethells, Matuku Res, 14.iii-10.v.1989, G. Hall (1, NZAC).

Distribution of Illeis gressitti

with the anterior margin dark. Metaventrite and abdomen largely dark brown. Morphology. Length 5.7–6.2 mm. Head between eyes 0.7– 0.8 times as broad as the eye width; frons flat, with reticulate microsculpture and minute punctures. Pronotum surface smooth, minutely and sparsely punctured. Scutellum triangular, finely punctured. Elytra slightly longer than wide, finely punctate; interspaces smooth and shiny. Prosternal intercoxal carinae parallel and extend to middle of the prosternum. Abdomen with the apical border of ventrite 5 broadly emarginate in males, straight in females; the apical margin of ventrite 6 rounded in both sexes. Male terminalia (Figs 79D–H). Female terminalia. Not examined. Measurements (n = 5): TL 5.70–6.20 (6.06 ± 0.22); PL 1.20–1.40 (1.26 ± 0.09); EL 4.50–5.00 (4.78 ± 0.18); PW 2.70– 2.90 (2.80 ± 0.07); EW 4.70–5.10 (4.82 ± 0.18); GD 2.00–2.40 (2.18 ± 0.15). Ratios: PL/PW 0.43–0.48 (0.45 ± 0.02); EL/EW 0.96–1.02 (0.99 ± 0.03); EL/PL 3.43–4.17 (3.81 ± 0.29); EL/ GD 2.00–2.50 (2.20 ± 0.18); TL/EW 1.21–1.32 (1.26 ± 0.06).

Types Illeis gressitti: New Guinea, N.E. W. Highlands, Corn Farm, 1500 m, Oct. 19, 1958, J.L. Gressitt (BPBM).

Distribution

Specimens examined

This species is widely distributed in eastern and southern parts of Australia, the island of New Guinea, and Kai Islands (Indonesia); introduced to New Zealand.

INDONESIA: Western New Guinea: Waigeu, 2500 ft, iv.1938, L.E. Cheeseman (BMNH). PAPUA NEW GUINEA: (NE) Wau, Morobe Distr., 1200m, 27.x.1961, J. Sedlacek (1, BPBM); (NE) Wau, Morobe Dist.,1200m, 7.ix.1961, J. Sedlacek (1, BPBM); Mt Kaindi, Wau, M. Dist. T. P. N. G, 18.i.1970, L. Roduni (1, UQIC); Upper Manki L.A., Bulolo M. Dist., In sticky traps, 30.vi.1972, F. R. Wylie & P. Shanahan (1, ANIC); Western Highlands, Jimi River, 4700ft, 16.vii-21.ix.1961, W. W. Brandt (2, ANIC); Prov. Morobe, Aseki, 1000–1300m, 13.x.1992, A. Riedel (1, NHMS).

Illeis gressitti Bielawski, 1961c (Fig. 79)

Illeis gressitti Bielawski, 1961c: 517. Type locality: Papua New Guinea, Western Highlands. Iablokoff-Khnzorian 1982: 294.

Diagnosis Illeis gressitti can be distinguished fromother species in the Australo-Pacific region by its very narrow frons, uniformly yellow elytra, and the pale pronotum bearing three black spots.

Description Colour pattern (Figs 79A–C). Head and head appendages yellow. Pronotum pale with three black spots in the middle, the prosternum and prothoracic hypomeron yellow. Scutellum black. Elytra yellow, anterior and lateral borders transparent. Ventral side mostly black, but the mesanepisternum, mesepimeron, metanepisternum and metepimeron areusually yellow, but can be completely black in some specimens. Mesoventrite yellow,

Distribution Endemic to New Guinea.

Illeis wyliei sp. nov. (Fig. 80) Diagnosis This species is similar to the Australian species Illeis flava, but can be distinguished by the entirely yellow lateral margins of the pronotum, a narrower frons and the distinctive male genitalia.

78

Ladybird Beetles of the Australo-Pacific Region

Megalocaria Crotch, 1871

Description Colour pattern (Figs 80A–C). Head and head appendages pale yellow, labrum occasionally brown. Pronotum pale, with black spots medially; lateral and anterior borders transparent without thickened margins. Prosternum yellow. Elytra yellow, without dark spots; Scutellum black. Pterothorax black ventrally, except for the pale mesepimeron. Abdominal disc black. Legs largely dark. Morphology. Length 4.3–5.3 mm. Head between the eyes 1.5 times as broad as the eye; frons flat or weakly concave with fine, reticulate microsculpture and minute hairs. Pronotum widest near the middle, with the lateral margins rounded; surface smooth, minutely and sparsely punctured; interspaces faintly reticulate, shiny. Scutellum triangular, finely punctured. Elytra slightly longer than wide, disc with irregular punctures; interspaces shiny and polished. Prosternal intercoxal carinae convergent anteriorly, and extending to the middle of the prosternum. Abdominal postcoxal line very short, not extending to the posterior margin of ventrite 1. Apical border of ventrite 5 straight in both sexes; the apical margin of ventrite 6 shallowly emarginate in males, almost truncated in females. Male terminalia (Figs 80D–G). Female terminalia (Fig. 80H). Measurements (n = 3): TL 4.3–5.3 (4.87 ± 0.51); PL 0.8–1.0 (0.93 ± 0.12); PW 2.20–2.30 (2.27 ± 0.06); EL 3.4–4.0 (3.80 ± 0.35); EW 3.4–4.0 (3.73 ± 0.31); GD 1.60–1.80 (1.73 ± 0.12). Ratios: PL/PW 0.36–0.43 (0.41 ± 0.04); EL/EW 1.00–1.05 (1.02 ± 0.03); EL/PL 4.00–4.25(4.08 ± 0.14); EL/GD 1.89–2.22 (2.20 ± 0.31); TL/EW 1.25–1.39 (1.30 ± 0.08).

Specimens examined Holotype male: PAPUA NEW GUINEA: ‘Upper Manki L.A., Bulolo. M. Dist., In sticky traps, F. R. Wylie & P. Shanahan, 4.viii.1972’ (ANIC). Paratypes: PAPUA NEW GUINEA: Upper Manki L.A., Bulolo. M. Dist., In sticky traps, F. R. Wylie & P. Shanahan, 14.vii.1972 (1, ANIC). Morobe Prov., Wau, Mt Kaindi, Near Edie Ck., 1900–2100m, 07.ii.1998, A. Riedel (1, NHMS). INDONESIA: Western New Guinea: Irian Jaya: Prov. Jayawijaya, Wamena, Kosarek, 25.9.1991, 1650–1700m, A. Riedel (1, NHMS).

Remarks Named after Dr Ross Wylie (Biosecurity Queensland Control Centre, Department of Agriculture and Fisheries) who collected the holotype of this species and many interesting ladybirds during his work in Papua New Guinea.

Distribution Endemic to New Guinea.

Distribution of Illeis wylei

(Figs 81–83)

Megalocaria Crotch, 1871: 6. Type species by monotypy, Neda reichei Mulsant, 1850. Archaeoneda Crotch, 1874: 169. Type species by original designation, Coccinella tricolor Fabricius, 1787. Synonymised by Iablokoff-Khnzorian 1982: 520.

Diagnostic combination Length 7–15 mm; dorsum strongly convex, glabrous. Elytra broadly explanate, with variable patterning. Anterior clypeal margin straight between distinct lateral projections. Antenna 11-segmented with a loosely articulated club; terminal antennomere elongate. Prothoracic hypomeron without foveae. Prosternal process extending beyond the procoxae, with parallel carinae. Anterior margin of the mesoventrite deeply emarginate anteriorly. Elytral margin without a bead; epipleuron broad and sometimes concave, but without delimited foveae. Tibia spur formula 0–2–2. Tarsal claw with a large basal tooth. Abdominal postcoxal line approaching the posterior margin of the ventrite, incomplete laterally; the oblique postcoxal line strongly reduced or absent. Penis consists of a single sclerite. Infundibulum absent.

Description Head. Frons as broad as, or slightly broader than the eye width. Eyes partially beneath the anterior margin of the pronotum, interior orbits strongly divergent anteriorly. Antenna 11-segmented, scape flat and slightly expanded; pedicel much shorter and narrower than the scape; antennal club threesegmented, loosely articulated, terminal antennomere elongate, rounded apically. Clypeus straight between two lateral projections. Terminal maxillary palpomere strongly securiform. Prothorax. Pronotum weakly convex, the anterior border deeply emarginate, the lateral border arcuate. Prosternum in front of the procoxae much longer than the prosternal process. Prosternal process narrow, ~0.3 times as broad as the procoxae diameter, the surface with longitudinal carinae, usually convergent anteriorly and extending forward. Prothoracic hypomeron without foveae. Pterothorax. Elytra much broader than the pronotum at base, broadly explanate laterally without a marginal bead. Mesoventrite narrow, at the middle about half as broad as the mesocoxal diameter; the anterior border strongly emarginate, without a raised ridge; the posterior margin straight. Mesal surface with clear articulation with the prosternal process. Postcoxal line of the metaventrite separate medially; discrimen complete. Elytral epipleuron broad, usually 0.4 times as broad as the elytron, strongly inclined with shallow impressions but not delimited foveae. Legs not visible dorsally; the apex of the middle and hind tibiae each with two spurs; tarsal claws each with a subquadrate basal tooth. Abdomen with six ventrites in both sexes. Abdominal postcoxal line separated medially, incomplete laterally; the oblique postcoxal line indistinct or absent. Ventrite 1 about twice as long as ventrite 2. Posterior margin of ventrite 5 straight in males, and arcuate in females. Male terminalia. Penis consists of a single thick sclerite, with membranous tissue apically. Tegmen stout; penis guide usually strongly curved apically, as long as or slightly longer than the parameres. Female terminalia. Spermatheca slim and long, lightly sclerotised, strongly C-shaped, surface smooth. Sperm duct short, membranous; infundibulum absent; bursa copulatrix

VII. Review of the Australo-Pacific Coccinellini

short, slightly wrinkled, the apex lightly sclerotised. Coxites broad at the base; styli very short.

Biology The biology and immature stages of the Australo-Pacific species are unknown. The Asian M. dilatata (Fabricius, 1775) has been recorded as an aphid predator (Iablokoff-Khnzorian 1982) but the African M. tetrasticta (Fairmaire, 1888) is known to feed on eggs and nymphs of plataspid bugs (Hemiptera, Plataspidae) (Dejean et al. 2002).

79

Distribution About 20 known species are distributed in Africa, Asia and the Australo-Pacific region.

Remarks Megalocaria includes the largest known ladybird beetles distributed exclusively in the Old World, some reaching up to 18 mm in length. The composition of this genus and its distinctiveness from related Docimocaria, Anisolemnia and other large-bodied Coccinellini require further research.

Key to species of Megalocaria from the Australo-Pacific region 1. – 2(1). –

Dorsal side yellow or orange, with black, or black and red markings (Fig. 83A). Body length less than 10 mm ............. M. tricolor Body black with orange or red markings (Fig. 81A). Body length more than 10 mm .......................................................................2 Each elytron with a single large fascia (Fig. 82A) ..........................................................................................................M. singularis Each elytron with four isolated orange or red spots (Fig. 81A) ....................................................................................... M. rufitarsis

Megalocaria rufitarsis (Sicard, 1929b) (Fig. 81)

Caria rufitarsis Sicard, 1929b: 192. Type locality: Papua New Guinea, Mt. Victoria. Megalocaria reichei: Iablokoff-Khnzorian 1982: 524 (lectotype).

Diagnosis This species is very similar to M. tibialis (Sicard, 1929b) and M. korschefskyi (Sicard, 1929b) known from Sulawesi, but can be distinguished in having four isolated spots on each elytron, and none in the apical area (Fig. 81A).

Description Colour pattern (Figs 81D–G). Frons yellow in male, with small dark markings at the base and close to clypeus in females; head appendages yellow. Pronotum black with pale markings along the lateral and usually along the anterior borders. Prothoracic hypomeron black, except for an anterior pale area. Scutellum black. Elytra black with eight large orange-red spots not occupying the apical area. Ventral side entirely black in females, black in males except for a pale mesanepisternum, mesepimeron, external borders of prosternum and abdomen. Elytral epipleuron mostly black, anteriorly yellowish. Legs black, except for yellow tarsi. Morphology. Length 12–14 mm. Frons between the eyes 1.2– 1.4 times as broad as the eye width, flat with sparse punctures. Pronotum distinctly narrowing anteriorly with the lateral margin weakly sinuate; anterior angles projecting; the posterior angles broadly rounded. Punctures on the pronotum as large as those on the head, separated by 1–3 diameters. Longitudinal carinae on the prosternal process convergent anteriorly, almost reaching the anterior border. Scutellum triangular, ~1/15 width of the elytra at the shoulders. Elytra broadly explanate laterally; disc with fine punctures as large as those on the pronotum, separated by 2–3 diameters. Elytral epipleuron ~3.5–4.0 times as broad as the metanepisternum. Posterior margin of ventrite 5 truncate in males, arcuate in females. Male terminalia (Figs 81D–G). Female terminalia (Figs 81H,I). Measurements (n = 5): TL 11.63–13.75 (12.73 ± 0.75); PL 2.13–2.63 (2.45 ± 0.21); EL 8.88–11.13 (10.05 ± 0.80); PW

6.25–7.00 (6.58 ± 0.30); EW 11.88–13.00 (12.58 ± 0.47); GD 5.00–6.00 (5.50 ± 0.39). Ratios: PL/PW 0.34–0.39 (0.37 ± 0.02); EL/EW 0.75–0.86 (0.80 ± 0.04); EL/PL 3.74–4.82 (4.12 ± 0.44); EL/GD 1.78–1.90 (1.83 ± 0.05); TL/EW 0.98–1.06 (1.01 ± 0.03).

Types C. rufitarsis: Victoria, N. Guinée/ rufitarsis/ Coll. A. Sicard (MNHN, lectotype).

Specimens examined PAPUA NEW GUINEA: NE Wau, 1200m, 13.vi.1973, J. L. Gressitt (1, BPBM); (NE) Wau, Morobe Distr., Mt. Missim, 1220m, 13.ii.1963, J. Sedlacek (1, BPBM); Wau (1, BPBM); Wareo, Finsch Haven, Rev. L. Wagner (1, SAM); Finsch Haven, N. E. New Guinea (1, SAM); Finsch Haven, N. E. New Guinea, Rev. L. Wagner (3, SAM).

Remarks Iablokoff-Khnzorian (1982) synonymised M. rufitarsis with the common and highly polymorphic Oriental species, M. reichei (Mulsant, 1850). However, we have found consistent differences between these species in the male genitalia and external morphology, and consider M. rufitarsis to be a distinct species. Megalocaria reichei does not occur in the study area.

Distribution Known only from New Guinea.

Distribution of Megalocaria rufitarsis

80

Ladybird Beetles of the Australo-Pacific Region

Megalocaria singularis (Weise, 1902) (Fig. 82)

Caria singularis Weise, 1902: 504. Type locality: New Guinea: Sattelberg. Anisolemnia singularis: Korschefsky 1932: 270. Megalocaria singularis: Iablokoff-Khnzorian 1982: 526.

Diagnosis This is the largest ladybird beetle in the study area, reaching up to 15 mm. It is easily distinguished by its unique elytral colour pattern (Fig. 82A).

Description Colour pattern (Figs 82D–G). Frons yellow in males, with large, triangular black marks in the central area in females. Head appendages yellow. Pronotum black, with a large pale area in the anterior-lateral angles, the lateral border narrowly black. Scutellum black, background colour of the elytra black, with narrow transverse pale stripes on either side of the scutellum, connected by large orange-red fascia (Fig. 82A). Ventral side body black, except for the pale mesepimeron, mesanepisternum, anterior border of prosternum and external borders of abdomen. Prosternal hypomeron black, except for pale patches anteriorly. Elytral epipleuron black, except for anterior half pale. Legs largely black, with the femur partially pale. Morphology. Length 11–15 mm. Frons between the eyes 1.3–1.5 times broader than the eye diameter. Lateral margin of the pronotum faintly sinuate before the anterior angles. Punctures on the pronotum smaller than those on the head, separated by 2 diameters. Longitudinal carinae on the prosternal process almost parallel, ending very close to the anterior border. Scutellum at the base ~1/12 times as broad as the elytra at the shoulders. Elytra broadly explanate laterally, with punctures as big as those on the pronotum, separated by 2–4 diameters. Elytral epipleuron four times as broad as the metanepisternum. Abdominal postcoxal line incomplete laterally; the oblique postcoxal line very short and indistinct in some specimens. Posterior border of ventrite 5 truncate in males, arcuate in females. Male terminalia (Figs 82D–G). Female terminalia (Figs 82H,I). Measurements (n = 3): TL 10.88–13.75 (12.75 ± 1.63); PL 2.00–2.50 (2.33 ± 0.29); EL 8.88–11.00 (10.21 ± 1.16); PW 6.50–7.25 (6.83 ± 0.38); EW 12.50–13.75 (13.08 ± 0.63); GD 5.25–6.25 (5.67 ± 0.52). Ratios: PL/PW 0.31–0.37 (0.34 ± 0.03); EL/EW 0.71–0.83 (0.78 ± 0.06); EL/PL 4.30–4.44 (4.38 ± 0.07); EL/GD 1.69–1.95 (1.80 ± 0.14); TL/EW 0.87–1.06 (0.97 ± 0.10).

Types ‘N. Guinea, Biro, 1898: Sattelberg, Huon-Golf; Holotypus’ (TMB, holotype).

Specimens examined INDONESIA: Western New Guinea: S. Arfak (1, MNHN); Arfak Mts. (BMNH). PAPUA NEW GUINEA: Wau, 1200m, 7.x.1965. J. M. Sedlacek (BPBM); Wau, 1250m, 30.viii.1965 (ANIC); Morobe Prov., Ca. 10km S Garaina, Saureri, 1500m, 24.iii.1998, A. Riedel (1, NHMS); NE Mt. Missim (1, CAS).

Distribution of Megalocaria singularis

Distribution Endemic to the island of New Guinea.

Megalocaria tricolor (Fabricius, 1787) (Fig. 83)

Coccinella tricolor Fabricius, 1787: 59. Type locality: ‘Insula Amsterdam.’ Neda tricolor: Mulsant 1850: 282; 1866: 200; Arrow 1927: 57. Archaioneda tricolor: Crotch 1874: 169; Korschefsky 1932: 271. Megalocaria tricolor: Iablokoff-Khnzorian 1982: 529. Archaioneda tricolor var. fijiensis Crotch, 1874: 169; Korschefsky 1932: 272; Timberlake 1943: 27.

Diagnosis This species is much smaller than other species in the genus, and is also the only Megalocaria known from the South Pacific islands of Tonga and Fiji. It is easily distinguished from other species by its unique yellow or tricoloured elytral colour pattern.

Description Colour pattern (Figs 83A–D). Head and appendages yellow. Pronotum yellow, with the base and most of the lateral margins black. Elytra in typical tricolour form, yellow with black and reddish markings as in Fig. 83A, the basal border and suture narrowly black; in some forms the elytra uniformly yellow, except for a black basal border and suture (Fig. 83B). Ventral side black, except for the pale mesepimeron, mesanepisternum and external borders of the abdomen. Prosternal hypomeron yellow, except for a black lateral border and posterior part. Elytral epipleuron entirely yellow, or yellow with black patches. Legs mostly yellow, femora partially black. Morphology. Length 7.0–9.5 mm. Frons 1.3–1.5 times broader than the eye diameter. Pronotum weakly convex, lateral border arcuate. Punctures on the pronotum smaller than those on the head, usually separated by 1 diameter. Prosternum densely hairy; the prosternal process with strongly convergent longitudinal carinae, ending ~4/5 way to the anterior border. Elytra lightly punctured, punctures as large as those on the pronotum, separated by 2–3 diameters. Elytral epipleuron ~2.5 times as broad as the metanepisternum. Abdominal postcoxal lines separated medially; the oblique postcoxal line absent (Fig. 83K). Ventrites 3 and 4 with distinct median projections posteriorly in females, truncate in males; posterior margin of ventrite 5 arcuate in both sexes. Male terminalia (Figs 83E–H). Female terminalia (Figs 83I,J). Measurements (n = 5): TL 7.10–9.50 (8.69 ± 0.71); PL 1.40– 1.80 (1.65 ± 0.10); EL 6.40–8.00 (7.26 ± 0.47); PW 3.90–4.90

VII. Review of the Australo-Pacific Coccinellini

81

Distribution of Megalocaria tricolor

(4.46 ± 0.25); EW 7.00–8.50 (7.78 ± 0.38); GD 3.50–4.45 (3.83 ± 0.31). Ratios: PL/PW 0.36–0.40 (0.37 ± 0.01); EL/EW 0.87–1.01 (0.93 ± 0.04); EL/PL 4.00–4.75 (4.40 ± 0.25); EL/ GD 1.78–2.03 (1.90 ± 0.08); TL/EW 0.89–1.18 (1.12 ± 0.08).

Specimens examined TONGA: Vavau I.: Neiafu, 0–100m, i.1980, N. L. H. Krauss (1, BPBM); Vavau I.: Neiafu, 0–100m, vi.1969, N. L. H. Krauss (1, BPBM); Tongatapu: Nuku’alofa, iii.1966, N. L. H. Krauss (2, BPBM); Iles Wallis, 1912, Bouge (4, MNHN); Niue I., airport, 28.ii.1979, T. G. Mautama (1, NZAC); Eua I. Tonga Is., 12.iv.1974, J. A. Litsinger (1, NZAC); Vavau I, Neiafu, 17.ii.1977, P. A. Maddison (1, NZAC); Tonga, i.1930, H. R. Rabone (3, AM). FIJI: Ovaiau, Levuka, 0–100m, xi.1975, N. L. H. Krauss (1, BPBM); Vanua Levu: Savusavu, 0–100m, iii.1978, N. L. H. Krauss (1, BPBM); I. Wallis (1, MNHN); Suva, 08.iii.1935, R. V. Fyfe. (1, ANIC); Lau Is, Lakeba I, Tubou v, sweeping, 22.vi.1977, J. S. Dugdale (1, NZAC); Viti Levu, Korulevu, 15.iii.1976, P.A. Maddison (1, NZAC).

Distribution Tonga, Samoa, Fiji, including Wallis Island.

Remarks Considering the known geographic distribution of this attractive species, the type locality ‘Amsterdam Island’ in the Pacific Ocean is probably erroneous. Iablokoff-Khnzorian (1982) claimed he examined specimens from New Caledonia, but this is probably an error and this species is not mentioned in Nattier et al. (2015).

Micraspis Chevrolat, 1836 (Figs 84–92)

Micraspis Chevrolat in Dejean, 1836: 435. Type species designated by Hope 1840: 157, Coccinella striata Fabricius, 1792. Alesia Mulsant, 1850: 343. Type species designated by Korschefsky 1932: 304, Coccinella striata Fabricius, 1792. Synonymised by Korschefsky 1932: 304.

Verania Mulsant, 1850: 358. Type species designated by Crotch 1874: 175, Coccinella comma Thunberg, 1781. Synonymised by Fürsch 1964: 71. Cisseis Mulsant, 1850: 129. Type species by monotypy, Coccinella furcifera Guérin-Méneville, 1835. Synonymised by IablokoffKhnzorian 1979: 72 [junior homonym]. Cissella Weise, 1895: 153. Replacement name for Cisseis Mulsant, 1850.

Diagnostic combination Length 3.0–5.3 mm, dorsum glabrous. Elytra colour pattern variable but usually with longitudinal black stripes on yellow background (Fig. 87A). Antenna usually shorter than the head width with a compact, three-segmented club. Anterior clypeal margin straight. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere strongly securiform. Prothoracic hypomeron without a fovea. Prosternal process with parallel carinae. Anterior border of the mesoventrite straight with a raised ridge. Scutellum very small. Elytral margin with a complete bead; epipleuron narrow, without foveae. Tibial spur formula 0–2–2. Tarsal claws each with a subquadrate basal tooth, rarely simple. Abdominal postcoxal lines separate medially, not strongly recurved, incomplete; the oblique postcoxal line usually distinct. Penis consists of a single sclerite. Infundibulum absent. Spermatheca small, cornu curved apically with a well-developed ramus and nodulus.

Description Body elongate to broadly oval; dorsum glabrous. Head. Frons broad, at least twice as broad as the eye width. Anterior margin of clypeus straight, or slightly arcuate between lateral projections. Antenna 11-segmented, scape not flattened, the pedicel distinctly narrower than the scape; antennal club compact, three-segmented, the penultimate antennomere about as long as the terminal segment; terminal antennomere subquadrate, apically rounded. Mandible bifid apically, the molar part with basal teeth. Terminal maxillary palpomere distinctly securiform. Prothorax. Pronotum convex with lateral margins narrowly explanate; the anterior margin arcuate medially. Prosternal process with parallel carinae extending forward slightly,

82

Ladybird Beetles of the Australo-Pacific Region

sometimes indistinct. Prothoracic hypomeron without delimited foveae. Pterothorax. Scutellum very small, usually less than 1/15 times as broad as the elytra at the shoulders. Elytra narrowly explanate laterally; epipleuron without foveae. Mesoventrite at the middle at least half as broad as the mesocoxal diameter; anterior border straight with anterior carinae; the mesal surface without clear articulation with the prosternal process. Postcoxal line on the metaventrite joined medially, complete laterally. Mid and hind tibiae each with two spurs apically. Tarsal claws with subquadrate basal teeth; rarely claws simple. Abdomen with six ventrites in both sexes. Postcoxal line separate medially, not recurved, incomplete laterally; oblique postcoxal line weak but usually present. Male terminalia. Penis stout, consists of a single sclerite. Female terminalia. Spermatheca small, cornu curved weakly apically, ramus and nodulus distinct and close to each other at the base. Sperm duct short, infundibulum absent. Coxites slim at base, with a long handle, styli very small.

Biology The biology of a few Indian species and the Australian species M. frenata has been investigated, showing that both adults and larvae mostly feed on aphids but are capable of feeding on pollen (Anderson 1982; Hawkeswood and Turner 2002) during the pre-breeding season. The larva was described by Ślipiński (2007).

Distribution A widely distributed Old World genus, with ~30 known species, distributed in Asia, Africa and the Australian region.

Remarks Micraspis is usually recognised by its extremely small scutellum, paired tibial spurs on the meso- and metatibiae, and the straight anterior margin of the mesoventrite.

Key to species of the genus Micraspis from the Australo-Pacific region Tarsal claws simple; antennomeres 10 and 11 at least partially fused ..............................................................................M. furcifera Tarsal claws appendiculate with a basal tooth; antennomeres 10 and 11 separated ........................................................................... 2 Surfaces of the pronotum and elytra dull, with dense reticulate microsculpture; elytra with a fine black suture and external margins, and dorsum with a faint medially interrupted longitudinal fascia (Fig. 85A) ................................................... M. aphidectoides – Surfaces of the pronotum and elytra shiny, with at most fine reticulation; elytral dorsal colour pattern different than above .......... 3 3(2). Elytron orange, usually with a darker suture and with two large black spots (Fig. 84A); pronotum with two small spots ......... M. allardi – Pronotal and elytral colour pattern different from above .................................................................................................................... 4 4(3). Each elytron with a single stripe or broader fascia (Figs 91A, 92A) .................................................................................................. 5 – Each elytron with an apically bifid stripe, or extensive black colouration (Figs 88A, 90A) .............................................................. 6 5(4). Elytron with a very narrow, linear dorsal stripe (Fig. 92A) ................................................................................................. M. lineola – Elytron with a broad, fascia-like dorsal stripe (Fig. 91A) ................................................................................................... M. lineata 6(4). Elytron with variable colouration, but usually yellow with an extensive central black mark that is sometimes interrupted into two smaller darker longitudinal spots (Fig. 86A); female abdominal ventrite 6 with a medial pocket ........................ M. bennigseni – Elytron with a sigmoid or basally bifid stripe; female ventrite 6 simple ............................................................................................ 7 7(6). Elytron with a sigmoid discal stripe (Fig. 89C) .................................................................................................................. M. frenata – Elytron with a discal stripe that is bifid at the base (Fig. 87A) ........................................................................................................... 8 8(7). Elytral external margins broadly explanate. Bifid dorsal stripe reaches the base of the elytron but is not connected to the black sutural area (Fig. 87A) ................................................................................................................................................ M. confusa – Elytral external margins narrower. Bifid dorsal stripe does not reach the base of the elytron but usually joins the black sutural area (Fig. 88A) ............................................................................................................................................................... M. flavovittata 1. – 2(1).

Micraspis allardi (Mulsant, 1866) (Fig. 84)

Lemnia allardi Mulsant, 1866: 249. Type locality: Northern India. Verania allardi: Crotch 1874: 177; Korschefsky 1932: 307; Gordon 1987: 22 (lectotype). Micraspis allardi: Iablokoff-Khnzorian 1982: 510; Hoang 1983: 59; Poorani 2002a: 335. Lemnia malaccensis Crotch, 1874: 177. Type locality: Gilolo. Synonymised by Weise 1912: 115.

Diagnosis This species could be easily identified by its yellow or reddish body, and four black maculae on elytra.

Description Colour pattern (Figs 84A–C). Head and appendages yellow. Background colour of the pronotum, scutellum and elytra yellow or orange, the pronotum with two, and the elytra with four black patches (Fig. 84A). Lateral border of elytra

yellow; suture black. Ventral side yellow, except for darker metaventrite, metanepisternum and external borders of the abdomen. Prothoracic hypomeron and elytral epipleura yellow. Legs usually yellow, with darker femora. Morphology. Length 3.4–4.0 mm. Frons ~2.5–2.8 times as broad as the eye diameter. Antenna 0.6 times as long as the head capsule, club compact; terminal antennomere with a rounded margin. Pronotum moderately convex, the anterior margin straight between distinct anterior angles; the lateral margin posteriorly strongly arcuate. Prosternal process with longitudinal carinae extending to the middle of the prosternum. Elytra moderately convex, much broader than the pronotum at the base; narrowly explanate laterally. Scutellum 1/20 of the elytral width, triangular. Mesoventrite with the anterior and posterior margins straight medially. Tarsal claw long, with a very small basal tooth. Abdominal postcoxal line (Fig. 84J) approaches the posterior margin of the ventrite, incomplete laterally; the oblique postcoxal line absent. Posterior margin of ventrite 5 slightly emarginate in males, truncate in females. Male terminalia (Figs 84D–G). Female terminalia (Figs 84H,I).

VII. Review of the Australo-Pacific Coccinellini

Distribution of Micraspis allardi

Measurements (n = 5): TL 3.40–4.00 (3.62 ± 0.24); PL 0.60–0.80 (0.70 ± 0.08); EL 2.50–3.00 (2.82 ± 0.19); PW 1.65– 1.95 (1.82 ± 0.12); EW 2.65–3.90 (3.21 ± 0.47); GD 1.40–1.90 (1.57 ± 0.20). Ratios: PL/PW 0.33–0.42 (0.38 ± 0.03); EL/EW 0.77–0.97 (0.89 ± 0.08); EL/PL 3.63–4.83 (4.06 ± 0.46); EL/ GD 1.58–2.07 (1.81 ± 0.18); TL/EW 1.03–1.28 (1.14 ± 0.10).

Types L. allardi: Type/ Type allardi Deyr (UCCC, lectotype); V. malaccensis: Type/gilolo/ Type malaccensis Cr (UCCC, holotype).

Specimens examined INDONESIA: Western New Guinea: Anggi, Tetaho, Kosmena, 1400–1750m, 26–28.iii.1993 (1, BPBM). Ambon Island: Amboina, 7.vi.1908 (1, BPBM). PAPUA NEW GUINEA: Mt. Lamington, 1300 to 1500 feet, C. T. McNamara (1, SAM); NE, Hagen town, 4.48S-144.17E, 1650M, 30.v.1966, J. L. Gressitt (1, BPBM); Baliem Tal, 1700m, iii.1992, J. Kolibac (1, NHMB); Mt. Lamington Dist., Northern Division, Papua, x.1929, C.T. McNamara (8, AM).

Distribution India, Pakistan, Myanmar, Indonesia, Philippines, Maluku, Papua New Guinea.

83

Morphology. Length 4.7–5.6 mm. Frons broad, ~2.8–3.0 times as broad as the eye diameter, surface convex and sparsely setose. Antennae as long as the frontal width. Pronotum with rounded lateral sided, margins explanate. The base of the pronotum slightly sinuate on either side close to posterior angles; the anterior border strongly arcuate medially; punctures on the disc and margins smaller than the eye facets; intervals reticulate and faintly shiny. Prosternal process with parallel longitudinal carinae extending to the middle of the prosternum. Scutellum ~1/20 of the elytral width at the shoulders. Elytral surfaces with variable punctures, interspaces densely reticulate, matt. External margins explanate; a reduced external bead sometimes visible near the shoulders; epipleura 2.5–3.0 times as wide as the metanepisternum. Abdomen with apical border of ventrite 5 shallowly emarginate in males, straight in females. Tarsal claws with a small basal tooth. Male terminalia (Figs 85D–H). Female terminalia (Figs 85I,J). Measurements (n = 6): TL 4.70–5.60 (5.18 ± 0.33); PL 0.70–0.80 (0.75 ± 0.05); EL 3.80–4.80 (4.38 ± 0.35); PW 2.10– 2.50 (2.33 ± 0.14); EW 3.50–4.70 (4.12 ± 0.43); GD 1.70–2.00 (1.92 ± 0.12). Ratios: PL/PW 0.29–0.35 (0.32 ± 0.02); EL/EW 1.00–1.10 (1.07 ± 0.03); EL/PL 5.38–6.29 (5.85 ± 0.37); EL/ GD 2.20–2.40 (2.29 ± 0.07); TL/EW 1.17–1.34 (1.26 ± 0.06).

Types M. aphidectoides: Ferguson R., May 1931, Handschin/ Verania aphidectoides m, Holotypus det Korschefsky/ Holotypus (NHMB, holotype).

Specimens examined AUSTRALIA: Northern Territory: Katherine, CSIRO Exp. Station, 5.ii.1970, K. Rottigan (ANIC): 35 mi W. of Katherine, at light, 25.ii.1966, J. Mahon (ANIC); Port Darwin (ANIC); Gregory National Park, 18–24.v.2001, T. Weir, P. Bouchard (ANIC). Western Australia: Kimberley Research Station, Ord River Valley, 9.iii.1982, E. Smith (ANIC); Mitchel Plateau, Mining Camp, 9.v.1983, I. Naumann & J. Cardale (ANIC).

Distribution Micraspis aphidectoides (Korschefsky, 1934) (Fig. 85)

This species has a very limited distribution across the northern parts of the Northern Territory and Western Australia.

Verania aphidectoides Korschefsky, 1934a: 108. Type locality: Australia, Northern Territory, Ferguson River. Micraspis aphidectoides: Iablokoff-Khnzorian 1979: 73; 1982: 498 (lectotype); Pope 1989: 676; Ślipiński 2007: 174.

Diagnosis Micraspis aphidectoides is easily distinguished from other members of the genus in having yellowish elytra with light brown stripes, strong microsculpture on the elytra, and distinctive male genitalia (Figs 85D,E).

Description Colour pattern (Figs 85A–C). Head and appendages yellow. Pronotum yellow, usually with dark medial area at the base. Scutellum dark brown. Elytra yellow, with two longitudinal brown markings on each elytron; the external borders and suture dark. Ventral side mostly brown, with the prosternum, elytral epipleura, mesepimeron, metepimeron and the lateral areas of abdominal ventrites lighter. Legs pale, with black tarsal claws.

Distribution of Micraspis aphidectoides

84

Ladybird Beetles of the Australo-Pacific Region

Micraspis bennigseni (Weise, 1903) (Fig. 86)

Verania bennigseni Weise, 1903: 231. Type locality: New Guinea: Madang (Friedrich Wilhelmshafen). Korschefsky 1932: 307. Micraspis bennigseni: Iablokoff-Khnzorian 1982: 519 (lectotype).

Diagnosis Micraspis bennigseni is easily recognised by its relatively rounded body and the elytral colour pattern with a single central macula (Fig. 86A).

Distribution of Micraspis bennigseni

Description Colour pattern (Figs 86D–G). Labrum light in males and dark in females. Frons yellow in males and with black patch in the middle in females. Pronotum black, with yellow areas around the anterior corners. Elytra yellow, with black borders and a large central black area of variable shape and size. Scutellum black, prosternum yellow, with black anterior margin in males and largely dark in females. Prothoracic hypomeron black, with anterior part yellow, Elytral epipleura yellow, with external borders black. Legs largely black. Pterothorax black, except for the pale mesepimeron. Morphology. Length 3.8–4.1 mm. Frons about twice as broad as the eye width. Eyes partially beneath the anterior margin of the pronotum, with the interior margin parallel. Clypeus straight medially between lateral projections. Antenna much shorter than the head capsule, club very compact, appears four-segmented; the terminal antennomere truncate. Pronotum moderately convex, the anterior margin straight medially; the lateral margin strongly arcuate posteriorly. Prosternal process with short longitudinal carinae. Elytra moderately convex, much broader than the pronotum at the base; the lateral margins distinctly explanate. Scutellum 1/20 width of the elytra at the bases, triangular with a basal margin as long as the lateral margin. Anterior margin of the mesoventrite straight. Metaventrite with discrimen not reaching the anterior margin; postcoxal line complete. Tibial spurs very short and difficult to see. Tarsal claw with a broad basal tooth. Abdominal postcoxal line (Fig. 86J) approaching the posterior margin and almost reaching the lateral borders; the oblique postcoxal line absent. Posterior margin of ventrite 5 strongly emarginate in males and truncate in females; ventrite 6 in females with a medial pocket. Male terminalia (Figs 86D–G). Female terminalia. Infundibulum cone-shaped (Fig. 86H,I); spermatheca with a relatively long cornu. Measurements (n = 3): TL 3.80–3.90 (3.85 ± 0.0.07); PL 0.70–0.7 (0.73 ± 0.04); EL 3.10–3.20 (3.15 ± 0.07); PW 2.00 (2.00 ± 0.00); EW 3.50–3.60 (3.55 ± 0.07); GD 1.70–1.80 (1.75 ± 0.07). Ratios: PL/PW 0.35–0.38 (0.36 ± 0.02); EL/EW 0.86–0.91 (0.89 ± 0.04); EL/PL 4.27–4.43 (4.35 ± 0.11); EL/ GD 1.78–1.82 (1.80 ± 0.03); TL/EW 1.06–1.11 (1.08 ± 0.04).

Types V. bennigseni: Friedr. Wilh. Hafen /v. Bennigsen / Verania bennigseni m. (ZMB, lectotype; DEI, paralectotype).

Specimens examined INDONESIA: Western New Guinea: Manokwari Prov., Mokwam, Arfak Mts., 1300–1650m, 17.iv.1993, A. Riedel (3, NHMS); Minyambou, 1500–1900M, 13–14.iv.1993; A. Riedel

(1, NHMS); Manokwari, Mokwam, Siyoubrig. 1400–1800m, 01.06.26S- 133.54.41E, 24–28.II.2007, A. Skale (1, NKME). PAPUA NEW GUINEA: Mt. Lamington, 1300 to 1500feet, C. T. McNamara (2, SAM); Morobe District, Wau, 6.x.1969, James. E. Tobler (2, CAS); Morobe Distr. Herzog Mts., Vagau, ca 4000 ft, 4–17.i.1965, M.A. Bcchus (2, BMNH); Wau, ii.1974 (2, CAS); Nondugl, 1600m, ix-xi.1951, G. Gyldenstolpe (1, CAS); Imondo, 18.v.1992, J. F. Grimshaw (1, NAQS); Bismark Gbg, Waghital, 5–7000ft., 1963, Cl. Voss (1, NHMB).

Distribution Endemic to the island of New Guinea.

Remarks We have examined several colour forms that differ from the type series, bearing a single circular macula on the elytra (Fig. 86A) but the male and female genitalia are identical to the type series. We have therefore decided to treat them as colour variations of M. bennigseni.

Micraspis confusa Pope, 1989 (Fig. 87)

Micraspis confusa Pope, 1989: 682. Type locality: Australia, Queensland, Bundaberg. Ślipiński 2007: 174.

Diagnosis This species is very similar to the southern vicariant M. flavovittata, but can be differentiated by the broadly explanate external margins of the elytra, and the dorsal bifid stripe extending anteriorly to the elytral base, but usually not connected to the suture.

Description Colour pattern (Figs 87A–C). Head yellow in males, with an anteromedial dark patch in females. Pronotum yellow, with basal border black and two short bands extending forwards. Scutellum black. Elytra yellow, with longitudinal stripes, external borders and suture black. Ventral side black or dark brown, except for the pale prothoracic hypomeron, mesepimeron, metepimeron and elytral epipleuron. Anterior and middle legs pale-coloured, femora darker; hind legs black. Morphology. Length 4.2–5.0 mm. Head largely exposed. Frons 2.0–2.3 times as wide as the eye diameter; flat and punctured, punctures as large as the eye facets and 1 diameter apart; interspaces reticulate. Anterior margin of the pronotum shallowly arcuate medially, the posterior margin slightly sinuate near rounded posterior angles. Pronotal disc very finely

VII. Review of the Australo-Pacific Coccinellini

85

species share similar colour patterns and have almost identical terminalia in both sexes. The Victorian M. flavovittata can be separated from the Queensland M. confusa by the narrowly explanate external margins of the elytra, and the dorsal bifid stripe usually not extending anteriorly to the elytral base but connected to the suture.

Description

Distribution of Micraspis confusa

punctate, intervals polished and shiny. Prosternal process transversely convex, surface with long carinae extending more than halfway to the anterior border of the prosternum. Elytra with punctures distinctly larger than those of the pronotum and head, the external margin comparably broadly explanate. Elytral epipleuron slightly concave. Anterior border of the mesoventrite shallowly emarginate. Posterior margin of ventrite 5 broadly emarginate in males, arcuate in females; ventrite 6 truncate apically in males, arcuate apically and bearing small tubercle on dorsal surface in females. Male terminalia (Figs 87D–G). Female terminalia. Not examined. Measurements (n = 3): TL 4.30–4.40 (4.33 ± 0.06); PL 0.70–0.90 (0.77 ± 0.12); EL 3.50–3.60 (3.57 ± 0.06); PW 2.00– 2.20 (2.10 ± 0.10); EW 3.50–3.60 (3.57 ± 0.06); GD 1.60–1.70 (1.63 ± 0.06). Ratios: PL/PW 0.32–0.45 (0.37 ± 0.06); EL/EW 0.97–1.03 (1.00 ± 0.03); EL/PL 3.89–5.14 (4.72 ± 0.72); EL/ GD 2.12–2.25 (2.19 ± 0.07); TL/EW 1.19–1.23 (1.22 ± 0.02).

Types M. confusa: Koebele and Perkins Colls./ Bundaberg, Australia, 21.xii.05/ R. C. L. Perkins, B. M. 1942–95/ Micraspis confusa sp. n., R. D. Pope det. 1989 (BPBM, holotype; BMNH, paratype).

Distribution

Colour pattern (Figs 88A–D). Head with frontal area yellow, sometimes with a dark patch anteriorly; occiput black. Pronotum anteriorly yellow, with the remaining part black; lateral margins black. Scutellum black. Elytron yellow, with lateral borders and suture broadly black, disc with a bifid stripe, usually not reaching the anterior or posterior margins but with the inner part connecting to the sutural dark area (Fig. 88A). Ventral side black, except for the pale mesepimeron, anterior part of prothoracic hypomeron; elytral epipleuron yellow, except for black external borders. Legs entirely black. Morphology. Length 4.5–5.0 mm. Head with frontal width 2.4–2.6 times that of the eye width, punctures as large as the eye facets, 1 diameter apart, intervals densely reticulate. Pronotum with punctures slightly smaller than those of head, 1–2 diameters apart; interspaces smooth or with reticulate microsculpture. Elytra with explanate external margins not quite reaching the apicosutural angles, conspicuously narrowed apically, the marginal bead faint and incomplete, sometimes absent. Discal punctures slightly larger than those of the pronotum, more deeply impressed; intervals smooth. Prosternal process with short and indistinct carinae usually extending anteriorly shortly before the base of the process. Mesoventrite with shallowly emarginate anterior margin. Scutellum triangular 1/15 times width of the elytra at the bases. Posterior margin of ventrite 5 broadly emarginate in males, truncate in females; ventrite 6 truncate apically in males, arcuate apically and bearing small tubercle on dorsal surface in females. Male terminalia (Figs 88E–H). Female terminalia (Fig. 88I). Measurements (n = 2): TL 4.7–5.2; PL 1.2–1.3; EL 4.0–4.1; PW 2.4–2.5; EW 3.6–3.8; GD 1.7–1.8. Ratios: PL/PW 0.50; EL/EW 1.05; EL/PL 3.33; EL/GD 2.35; TL/EW 1.24.

Types V. flavovittata: Type/ Austral, 53.55/ Verania flavovittata, (Type)/ flavovittata TYPE (BMNH, holotype).

This species is known only from a series of specimens collected in Bundaberg by R. Perkins and R. Koebele, who were searching for natural enemies of leafhoppers. No further specimens have been collected since their trip in 1904 (Pope 1989).

Micraspis flavovittata (Crotch, 1874) (Fig. 88)

Verania flavovittata Crotch, 1874: 176; Type locality: Australia: Melbourne. Korschefsky 1932: 309. Alesia flavovittata: Masters 1888: 90. Micraspis flavovittata: Iablokoff-Khnzorian 1979: 73; 1982: 518; Pope 1989: 681; Ślipiński 2007: 174.

Diagnosis Micraspis flavovittata is almost identical to M. confusa, and the taxonomic status of M. confusa requires further research. Both

Distribution of Micraspis flavovittata

86

Ladybird Beetles of the Australo-Pacific Region

Specimens examined AUSTRALIA: Victoria: Long Swamp, Discovery Bay Coastal Park Near Nelson, –38.092S 141.093E, 4.iv.2015, Reiner Richter (2, ANIC).

Remarks This species appears to be extremely rare and localised. It was once thought to be extinct (Pope 1989) but was recently rediscovered by Mr Reiner Richter in 2014, in swamp areas in western Victoria.

Micraspis frenata (Erichson, 1842) (Fig. 89)

black. Elytra yellow, with black external borders, suture and sigmoid dorsal stripe. Ventral side black, except for yellow mesepimeron, metepimeron, prothoracic hypomeron and elytral epipleuron. Anterior and middle legs usually yellow, femora at least partially black; hind legs black. Morphology. Length 3.7–4.7 mm. Frons between the eyes ~2.5 times as broad as the eye width. Eyes largely exposed, inner orbits parallel. Anterior border of the pronotum weekly arcuate medially, posterior borders evenly curved; disc finely punctured with interspaces smooth or finely reticulate, shiny. Prosternal process with carinae parallel and extending forward slightly beyond the process. Scutellum triangular, 1/16 times the width of the elytral bases. Elytral punctures as large as, or slightly larger than, the eye facets. External borders narrowly explanate, almost complete from the shoulders to the apex. Elytral epipleuron 2.0–2.2 times as broad as the metanepisternum. Anterior border of mesoventrite shallowly emarginate medially. Posterior border of ventrite 5 broadly but shallowly emarginated in males, usually slightly arcuate medially in females. Male terminalia (Figs 89D–F). Female terminalia (Figs 89G,H). Measurements (n = 10): TL 3.70–4.70 (4.25 ± 0.40); PL 0.60–0.80 (0.66 ± 0.07); EL 3.00–4.00 (3.58 ± 0.36); PW 1.80– 2.20 (2.03 ± 0.15); EW 2.80–3.60 (3.24 ± 0.30); GD 1.30–1.80 (1.56 ± 0.16). Ratios: PL/PW 0.30–0.36 (0.33 ± 0.02); EL/EW 1.06–1.14 (1.10 ± 0.03); EL/PL 4.88–5.83 (5.41 ± 0.39); EL/ GD 2.20–2.35 (2.29 ± 0.06); TL/EW 1.26–1.34 (1.31 ± 0.03).

Coccinella frenata Erichson, 1842: 239. Type locality: Australia: Tasmania. Verania frenata: Mulsant 1850: 362; Crotch 1871: 8, 1874: 175; Korschefsky 1932: 309; Bielawski 1962: 202. Alesia frenata: Masters 1888: 90. Micraspis frenata: Pope 1989: 679 (lectotype); Ślipiński 2007: 174; Nattier et al. 2015: 308. Coccinella bicruciata Montrouzier, 1861: 304. Type locality: New Caledonia, Balade and Art Island. Ceolophora vittipennis Fauvel, 1903: 328. Type locality: New Caledonia, Kanala. Syn. nov. Micraspis vittipennis: Chazeau 1978: 68. Micraspis lineata: Iablokoff-Khnzorian 1982: 514; Anderson and Hales 1983: 47.

Types

Diagnosis

M. frenata: Q, 27964/ Typus/ frenata Er. Van Diem. Schayer (ZMB, lectotype).

Micraspis frenata can easily be identified by the sigmoid dorsal stripe on the elytra, and the pronotum with a black basal area bilobed medially (Fig. 89A).

Description Colour pattern (Figs 89A–C). Head yellow, sometimes with an anteromedian dark patch. Pronotum yellow anteriorly, with base area black, lateral margin transparent. Scutellum

Distribution of Micraspis frenata

Specimens examined NEW CALEDONIA: 21.07S/164.57E, 30m Tiea Forest (GIE Fab Nicoli) Sclerophyllous forest, 30.i.2004, M. Wanat (ANIC); (N) 20.45S/164.53E, Tiendanite 50m, Solanaceae shrubs, 4.ii.2004, M. Wanat (ANIC); (S) 21.42S/165.41E, 20 m Moindou (ca. 3km E of) secondary Leucena bush, 30.i.2004, M. Wanat (ANIC).

VII. Review of the Australo-Pacific Coccinellini

AUSTRALIA: Queensland: Mareeba; Cairns; Chillagoe; Atherton; Tolga; Herberton; Townsville; Ayr; Charters Towers; Burleigh; Mackay; Marlborough; Barmount; Peak Downs; Rockhampton; Emerald; Edungalba; Biloela; Bundaberg; Biggenden; Mungallala; Mitchell; Kingaroy; Caloundra; Dalby; North Pine; Mt Glorious; Brisbane; Ormiston; Moreton Bay; Gatton; Toowoomba; Mt. Tamborine; Springbrook; Binna Burra. New South Wales: Kialoa; Wallangra; Moree; Graman; Glen Innes; Coffs Harbour; Narrabri; North Head; Coonabarabran; Port Macquarie; Ourimbah; Somersby; Lisarow; Colo Reserve; Gosford; Terrigal; Sydney; Moorna; Otford; Mt Keira; Wollongong; Kiama; Minnamurra Falls; Kangaroo Valley; Nowra; Batehaven; Cabbage Tree Ck.; Batemans Bay; Tuross; Wallaga Lake; Tathra. Australian Capital Territory: Canberra. Victoria: Woolonga; Millgrove; Warburton; Melbourne. Tasmania: Launceston; Hobart. South Australia: Lake Eyre; Marree; Renmark; Lyrup; Adelaide; Langhorne Ck.

Distribution This species is widely distributed in eastern and southern Australia and is also common in New Caledonia, possibly introduced.

Micraspis furcifera (Guérin-Méneville, 1835) (Fig. 90)

Coccinella furcifera Guérin-Méneville, 1835: pl. 51. Type locality: Australia. Cisseis furcifera: Mulsant 1850: 130. Verania furcifera: Crotch 1871: 8; Gordon 1987: 22 (lectotype). Alesia furcifera: Masters 1888: 90. Cissella furcifera: Weise 1895: 153; Korschefsky 1932: 377; Timberlake 1943: 16. Menevillidia furcifera: Brèthes 1923: 229. Micraspis furcifera: Iablokoff-Khnzorian 1979: 73, 1982: 515; Pope 1989: 675; Ślipiński 2007:174. Sospita flavolineata Mulsant, 1866: 113. Type locality: Australia. Synonymised by Crotch 1871: 8; Gordon 1987: 22 (lectotype). Verania gauthardi Mulsant, 1866: 241. Type locality: Australia, Central Africa. Synonymised by Crotch 1871: 8; Gordon 1987: 22 (lectotype).

metepimeron pale; metaventrite black except for a pale anterior area. Legs at least partially black. Morphology. Length 4.4–5.6 mm. Frons relatively long and flat, width 1.9–2.2 times the eye width. Frontal punctures as large as the eye facets, 1–2 diameters apart; interspaces weakly reticulate. Antenna with terminal and penultimate antennomeres at least partly fused. Pronotum convex, with punctures about as large as those on the frons, 1–2 diameters apart; interspaces shiny. Prosternal process without carinae. Elytra with very narrow explanate external margins, upturned at the external border to form a narrow gutter near the humeral angles; punctures similar in size to those on the pronotum, separated by 2–4 diameters; intervals smooth. Elytral epipleuron narrow, width 1.8–2.0 times as broad as the metanepisternum. Abdomen with apical border ventrite 5 very shallowly emarginate medially in males, slightly acuate medially in females. Male genitalia (Figs 90D–G). Female genitalia (Figs 90H,I). Measurements (n = 10): TL 4.40–5.60 (5.05 ± 0.44); PL 0.70–1.00 (0.81 ± 0.11); EL 3.50–4.70 (4.11 ± 0.41); PW 2.00–3.00 (2.40 ± 0.26); EW 3.40–4.30 (3.82 ± 0.32); GD 1.70–2.30 (2.01 ± 0.17). Ratios: PL/PW 0.23–0.40 (0.34 ± 0.05); EL/EW 1.00–1.14 (1.08 ± 0.05); EL/PL 4.60–6.29 (5.12 ± 0.63); EL/GD 1.85–2.32 (2.05 ± 0.14); TL/EW 1.19–1.62 (1.38 ± 0.12).

Types C. furcifera: Type / TYPE furcifera N. Holl. Guer., (UCCC, lectotype). S. flavolineata: Type/ Australia/ TYPE flavolineata Deyr., (UCCC, lectotype). V. gauthardi: Type/ S. Austral/ TYPE Gauthardi Bakewell (UCCC, lectotype).

Specimens examined AUSTRALIA: New South Wales: Bogan River, J. Armstrong (ANIC); Hay, xii.1972, on saltbush, V. Rogers (ANIC); Merran Creek, 17 km NE Swan Hill, 15.i.1986, Ch. Reid (ANIC). South Australia: 19 km N. Renmark, 14.xii.1995, K. Pullen (ANIC);

Diagnosis Micraspis furcifera differs from all its congeners in having simple pretarsal claws and antennomeres 10 and 11 at least partially fused. The elytral colour pattern with an oval spot situated behind the scutellum between the lateral stripe and the elytral suture is also unique.

Description Colour pattern (Figs 90A–C). Head yellow, with a dark marking at the base and often with triangular patches on the clypeus. Pronotum yellow, with a black transverse band along the base and two admedian spots, the anterior and lateral borders transparent. Scutellum black, or yellow with black external borders. Elytra with external margins transparent and yellow, suture black; disc with two longitudinal black fasciae, usually connected anteriorly and posteriorly and a small oval mark behind the scutellum (Fig. 90A). Pronotal hypomeron pale, except for the posterior area. Prosternum pale, with the prosternal process dark; mesoventrite, mesepimeron and

87

Distribution of Micraspis furcifera

88

Ladybird Beetles of the Australo-Pacific Region

8km NE Mannum, v.1989, Ch. Reid (ANIC); 30 mi SW Yunta, 19.vii.1969, G. Monteith (QM). Western Australia: 5 km S of Norseman, 25.xii.1985, P. Gullan (ANIC).

Distribution A rare species distributed across the southern part of Australia and often taken from salt bushes (Muehlenbeckia sp.).

Micraspis lineata (Thunberg, 1781) (Fig. 91)

Coccinella lineata Thunberg, 1781: 21, fig. 31. Type locality: ‘Capite bonae spei’ (probably erroneous). Verania lineata: Mulsant 1850: 360.

Diagnosis Micraspis lineata is very similar to M. frenata and M. lineola, but can be distinguished by having a single very broad and not sigmoid dorsal stripe.

Description Colour pattern (Figs 91A–C). Head and appendages yellow, frons entirely pale in males, usually with a black triangular patch medially in females. Pronotum with the basal half almost entirely black, with a black somewhat trapezoidal marking that extends forward close to, but not reaching, the anterior margin (Fig. 91A); scutellum black. Elytra with a broad black sutural area, and a broad anteriorly and posteriorly incomplete discal stripe; external margin narrowly black. Venter mostly black, except for the pale mesepimeron and metepimeron; legs black, with tarsi and sometimes tibiae brown to yellow. Morphology. Length 3.8–5.3 mm. Frons about twice the eye width. Antenna short, length 0.5–0.6 times the head capsule width. Frons flat, sparsely punctured. Pronotum widest before the base, strongly narrowing anteriorly; disc convex and punctate; interspaces shiny and smooth. Prosternal process with parallel carinae extending to about the middle of the prosternum. Scutellum 1/20 width of the elytra at the humeri. Elytra convex, with narrowly explanate lateral margins; disc punctate with punctures as large as those on the pronotum; interspaces shiny; elytral epipleuron 3.5–3.7 times the width of the metanepisternum. Anterior margin of prosternum straight, with marginal stria. Abdominal ventrite 5 truncate in both sexes. Male terminalia (Figs 91D–G). Female terminalia (Figs 91H,I). Measurements (n = 9): TL 3.80–5.30 (4.66 ± 0.55); PL 0.70–1.00 (0.88 ± 0.11); EL 3.10–3.80 (3.52 ± 0.25); PW 1.80– 2.30 (2.14 ± 0.17); EW 2.90–3.70 (3.37 ± 0.23); GD 1.60–2.00 (1.74 ± 0.12). Ratios: PL/PW 0.36–0.50 (0.41 ± 0.04); EL/EW 0.97–1.12 (1.04 ± 0.05); EL/PL 3.50–4.50 (4.01 ± 0.38); EL/ GD 1.80–2.18 (2.03 ± 0.12); TL/EW 1.22–1.56 (1.38 ± 0.14).

Specimens examined INDONESIA: Western New Guinea: Baliem Dist., Ilugwa, Melanggama, 9–12.ix.1990, 1900–2200m, A. Riedel (7, NHMS). PAPUA NEW GUINEA: NE Korgua, 1450m, 30.vi.1963, H.C. (1, BPBM); NE Koibuga, 1500m, 5.vii.1963, H. C. (3, BPBM); Nondugl, 1600m, ix-xi.1951, G. Gyldenstolpe (3, NHMS); Bismark Gbg, Waghital 5–7000ft, 1963, Cl. Voss (1, NHMB).

Distribution of Micraspis lineata

Distribution Endemic to the island of New Guinea.

Micraspis lineola (Fabricius, 1775) (Fig. 92)

Coccinella lineola Fabricius, 1775: 79. Type locality: Australia. Verania lineola: Crotch 1871: 8; 1874: 176; Froggatt 1902: 901; Korschefsky 1932: 309; Timberlake 1943:27. Micraspis lineola: Iablokoff-Khnzorian 1979: 73, 1982: 506; Pope 1989: 678; Ślipiński 2007: 174. Coccinella strigula Boisduval, 1835: 601. Type locality: Australia. Synonymised by Crotch 1874: 176. Verania strigula: Mulsant 1850:366, 1866: 243.

Diagnosis Micraspis lineola is very similar to M. lineata, but can be distinguished from that species in having elytra with a very narrow dorsal stripe, and only narrowly darkened suture and external margins.

Description Colour pattern (Figs 92A–C). Head entirely pale in males, usually with a dark anteriomedian patch in females. Pronotum yellow, with a narrow median transverse black stripe and sometimes admedian smaller patches. Elytral external borders and suture very narrowly black, dorsal stripe straight and very narrow. Scutellum black. Prosternal hypomeron, mesepimeron and epipleura yellow, except for the epipleural external margin. Mesoventrite and metaventrite black. Prosternum yellow in males and black in females. Anterior and middle legs pale, hind femur partially dark in males. All legs at least partially black in females. Morphology. Length 3.7–4.5 mm. Frons 2.5–2.7 times wider than the eye width; flat and sparsely punctured; interspaces reticulate. Pronotum widest before the base, narrowing anteriorly; disc convex, punctured; interspaces smooth and shiny. Prosternal process with carinae converging weakly, extending to about the middle of the prosternum. Scutellum width 1/20 of the elytral width at the humeri. Elytra punctured and shiny; epipleuron ~2.5–2.8 times wider than the metanepisternum. Mesoventrite weakly emarginate medially. Abdominal ventrite 5 apically emarginate in males, truncate in females. Male genitalia (Figs 92D–G). Female genitalia (Fig. 92H). Measurements (n = 13): TL 3.70–4.50 (4.10 ± 0.25); PL 0.70– 1.00 (0.82 ± 0.11); EL 3.00–3.60 (3.27 ± 0.19); PW 1.80–2.10

VII. Review of the Australo-Pacific Coccinellini

89

Distribution of Micraspis lineola

(1.97 ± 0.09); EW 3.10–3.70 (3.42 ± 0.20); GD 1.50–2.00 (1.77 ± 0.17). Ratios: PL/PW 035–0.53 (0.42 ± 0.06); EL/EW 0.91–1.00 (0.96 ± 0.03); EL/PL 3.10–5.14 (4.06 ± 0.56); EL/ GD 1.870–2.07 (1.86 ± 0.12); TL/EW 1.14–1.24 (1.20 ± 0.03).

Specimens examined INDONESIA: Biak Islands: Korim, Roidifu, 100m, 2.ii.2001, A. Riedel (1, NKME); Biak Is., Korim, Wouna, 21–22.iv.1993, A. Riedel (5, NHMS); Korim, Nernu, 100–150m, 12–14. xii.2000, A. Riedel (1, NHMS). PAPUA NEW GUINEA: NE Lae, 15m, 20.i.1962, G. Monteith & J. Sedlacek (2, BPBM); NE Lae, Sirguawa R 147.10E- 6.45S, 30m, 11.iv.1966, P. Shanahan (1, BPBM); NE Feramin, 1450m, 27.viii.1963, R. Straatman (1, BPBM); NE Feramin, 1400m, 24–25.viii.1963, R. Straatman (1, BPBM); Angoram, Sepik river, 29.v.1970, H. Pelz (2, CNC); Finschhafen, 15.v.1944, E. S. Ross (5, CAS); NE Markham River, 50m, 20–25.i.1962, J. Sedlacek (1,BPBM); SE Mamai Plt’n nr. Port Glasgow, 7.ii.1965, R. Straatman (1, BPBM). FIJI: Viti Levu, Sigatoka Res. Station, 28.ii.1977, J.C. Watt (NZAC). TONGA: Tongatapu, Vaini Cout. Experimental Station (NZAC); Eua, 14.iv.1974, J.A. Litsinger (NZAC); Tongatapu, 10.viii.1973, J.A. Litsinger (NZAC). AUSTRALIA: Northern Territory: Cooper Creek, 11 km S of Nimbuwah Rock, 3.vi.1973, Upton (ANIC); 18 km E of Oenpelli, 1.vi.1973, Matthews (ANIC). Queensland: Lockerbie, 6.vi.1969, G. Monteith (QM); Iron Range NP, 1 km N of Lamond Hill, 15.vii.1998, A. Calder (ANIC).

Distribution Widely distributed species in New Guinea, Pacific islands and northern Australia.

Microcaria Crotch, 1871 (Figs 93–97)

Microcaria Crotch, 1871: 7. Type species designated by Rye 1873: 329, Coccinella (Daulis) mulsanti Montrouzier, 1861.

Bothrocalvia Crotch, 1874: 143. Type species by original designation, Coccinella albolineata Gyllenhal in Schönherr, 1808. Syn. nov.

References: Chazeau 1978; Iablokoff-Khnzorian 1982: 218–270; Pope 1989: 662; Poorani 2002a: 321, 327. Diagnostic combination Length 5.0–6.8 mm, elytra convex, glabrous. Elytral colour pattern extremely variable. Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly shorter than head capsule with moderately compact threesegmented club. Terminal maxillary palpomere securiform. Pronotal disc evenly convex transversely; lateral margins slightly concave, within upturned and slightly thickened external borders. Prothoracic hypomeron with foveae near the anterior angles; prosternal process with distinct carinae extending anteriorly. Anterior margin of the mesoventrite deeply emarginate medially. Elytra with narrow raised margins; epipleuron distinctly foveate at the level of the hind coxae. Tibial spurs formula 0–2–2. Abdominal postcoxal line not recurved and almost complete laterally; oblique postcoxal line absent. Penis enlarged near the middle, apically attached to a membranous piece. Infundibulum absent; spermatheca with cornu weakly C-shaped; nodulus poorly developed, ramus distinct.

Description Body moderately convex, oval, dorsum glabrous. Head strongly withdrawn into the prothorax. Eyes largely beneath the pronotum, with inter orbits strongly divergent anteriorly. Frons narrow, 1.5–1.7 times the width of the eye. Antenna 11-segmented, 0.7–0.8 times the head width, scape slightly enlarged, the last three antennomeres forming a narrow but rather compact club; the terminal antennomere rounded apically, longer than the penultimate. Anterior clypeal margin straight. Terminal maxillary palpomere strongly securiform. Mandible bifid apically, the molar part with a basal tooth. Prothorax. Pronotum convex; the anterior margin strongly emarginate medially, forming blunt anterior angles. Prothoracic

90

Ladybird Beetles of the Australo-Pacific Region

hypomeron with distinct foveae. Prosternal process with admedian carinae extending forward to the middle of the prosternum. Pterothorax. Scutellum ~1/10–1/12 times as broad as the elytra at the shoulders. Elytra convex, slightly broader than the base of the pronotum, lateral margins not explanate. Elytral epipleuron slightly inclined, with distinct foveae at the middle. Mesoventrite strongly emarginate medially, with a complete raised ridge; the mesal surface with clear articulation with the prosternal process. Postcoxal line of the metaventrite joined medially, complete laterally. Apex of the middle and hind tibiae each with two spurs; tarsal claws with subquadrate basal teeth. Abdomen with six ventrites in both sexes. Postcoxal lines joined or separate medially, complete laterally, not recurved; oblique postcoxal line absent. Male terminalia. Penis consists of a single, apically strongly narrowed sclerite, and a ventrally attached apical membranous piece bearing several smaller sclerites; the basal sclerite is strongly swollen near the middle forming a partly membranous ventral sac (Fig. 93D). Female terminalia. Coxites with short apical styli; infundibulum absent; spermatheca with a weakly C-shaped

cornu; nodulus poorly developed, ramus distinct, both close together (Fig. 93K).

Biology Some species have been recorded as feeding on psyllids, aphids and nymphs of Cicadellidae (Chazeau 1978; Pope 1989).

Distribution Microcaria is a widely distributed Asian and Australian genus, extending from China and India to Australia, to the Pacific Islands (Chazeau 1978; Pope 1989).

Remarks Microcaria is externally identical to Coelophora and can be distinguished from that genus by the male and female genitalia only. Microcaria is distinguished from Coelophora in having a penis consisting of a medially expanded single sclerite, and the spermatheca with a cornu only weakly C-shaped, and a distinctly projecting ramus.

Key to species of Australo-Pacific Microcaria 1. – 2(1). – 3(2). – 4(3). –

Elytra uniformly reddish (Fig. 96A) ............................................................................................................................... M. versipellis Elytra uniformly black or with other colour pattern ........................................................................................................................... 2 Antennal club very narrow (Fig. 95J) ....................................................................................................................... M. salomonensis Antennal club distinct ......................................................................................................................................................................... 3 Elytra black, or with small reddish patch at apex or/and near the humeri on each elytron (Fig. 93A) .............................. M. jansoni Elytra with different colour pattern (Figs 94A–D) ............................................................................................................................. 4 Elytra red, with transverse black bands that are sometimes confluent apically (Fig. 97A) ...................................................M. vivida Elytra yellow or orange, with spots or longitudinal stripes (Figs 94A–D) ....................................................................... M. mulsanti

This species is easily distinguished by having a uniformly orange pronotum and black elytra bearing orange or red apical spots.

bead. Anterior margin of the mesoventrite roundly emarginate medially. Postcoxal lines joined medially. Elytral epipleuron 2.5 times as broad as the metanepisternum, with shallow foveae for the middle and hind legs. Posterior margin of ventrite 5 truncate in both sexes. Male terminalia (Figs 93D–I). Female terminalia (Figs 93J,K). Measurements (n = 8): TL 4.90–6.10 (5.65 ± 0.37); PL 0.80–1.00 (0.96 ± 0.07); EL 4.10–5.10 (4.69 ± 0.31); PW 2.55–3.55 (2.93 ± 0.19); EW 4.30–5.40 (5.00 ± 0.35); GD 2.40–3.05 (2.78 ± 0.20). Ratios: PL/PW 0.29–0.36 (0.33 ± 0.02); EL/EW 0.90–1.00 (0.94 ± 0.03); EL/PL 4.50–5.33 (4.91 ± 0.28); EL/GD 1.66–1.72 (1.69 ± 0.02); TL/EW 1.08– 1.20 (1.13 ± 0.04).

Description

Types

Colour pattern (Figs 93A–C). Head and appendages light brown; pronotum uniformly reddish; scutellum reddish to black; elytra black except for two reddish spots apically. Ventral side reddish except for pale mesepimeron and black external borders of elytral epipleura. Legs yellow-reddish. Morphology. Length 5–6 mm. Frons 1.0–1.5 times broader than the eye width. Pronotum moderately convex, densely punctured; punctures 1–2 diameters apart, interspaces densely reticulate. Prosternal process with admedian carinae strongly convergent anteriorly, extending forward to the middle of the prosternum. Elytra slightly broader than the pronotum at the base. Lateral borders narrowly explanate, with a thick marginal

C. jansoni: Q, C. York/ TYPE Jansoni C. York/ TYPE (UCCC, lectotype).

Microcaria jansoni (Crotch, 1874) (Fig. 93)

Coelophora jansoni Crotch, 1874: 155. Type locality: Australia, Cape York. Lea 1902: 490; Weise 1923: 132; Korschefsky 1932: 293; Gordon 1987: 19 (lectotype); Pope 1989: 663; Ślipiński 2007: 160. Microcaria jansoni: Timberlake 1943: 33. Lemnia jansoni: Iablokoff-Khnzorian 1982: 247.

Diagnosis

Specimens examined PAPUA NEW GUINEA: Lae, 30.i.1971, R. E. Parrott (1, CNC); Dogura, 01.x.1958, E. L. Cassidy (1, CAS); Weam, viii.1976, H. Ohlmus (1, ANIC); Misima Island, H. K. Bartlett (4, SAM). SOLOMON ISLANDS: Bougainville, Kukugai village (Ruin), 17.x1960–2.ii.1961, W.W. Brandt (1, ANIC); Gizo I., Gizo, 0–100M, ii.1984, N. L. H. Krauss (1, BPBM);

VII. Review of the Australo-Pacific Coccinellini

91

Distribution of Microcaria jansoni

Gizo I., 70M, 18.vii.1964, J. & M. Sedlacek (3, BPBM); Guadalcanal: Honiara, 0–200m, xii.1975, N. L. H. Krauss (3, BPBM); Honiara, 200 m, i.1974, N.H. Krauss (6, BMNH). Kukum, 18.x.1965, E.S. Brown (BMNH); Kolombangara Isl., Ringi Coce, Main Road near R6 Road, 16.i.1970, J.E. Tobler (3, CAS); Rossel Island, Arbaleti, 2.x-2.xi.1963, W. W. Brandt (1, ANIC). Malaita Isl., Auki 0–200m, xii.1975, N. L. Krauss (1, ANIC). AUSTRALIA: Queensland: Darnley I.; Yorke I.; Yam I.; Moa I.; Sue Islet; Somerset; Dividing Range 15 km W. of Captain Billy Creek; Iron Range; Port Stewart; Kalpower Crossing; Endeavour R.; Mt Webb Natl Park; Cooktown; Mossman; Julatten; Kuranda; Mission Beach; Leichardt Creek; Palm I., Magnetic I.; Townsville; Bowen; Yeppoon.

Distribution Papua New Guinea, Solomon Islands, and northern Queensland, Australia.

Microcaria mulsanti (Montrouzier, 1861) (Fig. 94)

Coccinella (Daulis) mulsanti Montrouzier, 1861: 304. Type locality: New Caledonia, Art Island. Daulis mulsanti: Mulsant 1866: 211. Microcaria mulsanti: Crotch 1871: 7; Timberlake 1943: 33. Coelophora mulsanti: Crotch 1874: 152; Fauvel 1903: 327; Korschefsky 1932: 294; Chazeau 1978: 63 (lectotype); Pope 1989: 668; Ślipiński 2007: 160; Nattier et al. 2015: 306. Lemnia mulsanti: Iablokoff-Khnzorian 1982: 255. Coelophora ochracea Mulsant, 1866: 262. Type locality: Australia. Synonymised by Crotch 1874: 152.

Diagnosis Microcaria mulsanti is a widely distributed and variable species, but is identified by its relatively large orange body, with black isolated spots or longitudinal fasciae on the elytra (Figs 94A–D).

Description Colour pattern (Figs 94A–G). Head and appendages yellow. Pronotum yellow, with basal and posterior- lateral borders black. Scutellum usually yellow with a dark lateral margin. Elytra background yellow, usually with 10 black spots, or with four longitudinal short stripes at the base and six spots, or in some specimens some spots are missing. Prosternum, mesoand metaventrite, external borders of the elytral epipleuron and central part of the abdomen are black, the remaining parts of the ventral side and appendages are yellowish. Morphology. Length 5–7 mm. Head dorsally mostly covered by the pronotum. Frons 1.7–2.0 times broader than the eye width; surfaces with punctures as large as the eye facets, interspaces reticulate. Lateral borders of the pronotum evenly arcuate. Pronotal punctures as large those on the head, with reticulate interspaces. Prothoracic hypomeron with a shallow fovea anteriorly. Prosternal process with longitudinal carinae extending forward to the middle of the prosternum. Scutellum with a basal border as long as the lateral borders. Elytra slightly broader than the pronotum; punctures much larger than those on the pronotum, interspaces polished and shiny. Elytral epipleuron yellow, with an external black margin, twice as wide as the metanepisternum; fovea for mid and hind femoral apices shallow. Anterior margin of the mesoventrite emarginate medially, posterior margin arcuate. Abdominal postcoxal line (Fig. 94P) meeting at the middle. Posterior margin of ventrite 5 almost truncate in both sexes. Male terminalia (Figs 94H–N). Female terminalia (Fig. 94O). Measurements (n = 8): TL 5.30–6.80 (6.19 ± 0.50); PL 1.00–1.30 (1.18 ± 0.12); EL 4.30–5.60 (5.00 ± 0.42); PW 2.70– 3.50 (3.21 ± 0.29); EW 4.60–5.60 (5.11 ± 0.44); GD 2.50–3.20 (2.84 ± 0.25). Ratios: PL/PW 0.34–0.38 (0.37 ± 0.02); EL/EW 0.93–1.06 (0.98 ± 0.04); EL/PL 3.92–4.67 (4.27 ± 0.29); EL/ GD 1.65–1.92 (1.76 ± 0.08); TL/EW 1.15–1.30 (1.21 ± 0.04).

Types C. mulsanti: 1207 Daulis mulsanti Montrouzier Art Nov. Caledonia (MNHN, lectotype).

92

Ladybird Beetles of the Australo-Pacific Region

Distribution of Microcaria mulsanti

C. ochracea: 28003/ Nov. Holl. Cuming/ Mulsanti Montr. var./ Coelophora ochracea (ZMB, lectotype).

Specimens examined PAPUA NEW GUINEA: Amazon Bay area, Dogon. 2300ft., 13.ix-11.xii.1962, W. W. Brandt (2. ANIC); NE Mt, Kaindi, 2350m, 31.xii.1964, J. Sedlacek (1. BPBM); NEW CALEDONIA: Noumea: Anse Vata, 30m, 8.viii.1979, G. A. Samuelson (2, BPBM); Anse Vata, 20–50m, 20.x.1967, J. & M. Sedlacek (1, BPBM); New Caledonia:3km E of Poindimie, 12.1945, A. E. Milliron (1, BPBM); Noumea (Anse Vata), on citrus, x.1955, J. Rageau (11, CAS); Noumea (Anse Vata),on temple tree, ix.1955, J. Rageau (2, CAS); Noumea, 11.x.1899, Koebele collection (10, CAS); Anse Vata, 1.xi.1958, C. R. Joyce (1, BPBM); Noumea, 11.i.1959, N. L. H. Krauss (1, BPBM); Noumea: Ile martre, vii.1950, N. L. H. Krauss (1, BPBM); Noumea, 28.v.1945, H. E. Milliron (1, BPBM). Loyalty Islands: Mare I., La Roche, iii.1959, N. L. H. Krauss (1, BPBM); Ouvea, Fayaoue, 0–50m, xii.1968, N. L. H. Krauss (1, BPBM); (S) 21.37S/165.49E 450m Col d’Amier (Top of hill), light, 9.ii.2004, M. Wanat (3, ANIC); (S)22.20S/166.58E, 80m Port Boise (Ca. 5 km N of) maquis, light, 14.ii.2004, M. Wanat (1, ANIC); Mont-Koghi, 15.xii.1981, R. Catala (1, NMHN). AUSTRALIA: Queensland: 16.56S-149.11E, NE Herald Cay, 9–11.iv.1994, S. Donaldson (5, ANIC); 16.54S-149.53E, Coringa Islet, 13–15.iv. S. Donaldson (6, ANIC); 16.57S-149.10E, Coral Sea, Herald NE Cay, x.1984, J. Hicks (1, ANIC); 20.50S-153.02E, Jimbooba, QLD, 8.viii.1991, W.Dressier, (2. ANIC); Cairns, J. Sedlacek (1, ANIC); Magnetic I., A. M. Lea (1, SAM). Northern Territory: Tindal, 14.31S-132.22E, light trap, 1–20.xii.1967, W. J. M. Vestjens (1, ANIC); 12.52S-132.50E, Koongarra, 15km E. of Mt. Cahill, 24.v.1973, T. Weir & T. Angeles (1, ANIC); 12.17S-133.20E, Cooper Greek, 11km S. by W. Nimbuwah Rock, 3.vi.1973, Feehan & Upton (1, ANIC); 16.18S-132.27E, Nitmiluk NP., end of first gorge, 19.vi.1992, C. Reid, Beating bushes (1, ANIC); 12.20S-133.19E, Nabarlek Dam, N.T. 15km S. By W. of Nimbuwah Rock, 2.vi.1973, at light, E.G. Matthews (1. ANIC); 12.51S-132.47E, 10km E by N. of Mt. Cahill. 21.v.1973, E. G. Matthews (3. ANIC); 12.23S-132.57E, 5km

NNW of Cahills Crossing, East Alligator River, 28.v.1973, E. G. Matthews (1, ANIC); 12.17S-133.13E, Birraduk Creek, N.T. 18km E by N. of Oenpelli, 4.vi.1973, Feehan & Upton (2, ANIC); 16.08S-136.06E, 22km W of Borroloola, 16.iv.1976, at light, J. E. Feehan (1, ANIC); Cattle Creek, 16.32S-136.10E 54km s by W of Borroloola, 27.x.1975, M. S. Upton (1, ANIC); Port Darwin (1, ANIC); Katherine, 17–18.viii.1973, L. P. Kelsey (1. ANIC); Surprise Creek, 16.25S-136.05E, 45km SW by S of Borroloola, 5.xi.1975, M. S. Upton (1, ANIC); Fanny Bay, Darwin, 3.iii.1967, M.S. Upton (1, ANIC); Batten Point, 15.54S-136.32E, 30km NE by E of Borroloola, 30.x.1975, M. S. Upton (1, ANIC); Goose Lagoon, 16.10S-136.15E, 11km SW by S of Borroloola, 31.x.1975, M. S. Upton (1, ANIC). Western Australia: 14.45S-125.47E, l0km NW by N of Mining Camp, Mitchell Plat., 11–17.v.1983, J. Balderson (1, ANIC); 17.19S-122.10E, 8km S of Cape Bertholet, West Kimberley, 18.iv.1977, D. H. Colless (1, ANIC).

Distribution New Guinea, New Caledonia and northern Australia.

Microcaria salomonensis (Korschefsky, 1940) comb. nov. (Fig. 95)

Coelophora salomonensis Korschefsky, 1940: 1. Type locality. Solomon Islands.

Diagnosis This species can be easily distinguished from other species in the study area by its antenna gradually thickening apically and not bearing a distinct club (Fig. 95J).

Description Colour pattern (Figs 95A–C). Head and head appendages brown. Pronotum with a large black subtrapezoidal median marking extending from the posterior margin to the anterior margin, but often not touching the margin, which is narrowly

VII. Review of the Australo-Pacific Coccinellini

93

Distribution of Microcaria salomonensis

translucent. Scutellum entirely black. Elytra orange, with extensive dorsal black markings not reaching the lateral margins, but narrowly extending towards the scutellum and apex, forming a blackened suture. Ventral side background yellowish, except the metaventrite and median part of abdominal ventrites 1 and 2 are dark pitchy brown to black. Elytral epipleuron and legs yellow. Morphology. Length 5.3–5.7 mm. Head deeply inserted within prothorax, usually only with the clypeus and labrum visible externally. Frons 1.2–1.4 times the eye width, with punctures very shallowly impressed, smaller than the eye facets. Terminal maxillary palpomere weakly securiform (Fig. 95K); antenna 11-segmented without a distinct club (Fig. 95J). Pronotum trapezoidal, widest near the base and distinctly narrowing anteriorly; anterior angles broadly rounded; lateral margins narrowly explanate with a fine bead; disc faintly convex with fine punctation, punctures shallowly impressed, separated by 2–4 diameters; interspaces reticulate. Prosternal process with subparallel carinae extending forward to the middle of the prosternum; hypomeron concave but without a clear fovea. Elytra convex with punctures as large as those on the pronotum; interspace reticulate; lateral margins narrowly explanate with a complete but very fine bead. Epipleuron 3.2– 3.6 times wider than the metanepisternum, shallowly impressed but not foveate. Mesoventrite narrower than the coxal width, with the anterior margin medially deeply emarginate with entire marginal carinae. Abdominal postcoxal line reaching the posterior margin of the ventrite, incomplete laterally. Posterior margin of ventrite 5 almost truncate in both sexes. Male terminalia (Figs 95F–H). Penis consists of a single, simple sclerite (Fig. 95F); penis guide sword-shaped, longer than the very narrow parameres (Figs 95H,I). Female terminalia (Figs 95D,E). Sperm duct very short, broadened at the base forming an unsclerotised infundibulum; spermatheca with the cornu strongly curved and comparably smooth, the ramus short and nodulus indistinct. Measurements (n = 3): TL 5.30–5.75; PL 0.90–1.10; EL 4.30–4.70; PW 2.62–2.82; EW 4.55–4.72; GD 2.10–2.20. Ratios: PL/PW 0.37–0.41; EL/EW 0.90–0.95; EL/PL 3.85– 4.00; EL/GD 1.95–2.01; TL/EW 1.10–1.12.

Types C. salomonensis: British Solomons, March 1932 R.J.A.W. Lever/ Typus/ Syntype/ Pres. by Imp. Inst. Ent. B.M. 1944– 97/ Coelophora salomonensis m. der R. Korschefsky 1938 (BMNH, syntypes).

Specimens examined SOLOMON ISLANDS: Guadalcanal Isl., Honiara, 0–200m, i.1984, N. Krauss (BPBM); same locality (BMNH); same locality Rua Vatu, 22.vi.1954, E.S. Brown (BMNH); same island, Kaukau, 20.viii.1934, R.A. Lever (BMNH); Malaita Isl., Auki, 24.i.1970, J. Tobler (CAS); Florida Is., 1.iv.1944 (CAS).

Distribution Endemic to Solomon Islands.

Remarks The taxonomic placement of this species is uncertain and requires further research. It certainly does not belong to Coelophora in the currently restricted sense. It is here provisionally placed in Microcaria, based on the carinate prosternal process, emarginate anterior margin of the mesoventrite, tibial spur formula 0–2–2, and a shallowly foveate prothoracic hypomeron. However, the morphology of the male genitalia does not support this placement, as both the penis and the tegmen do not conform to the fairly consistent forms typically found in Microcaria.

Microcaria versipellis (Mulsant, 1850) comb. nov. (Fig. 96)

Coelophora versipellis Mulsant, 1850: 394. Type locality: Java. Bielawski and Chûjo 1968: 121. Lemia (Artemis) versipellis: Iablokoff-Khnzorian 1982: 240.

Diagnosis Microcaria versipellis is distinguished from other species treated here by its uniformly orange or reddish body.

94

Ladybird Beetles of the Australo-Pacific Region

Diagnosis This species is easily distinguished from similar Coelophora and Microcaria species by its dorsal colour pattern with six transverse black patches. However, this species can be easily confused with the sympatric species Harmonia novaehebridensis (Fig. 55A), but both species can be distinguished by the generic characters, with Harmonia having a smooth prothoracic hypomeron, mesoventrite with a straight anterior margin and no tibial spurs. Distribution of Microcaria versipellis

Description Colour pattern (Figs 96A–C). Whole body orange or reddish on the ventral and dorsal sides, except for the narrowly black elytral suture; lateral margins of abdomen, head appendages and legs yellowish. Morphology. Head mostly covered by the pronotum from above. Frons flat, 1.2–1.5 times as broad as the eye width. Pronotum moderately convex; anterior border slightly arcuate medially; lateral margins more distinctly arcuate posteriorly than anteriorly; surfaces distinctly punctate with punctures as large as the eye facets; interspaces reticulate. Prothoracic hypomeron with a large fovea anteriorly. Prosternal process with longitudinal carinae slightly convergent anteriorly, ending at 2/3 to the anterior border. Elytra distinctly punctate, punctures variable, mostly larger than those on the pronotum; interspaces smooth and shiny; epipleuron 1.5 times broader than the mesanepisternum, with a shallow fovea. Anterior margin of the mesoventrite strongly emarginate, posterior margin arcuate. Abdominal postcoxal line meets at the middle. Posterior margin of ventrite 5 shallowly emarginate in males and arcuate in females. Male terminalia (Figs 96D–H). Female terminalia (Figs 96I,J). Measurements (n = 4): TL 5.30–6.10; PL 0.95–1.20; EL 3.80–4.50; PW 3.10–3.30; EW 4.70–5.20; GD 2.80–3.10. Ratios: PL/PW 0.30–0.37; EL/EW 0.80–0.91; EL/PL 3.75– 4.00; EL/GD 1.30–1.52; TL/EW 1.12–1.20.

Specimens examined INDONESIA: Ambon Island: v.1908, F. Muir (13, BPBM); 15.iv.1963, A. M. R. Wegner (1, MIIZ); Alt. ca. 70m, at light, 22.ix.1960, A. M. R. Wegner (1, MIIZ).

Distribution The Philippines and Indonesia: Java, Borneo, Ambon.

Remarks The type material of C. versipellis is apparently lost and the interpretation of the species here is based on a series of specimens from Ambon identified by R. Bielawski and deposited in MIIZ.

Microcaria vivida (Crotch, 1874) comb. nov. (Fig. 97)

Coelophora vivida Crotch, 1874: 155; Corrected type locality: Vanuatu. Korschefsky 1932: 298. Coelophora auberti Chazeau, 1978: 66. Type locality: NouvellesHébrides, Ile Pentecôte. Syn. nov.

Description Colour pattern (Figs 97A–C). Head and pronotum orange or reddish, sometimes the lateral edges of pronotum are dark. Scutellum and elytra orange-reddish, each elytron with three black patches or fasciae, the external borders and suture orange-reddish. Ventral side black, except for the orangereddish prothoracic hypomera, meso- and metanepisternum and mesepimeron, and the external borders of the prosternum and abdomen. Legs orange-reddish. Morphology. Length 5.0–5.5 mm. Head deeply inserted in the prothorax with the eyes mostly covered from above. Eyes large; frons flat, 1.3–1.5 times width of the eye; punctures as large as the eye facets, interspaces reticulate. Pronotum weakly convex, lateral borders arcuate with a very fine marginal bead. Prothoracic hypomera shallowly foveate. Prosternal process with lateral carinae convergent anteriorly, ending at halfway to the anterior margin. Scutellum transverse. Elytra epipleura ~2.5 times broader than the mesanepisternum, with a very weak impression for the hind legs. Anterior margin of the mesoventrite deeply emarginate, posterior margin slightly emarginate. Posterior margin of ventrite 5 truncate in both sexes; ventrite 6 shallowly emarginate in males and arcuate in females. Male terminalia (Figs 97D–G). Penis broadened at the middle with the apical third very slim; penis guide as long as the parameres, and evenly tapering from the base to the apex. Female terminalia (Fig. 97H). Spermatheca comparably small and weakly curved; ramus and nodulus distinct; sperm duct slim. Measurements (n = 5): TL 5.10–5.50 (5.30 ± 0.16); PL 1.00–1.10 (1.02 ± 0.04); EL 4.10–4.50 (4.28 ± 0.16); PW 2.60– 2.90 (2.74 ± 0.11); EW 4.30–4.70 (4.50 ± 0.16); GD 2.20–2.60 (2.34 ± 0.17). Ratios: PL/PW 0.34–0.41 (0.37 ± 0.02); EL/EW 0.93–0.98 (0.95 ± 0.02); EL/PL 3.82–4.50 (4.20 ± 0.27); EL/ GD 1.73–1.91 (1.83 ± 0.07); TL/EW 1.15–1.23 (1.18 ± 0.04).

Types C. vivida: New Holland/vivida Cr/Type (BMNH, syntype?). C. auberti: Nouvelles-Hébrides, Ile Pentecôte, Baie Melsisi, 1935–1936, E. Aubert de la Rue (MNHN, holotype).

Specimens examined VANUATU: Efate Vila 0–100m, ii.1970, N. L. H. Krauss (BPBM); Efate I: Klem Butte, 150–200m, 5.xii.1978, N.L.H. Krauss (BPBM); Espiritu Santo Narango, 90m, vi.1960, W.W. Brandt (BPBM); Espiritu Santo I: Luganville. 0–100m, i.1984, N. L. H. Krauss (BPBM); Efate I. (NW): Limestone, Plateau, N. Of Maat, 100m, 20.viii.1957, J.L. Gressit (BPBM); Tanna, ix.1930, L. E. Cheesman, B.M. 1931–30 (BMNH).

Distribution Endemic to Vanuatu.

VII. Review of the Australo-Pacific Coccinellini

95

Distribution of Microcaria vivida

Remarks The type locality of Coelophora vivida given by Crotch as ‘Australia’ is certainly wrong as this species does not occur in Australia. Fortunately, the species is very characteristic and its identity and distribution in Vanuatu can be established without doubt. Accordingly, the type locality of Coelophora vivida Crotch, 1874 is here corrected to Vanuatu. Chazeau (1978) was not aware of the problems with type locality of C. vivida and described this species as new under the name of C. auberti.

Oenopia Mulsant, 1850 (Fig. 98)

Oenopia Mulsant, 1850: 420. Type species, designated by Sicard 1907: 13, Oenopia cinctella Mulsant, 1850. Gyrocaria Timberlake, 1943. Type species by original designation, Coelophora guttata Blackburn, 1892. Synonymised by Iablokoff-Khnzorian 1979: 69. Synharmonia Ganglbauer, 1899: 994. Type species, designated by Chapin 1965: 223, Coccinella conglobata Linnaeus, 1758. Synonymised by Iablokoff-Khnzorian 1982: 398. Protocaria Timberlake, 1943. Type species by monotypy, Protocaria scalaris. Synonymised with Synharmonia by Chapin 1965: 223. Pseudoharmonia Savoiskaya 1969: 37. Type species by monotypy, Pseudoharmonia montana Savoiskaya, 1969. Synonymised by Iablokoff-Khnzorian 1982: 398.

References: Bielawski 1964a: 83–86; Pope 1989: 659; Vandenberg 1996; Poorani 2002b; Poorani and Booth (2005). Diagnostic combination Length 3–5 mm; dorsum convex and glabrous. Elytra black, with yellowish colour pattern of two or four spots (Fig. 98A). Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly shorter than the head capsule with a moderately compact three-segmented club. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron with a fovea near the anterior angles. Prosternal process with distinct carinae extending anteriorly. Anterior

margin of the mesoventrite deeply emarginate medially. Elytral margin with narrow raised and externally thickened margins; epipleuron not foveate. Tibial spurs formula 0–2–2. Abdominal postcoxal line not recurved and incomplete laterally (Fig. 98K); oblique postcoxal line short and hardly visible. Penis consists of a single sclerite with basal capsule strongly developed (Fig. 98E). Ovipositor, coxites with a club-handle shape; infundibulum present; spermatheca sclerotised and bicameral (Fig. 98I).

Description Body broadly oval, strongly convex; dorsum glabrous. Head. Eyes mostly exposed, with inner margins parallel; frons broad, at least twice as long as the eye diameter. Clypeus straight or weakly arcuate between lateral projections. Antenna short; scape not expanded, pedicel slightly narrower than the scape; club compact, terminal antennomere longer than the penultimate. Terminal labial palpomere weakly securiform. Prothorax. Pronotum with an anterior border strongly arcuate medially. Lateral margins narrowly explanate with a distinct bead. Prothoracic hypomeron with a well-delimited fovea; Prosternal process with parallel carinae extending beyond the procoxae. Pterothorax. Scutellum 1/8–1/9 times as broad at the base as the elytra at the shoulders. Elytra slightly broader at the base than the pronotum; the external margins narrowly explanate with complete beads; epipleura strongly descending externally with shallow fossa to receive the hind femora. Mesoventrite broad, at the middle as broad as the coxa diameter; the anterior border shallowly emarginate medially, mesal surface with a deep fossa to receive the prosternal process; the posterior border arcuate medially. Metaventral postcoxal line complete. Middle and hind femur broad and short, completely beneath elytra; tibial spurs present on middle and hind tibiae; tarsal claws with a basal tooth. Abdomen with six ventrites in both sexes; the abdominal postcoxal line reaching the posterior margin of the ventrite, incomplete laterally; the oblique postcoxal line very faint to absent. Posterior margin of ventrite 5 usually broad emarginate in males and truncate in females.

96

Ladybird Beetles of the Australo-Pacific Region

Male terminalia. Penis consists of a single sclerite, strongly constricted apically; the capsule enlarged with the outer arm much longer than the inner arm, the inner arm strongly bent (Fig. 98E). Tegmen short, penis guide shorter than the parameres, deeply emarginate apically (Fig. 98G); parameres swollen apically and densely setose. Female terminalia. Spermatheca sclerotised and bicameral; cornu distinct, longitudinally grooved; nodulus bulbous, connected with the cornu by a slim tube, ramus slim. Sperm duct slim, short; infundibulum short. Coxites slim, broader apically, styli distinct.

Biology Species of Oenopia are polyphagous, but feed mostly on aphids with additional prey including psyllids and plant hoppers (Poorani 2002b).

Distribution A moderately large genus, with ~30 species distributed in Europe, Asiatic Russia, Africa, India, Sri Lanka China, Japan, Philippines, New Guinea and Australia.

Remarks Pope (1989), Vandenberg (1996) and Poorani (2002b) gave detailed accounts of the constitution of this genus and its characteristics. The single species Oenopia guttata, occurring in New Guinea and northern Queensland, is easily recognised by its small rounded body with characteristic maculate colour pattern (Fig. 98A), prothoracic hypomeron with weak foveae, emarginate mesoventrite, and tibial apices with spurs.

Oenopia guttata (Blackburn, 1892)

Gyrocaria minuta Bielawski, 1964a: 88. Type locality: New Britain. Synonymised by Iablokoff-Khnzorian 1982: 400.

Description Colour pattern (Figs 98A–D). Head yellow in males, mostly black in females. Medial part of the pronotum black, lateral parts yellow; scutellum black, elytra black, each with two elytral patches never reaching the external borders or suture; rarely elytra with only an apical yellow patch. Pronotal hypomera yellow, mesepimeron usually yellow, remainder of the underside black; anterior and middle legs usually entirely yellow, hind legs at least in part black. Morphology. Length 3.5–4.0 mm. Eyes almost completely exposed; frons flat, 2.0–2.2 times the eye width. Antenna length half the width of the head capsule; the terminal antennomere obliquely truncate apically. Lateral border of the pronotum weakly arcuate with a sharp angle posteriorly; disc punctate with interspaces densely reticulate and faintly shiny. Prosternal process broad, surface with long parallel carinae almost reaching the anterior border. Scutellum 1/8–1/9 times as broad at the base as the elytra at the shoulders. Abdominal postcoxal line extending to the posterior margin of the ventrite and complete laterally; the oblique postcoxal line very short and faint, sometimes absent. Posterior margin of ventrite 5 broadly emarginate in males, almost truncate in females. Male terminalia (Figs 98E–H). Female terminalia (Figs 98I,J). Measurements (n = 12): TL 3.50–4.00 (3.75 ± 0.16); PL 0.65–0.85 (0.73 ± 0.06); EL 2.80–3.15 (3.02 ± 0.11); PW 2.00– 2.30 (2.17 ± 0.10); EW 3.00–3.50 (3.19 ± 0.17); GD 1.50–1.85 (1.74 ± 0.11). Ratios: PL/PW 0.30–0.37 (0.34 ± 0.03); EL/EW 0.90–1.02 (0.95 ± 0.04); EL/PL 3.71–4.62 (4.13 ± 0.31); EL/ GD 1.68–1.87 (1.74 ± 0.06); TL/EW 1.13–1.27 (1.18 ± 0.04).

Types

(Fig. 98)

Coelophora guttata Blackburn, 1892: 239. Type locality: Australia, N. Queensland. Lea 1902: 490; Weise 1923: 132. Gynocaria guttata: Timberlake 1943: 39; Bielawski 1964a: 83. Oenocaria guttata: Iablokoff-Khnzorian 1982: 400; Pope 1989: 659; Ślipiński 2007: 176.

Distribution of Oenopia guttata

C. guttata: Type/4427/ Coelophora guttata Blackb. (BMNH, lectotype). G. minuta: New Britain, Gazelle Pen., Karawat, 60 m., Aug. 31, 1955, J.L. Gressitt coll. (BPBM, holotype).

VII. Review of the Australo-Pacific Coccinellini

Specimens examined PAPUA NEW GUINEA: Wau, 4000 ft, 24–30.vi.1974, H. Howden (BMNH); Morobe Dist., Herzog Mts., Vagau, 4000 ft, 4–17.i.1965, M.E. Bacchus (BMNH); East High. Prov., Arona Valley, 1300 m, 3.i.1977, E. Smith (BMNH); NE central district, Tapini, 800–1000m, xi.1968 N. L. H. Krauss (3, BPBM); (NE) Wau, Morobe Distr., 1150m, 2.x.1961, J. Sedlacek (1, BPBM); Wareo, Finsch Haven, New Guinea, Rev. L. Wagner (9, SAM); Papua Tapini, 1200m, xi.1961, N. L. H. Krauss (1, BPBM); Western Highlands, Jimi river, 4700ft, 16.vii-21.ix.1961, W. W. Brandt (1, ANIC); (NE) Kundiava 1400–1500m, xi.1971, N. L. H. Krauss (1, BPBM); Amazon Bay area, Komania, 3400ft., 11–26.xi.1962, W. W. Brandt (2, ANIC); Upper Manki L. A., in sticky trap, Bulolo M. Dist., 8.ix.1972, F. R. Wylie & P. Shanahan (1, ANIC); Morobe Prov., Wau, MI. Kaindi, 1300– 1400m, 01.iv.1998, A. Riedel (1. NHMS); New Britain Prov., 30km SW Kokopo, Arabam, 200m 04.35.75s- 152.06.84E. 21.ii-4.iii.2000, A. Weigel (1, NKME); Matupit, Bismark Arch. Ralum, 2.xi.1896, Dahl S. (1, ZMB). SOLOMON ISLANDS: Bougainville, Buin, 11.i.1970, James E. Tobler (1, CAS). AUSTRALIA: QUEENSLAND: Kuranda, 10.viii.1904, Koebele, (BPBM); Julatten, vi.1985, B.P. Moore (ANIC); Redlynch, 20.vii.1986, D. Sands (ANIC); Cairns (ANIC); East Palmerston, v.1964, G. Snowball (ANIC); South Johnstone (ANIC).

Distribution Papua New Guinea, Solomon Islands and northern Queensland, Australia.

Olla Casey, 1899 (Fig. 99)

Olla Casey, 1899: 93. Type species by subsequent designation of Korschefsky 1932: 88, Coccinella abdominalis Say, 1824.

References: Gordon 1985: 826; Vandenberg 1992; 2002. Diagnostic combination

Length 3–6 mm; dorsum convex and glabrous. Elytra grey to yellowish, with a black colour pattern (Fig. 99A). Anterior clypeal border straight between lateral projections. Antenna 11-segmented; slightly longer than the frons between the eye margins, with a compact three-segmented club. Terminal maxillary palpomere strongly securiform. Prothoracic hypomeron without a fovea near the anterior angles. Prosternal process with distinct carinae extending anteriorly. Anterior margin of the mesoventrite straight medially. Elytral margin with narrow margins; epipleuron not foveate. Tibial spurs formula 0–2–2. Abdominal postcoxal line not recurved, complete laterally; oblique postcoxal line short and hardly visible. Penis consists of a single, apically split sclerite and its base with strongly recurved inner arm of capsule (Fig. 99D). Ovipositor, coxites shaped like a club handle; infundibulum absent; spermatheca weakly sclerotised, with a strongly developed ramus and poorly developed nodulus (Fig. 99H); infundibulum absent.

Description Body elongate-oval, convex; glabrous. Head. Eyes almost entirely exposed, inner orbit parallel; frontal width broader than the eye diameter. Anterior border

97

straight between lateral projections. Antenna 11-segmented, much shorter than the head capsule; scape not expanded; club strongly three-segmented, the terminal antennomere slightly expanded. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere strongly securiform. Prothorax. Pronotum convex, glabrous; anterior border deeply emarginate, lateral border weekly acuate, posterior angle blunt. Prosternal hypomera without foveae. Prosternum weekly convex medially, the prosternal process very narrow and rounded apically, surface with distinct carinae. Pterothorax. Mesoventrite with almost straight anterior margin; mesal surface with a shallow fossa, the posterior margin straight. Metaventral postcoxal line straight medially, complete, and strongly curved near the lateral border. Scutellum triangular. Elytra finely punctuated, with the lateral borders very narrowly explanate with a thickened margin. Elytral epipleuron smooth, without a distinct fovea. Legs slim and long, mid and hind tibiae with two apical spurs; tarsal claws each with a subquadrate basal tooth. Abdomen with six ventrites in both sexes. Abdominal postcoxal line separated medially, not recurved; the oblique postcoxal line very fine. Posterior border ventrite 5 strongly arcuate in females and truncate in males. Male terminalia. Penis stout, capsule with the inner arm not recurved, straight and in line with the penis tube, the outer arm strongly arcuate; its apex separated into two pieces. Tegmen with a long capsule; the penis guide broad and bifid apically, the parameres slim and long. Female terminalia. Spermatheca short, with a long ramus, nodulus indistinct. Sperm duct membranous, infundibulum absent. Coxites broad at the base, the external border almost rounded, with slim and long handle.

Biology Olla species are apparently aphid and psyllid predators. Olla v-nigrum has been used a biological control agent against aphids and psyllids, including the Asian citrus psyllid (Diaphornia critri Kuwayama) in Florida (Michaud 2002).

Distribution Five described species are distributed in the New World, from USA to Chile; introduced to Hawaii, New Caledonia and Midway Islands (USA) (Gordon 1985). It has just been found for the first time in Australia (Queensland).

Olla v-nigrum (Mulsant, 1866) (Fig. 99)

Harmonia v-nigrum Mulsant, 1866: 64. Type locality: Mexico. Coccinella v-nigrum: Crotch 1874: 109. Olla v-nigrum: Timberlake 1943: 24; Gordon 1985: 826; Vandenberg 1992: 370; Nattier et al. 2015: 309. Coccinella abdominalis Say, 1824: 95. Type locality: USA, Arkansas.

Description Colour pattern (Figs 99A–C). Head and appendages yellow or light brown, frons with a pair of rounded spots near the base; clypeus and labrum brown. Pronotum pale yellow, with seven black marks; scutellum plate. Elytra pale yellow, with a darker suture, each with eight often irregular spots (Fig. 99A). Ventral surface yellowish brown, except the metanepisternum,

98

Ladybird Beetles of the Australo-Pacific Region

Distribution of Olla v-nigrum

mesepimeron and metepimeron are creamy white. Prothoracic hypomeron and elytral epipleuron are light. Morphology. Length 4.4–6.0 mm. Frons 1.7–1.9 times the eye width. Prosternal process with median carinae converging anteriorly. Elytral epipleuron moderately concave, descending externally, 2.2 times as wide as the metanepisternum. Abdominal postcoxal line (Fig. 99I) reaching the posterior margin of the ventrite; the oblique postcoxal line fine and barely visible. Male terminalia (Figs 99D–F). Female terminalia (Figs 99G,H). Measurements (n = 8): TL 4.40–6.00 (5.16 ± 0.58); PL 0.60–0.90 (0.81 ± 0.11); EL 3.70–5.10 (4.34 ± 0.52); PW 2.30– 2.80 (2.55 ± 0.19); EW 3.60–4.80 (4.10 ± 0.42); GD 1.90–2.30 (2.10 ± 0.17). Ratios: PL/PW 0.26–0.38 (0.32 ± 0.04); EL/EW 1.03–1.09 (1.06 ± 0.03); EL/PL 4.56–6.17 (5.39 ± 0.60); EL/ GD 1.86–2.22 (2.06 ± 0.12); TL/EW 1.21–1.32 (1.26 ± 0.04).

Specimens examined NEW CALEDONIA: Province Sud, Noumea, 12.xi.2001, Johanson, Pape & Viklund (1, NHRS); Province Sud. Fort Teremba, coastal mangrove, 21.xi.2001, Johanson, Pape & Viklund (3, NHRS); 21.42S, 165.41E, 20 m, Moindou (ca. 3 km E of) secondary Leucera bush, 30.i.2004, M. Wanat (2, ANIC); Santo, Vanuatu Shark Bay, Sanma Province, 15.15S 167.05E, general collecting to light, 19.xi.2003, R. Bashford (1, ANIC); (S) 21.42S/165.41E, 20 m Moindou (ca. 3km E of) secondary Leucaena bush, 30.i.2004, M. Wanat (23, ANIC); (S) 22.11S/ 166.30E, Koghi Mts 400–450 m, ex Hibbertia lucens, 11.ii.2004, M. Wanat (5, ANIC); (N) 20.45S/164.53E, Tiendanite 50m, Solanaceae shrubs, 4.ii.2004, M. Wanat (2, ANIC); (N) 20.45S/164.53E, Tiendanite 50m, ad lucem, 4.ii.2004, M. Wanat (1, ANIC); (N) 21.09S/165.19E, Aoupinie 420m stream valley, 7.ii.2004, M. Wanat (1, ANIC); (N) 21.09S/165.19E, Aoupinie 420m-530m, road to sawmill, 7.ii.2004, M. Wanat (1, ANIC); 21.07S/164.57E, 30m Tiea Forest, Sclerophyllous forest, 30.i.2004, M. Wanat (2, ANIC); (S) 22.21S/ 166.58E 0–40 m Port Boise (Gite Kanua) forest at sea shore, 14.02.2004, M. Wanat(1, ANIC); (S) 21.37S/165.49E 450m Col d’Amieu (Top of hill), 09.ii.2004, M. Wanat (1, ANIC).

AUSTRALIA: Queensland: Bowen, 27.ix.2018, Monsour, on Leaucaena infested by psyllids (20, ANIC).

C.

Distribution North America: south-eastern Canada to Florida (US) and west to southern British Columbia (Canada) and southern California (US) (Gordon 1985; Vandenberg 1992, 2002); and from Mexico to Argentina, Puerto Rico and Bahamas. Introduced to Hawaii and New Caledonia. In September 2018 a thriving population of this species was found by Mr Christopher Monsour (Prospect Agriculture Pty Ltd) near Bowen (Queensland) feeding on psyllids on a rivier tamarind (Leucaena leucocephala) (Ślipiński, pers. obs.). The origin of this population is unknown, but it appears that the species is well established in this area.

Phrynocaria Timberlake, 1943 (Figs 100–109)

Phrynocaria Timberlake, 1943: 34. Type species by original designation, Coccinella congener Billberg, 1808. Phrynolemnia Iablokoff-Khnzorian, 1984: 204. Type species by original designation, Coccinella congener Billberg, 1808. Synonymised by Ślipiński 2007: 176.

Diagnostic combination Length 3–5 mm. Dorsum moderately to strongly convex, glabrous. Elytral colour pattern variable. Anterior clypeal margin straight between lateral projections. Antenna long, 11-segmented, with a loosely articulated club. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron with distinct foveae near the anterior angles. Prosternal process with or without admedian carinae. Anterior margin of the mesoventrite deeply emarginate medially. Elytral margin with a very narrow bead; epipleuron with foveae. Tibial spur formula 0–0–0. Tarsal claws with a large basal tooth. Abdominal postcoxal line not recurved; oblique postcoxal line absent. Penis consists of a long basal sclerite and a shorter

VII. Review of the Australo-Pacific Coccinellini

apical section bearing several sclerites. Infundibulum absent. Spermatheca long and C-shaped.

Description Body oval to hemispherical, strongly convex; dorsum glabrous. Head. Eyes large, finely facetted, largely beneath thepronotum, the interior orbits strongly divergent anteriorly. Frons much narrower than the eye diameter; antennae long, 11-segmented, club three-segmented, loosely articulated. Anterior clypeal margin truncate between two lateral projections. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere relatively weakly expanded, securiform. Prothorax. Pronotum moderately convex, lateral borders strongly arcuate with the lateral margins upturned and thickened. Prothoracic hypomeron with distinct foveae. Prosternal process ~0.2 times as broad as the procoxa, the surface with or without longitudinal carinae. Pterothorax. Scutellum large, triangular, ~1/8 times as broad as the elytra at the shoulders. Elytra much broader than the pronotum. Lateral borders narrowly explanate, with a complete but very narrow bead. Elytral epipleuron strongly curved, with shallow foveae to receive the middle and hind femora. Mesoventrite 0.8 times as broad as the coxa diameter, the anterior border deeply notched to receive the apex of the prosternal process. Postcoxal line on the metaventrite joined apically with two or three distinct arches in the medial section, complete laterally. Apex of the middle and hind tibiae without tibial spurs; tarsal claws with a basal subquadrate tooth. Abdomen with six ventrites in both sexes; ventrite 1 more than twice as long as ventrite 2; intercoxal process almost truncate apically, at the middle ~0.2 times the width of the abdomen; postcoxal line joined medially, reaching the posterior

99

margin of ventrite 1, usually complete laterally; oblique postcoxal line absent. Posterior margin of ventrite 5 usually truncate in both sexes. Ventrite 6 broad and very short. Male terminalia. Penis consists of a long basal sclerite, and shorter apical section bearing several sclerites; medial section of the long sclerite is usually with lateral projections. Tegmen with short phallobase; penis guide broad and flat, shorter than the parameres. Female terminalia. Spermatheca long and curved, C-shaped. Ramus indistinct, nodulus forming a short tube. Sperm duct without an infundibulum. Apex of the bursa copulatrix usually lightly sclerotised, forming an apical broad flat tube.

Biology Houston (1988) described larvae of P. astrolabiana and P. gratiosa and summarised the published information on the biology of these species. Most observations associate Phrynocaria with various Coccidae scale insects (Coccus viridis, Pulvinaria and Lecanium), but it was successfully reared on Hyperomyzus aphids under laboratory conditions (Houston 1988). Chazeau (1980) also observed P. quadrivittata feeding on green scale insects, Coccus viridis, in New Caledonia.

Remarks The composition of this genus is uncertain and the relationships between Coelophora, Phrynocaria and Lemnia require further research. However, preliminary molecular research (Tomaszewska et al., unpublished) supports the separate Coelophora and Phrynocaria, but the position of Lemnia remains unresolved. Phrynocaria differs from Coelophora in having a very narrow frons, the middle and hind tibiae without spurs, and a flat penis guide, shorter than the parameres.

Key to the Phrynocaria species of the Australo-Pacific Region 1. – 2(1). – 3(2). – 4(3). – 5(4). – 6(5). – 7(5). – 8(1). – 9(8). –

Prosternal process with distinct admedian carinae extending forward beyond the base of the process ............................................. 2 Prosternal process without admedian carinae ..................................................................................................................................... 7 Large species, length 5.8–6.3 mm........................................................................................................................................................ 3 Smaller species, length less than 5.5 mm............................................................................................................................................. 4 Elytra reddish with a black suture and a single black stripe (Fig. 109A)............................................................................. P. wallacii Elytra black or orange with central black macula (Fig. 107 A–D)......................................................................................... P. moseri Pronotum uniformly orange; elytra black with orange apical spots (Fig. 106A); ventral side and legs yellow or orange .................. ��������������������������������������������������������������������������������������������������������������������������������������������������������������������������� P. kolombangara Pronotum with various black markings, at least with distinctly blackened lateral margins; in forms with an almost uniformly yellow or orange pronotum, at least part of the ventral side and hind legs are dark ............................................................................... 4 At least the femora of hind legs are dark ............................................................................................................................................ 5 Hind legs yellow or orange ................................................................................................................................................................. 6 Male genitalia as in Fig. 105F ..............................................................................................................................................P. gratiosa Male genitalia as in Figs 101D–G ........................................................................................................................................... P. aurita Elytra black, each with a dorsal longitudinal reddish patch, not connected to the suture or lateral margin .................... P. ambonina Elytra black, each with basal and apical reddish patch (Fig. 104A) .....................................................................................P. georgia Pacific species .............................................................................................................................................................. P. quadrivittata Papuan and Australian species ............................................................................................................................................................ 8 Elytra yellow, each with a black sutural stripe and two lateral stripes (Fig. 103A); mesoventrite with a median carina; male genitalia as in Figs 103D–G ..........................................................................................................................................................P. crotchi Elytra variable (Fig. 102), but without a pattern as above; mesoventrite without a median carina or with a short anterior carina only; male genitalia as in Figs 102F–J ........................................................................................................................... P. astrolabiana

Phrynocaria ambonina sp. nov. (Fig. 100) Diagnosis This species can be distinguished by the unique colour pattern on its elytra (Fig. 100A) and its distinctive male terminalia.

Description Colour pattern (Figs 100A–C). Head and appendages yellow in males, frons with darker central patch in females. Pronotum orange-yellow, with a narrow black patch at middle. Scutellum black. Elytra black, with two large orange-yellow patches; elytral epipleuron pale with external borders black. Ventral side yellow except a pale mesepimeron in males; in females

100

Ladybird Beetles of the Australo-Pacific Region

Distribution of Phrynocaria ambonina

the metaventrite and metanepisternum usually partially black. Legs yellow. Morphology. Length 4.0–4.2 mm. Frons narrower than the eye width. Anterior border of the pronotum straight medially, lateral borders strongly and evenly arcuate. Prosternal process with short carinae that end half way to the anterior border. Prothoracic hypomeron with distinct foveae. Elytra strongly convex, weakly explanate laterally, punctures large and deeply impressed; elytral epipleura with foveae. Anterior border of the mesoventrite strongly excavated, the posterior border slightly arcuate. Postcoxal line on metaventrite with two curves medially. Abdominal ventrite 5 truncate apically in both sexes, ventrite 6 truncate in males and arcuate in females. Male terminalia (Figs 100D–G). Female terminalia (Fig. 100H). Measurements (n = 3): TL 4.00–4.20 (4.13 ± 0.12); PL 0.60–0.70 (0.67 ± 0.06); EL 3.30–3.60 (3.47 ± 0.15); PW 2.20–2.40 (2.33 ± 0.12); EW 3.50–3.60 (3.57 ± 0.06); GD 2.00–2.10 (2.03 ± 0.06). Ratios: PL/PW 0.25–0.32 (0.29 ± 0.03); EL/EW 0.94–1.00 (0.97 ± 0.03); EL/PL 4.71–6.00 (5.24 ± 0.68); EL/GD 1.65–1.80 (1.71 ± 0.08); TL/EW 1.14– 1.17 (1.16 ± 0.01).

margins black. Ventral side mostly dark, with prosternum, mesepimeron and mesoventrite lighter, prothoracic hypomera and elytral epipleura yellow. Legs dark, tarsi usually light brown. Morphology. Length 4.8–5.1 mm. Eyes very large and relatively coarsely facetted; frons 0.5–0.7 times the eye width, flat and finely punctured, interspaces densely reticulate and weakly shiny. Pronotum broadest at the base, faintly narrowing anteriorly; anterior angles blunt. Pronotal disc with minute and broadly separated punctures about as large as those on thehead; interspaces finely reticulate, strongly shiny. Prosternal process with converging carinae almost reaching the anterior margin of the prosternum. Mesoventrite deeply emarginate anteriorly. Scutellum triangular, strongly pointed apically, smooth. Elytra broadly circular and convex, disc with a mixture of smaller and larger punctures, interspaces smooth and shiny. Epipleura about twice the width of the metanepisternum, shallowly foveate to receive the legs. Abdominal postcoxal line reaches the posterior margin of the ventrite and is almost complete laterally; ventrite 5 straight in both sexes, ventrite 6 truncate in males, arcuate in females. Male terminalia (Figs 101D–G). Female terminalia (Figs 101H,I). Measurements (n = 2): TL 4.80–5.11; PL 1.09–1.10; EL 3.52–3.64; PW 2.65–2.74; EW 4.14–4.21; GD 2.01. Ratios: PL/PW 0.38–0.41; EL/EW 0.82–0.91; EL/PL 3.21–3.25; EL/ GD 1.56–1.62; TL/EW 1.12–1.21.

Types L. aurita: Holotypus 1982 Lemnia aurita Khnzorian/ New Guinea, Wau McAdam park, 18–24.iv.1965/ Coll. Dr J. Balogh et Dr J.J. Szent-Ivany/ Holotypus Lemnia aurita Khnz. (TMB, holotype male).

Specimens examined

Holotype: INDONESIA: Amboina, xi.1908, F. Muir (BPBM). Paratypes: same data as holotype (1, BPMB; 1, ANIC); same locality and collector, i.07, x.07, no collecting date (3, BPBM).

INDONESIA: Western New Guinea: Dutch New Guinea, Japen I., Mt. Baduri, 1000ft, viii.1938, L.E. Chessman (BMNH). PAPUA NEW GUINEA: (NE) Wau, Morobe Distr., 1200m, 11–20.xi.1961, J. Sedlacek (BPBM); New Britain, Gazelle Pen., Gaulim, 130m, 28.xi.1962, J. Sedlacek (BPBM).

Distribution

Distribution

Known only from Ambon Island, Indonesia.

Endemic to the island of New Guinea.

Specimens examined

Phrynocaria aurita (Iablokoff-Khnzorian, 1982) comb. nov. (Fig. 101)

Lemnia (Microlemnia) aurita Iablokoff-Khnzorian, 1982: 262. Type locality: New Guinea, Wau, McAdam National Park.

Remarks We have examined three female specimens that probably represent a different colour form of P. aurita (Fig. 101C). These specimens differ considerably in having a large central dark

Diagnosis Phrynocaria aurita can be distinguished by its almost entirely orange pronotum and elytra with blackened margins.

Description Colour pattern (Figs 101A,B). Head and head appendages yellow. Pronotum orange or yellow, basal margin black, lateral margins darkened at the basal half; disc with black admedian converging stripes or an entire trapezoidal black marking medially. Scutellum black. Elytra orange, with suture and

Distribution of Phrynocaria aurita

VII. Review of the Australo-Pacific Coccinellini

pronotal area, and most of the elytra being black with yellow or orange maculae. However, without male genitalia and with some variation in spermatheca morphology, we were not able to make a firm identification of these specimens.

Phrynocaria astrolabiana (Weise, 1902) (Fig. 102)

Coelophora astrolabiana Weise, 1902: 492. Type species: New Guinea, Astrolabe Bay. Korschefsky 1932: 290; Iablokoff-Khnzorian 1982: 220. Phrynocaria astrolabiana: Pope 1989: 672; Ślipiński 2007: 178.

Diagnosis Phrynocaria astrolabiana is diagnosed by the prosternal process without admedian carinae, the flat mesoventrite without median carina, a dorsal colour pattern without longitudinal fasciae, and distinctive male terminalia.

101

admedian carinae. Elytra strongly convex, much broader than the pronotum at the base; lateral borders narrowly explanate. Mesoventrite with the anterior border deeply notched medially; posterior border slightly arcuate. Metaventrite with complete discrimen, postcoxal line complete and with two curves medially. Elytral epipleuron with shallow foveae to receive the apices of the hind femora. Posterior margin of abdominal ventrite 5 truncate in both sexes; ventrite 6 emarginate in males and arcuate in females. Male terminalia (Figs 102F–J). Female terminalia (Figs 102K,L). Measurements (n = 5): TL 4.40–5.30 (4.90 ± 0.29); PL 0.60–0.90 (0.88 ± 0.08); EL 3.50–4.52 (4.10 ± 0.32); PW 2.42–2.70 (2.50 ± 0.13); EW 4.12–4.52 (4.23 ± 0.08); GD 2.01–2.54 (2.08 ± 0.19). Ratios: PL/PW 0.25–0.40 (0.33 ± 0.04); EL/EW 0.90–1.02 (0.95 ± 0.05); EL/PL 4.20–6.02 (4.88 ± 0.72); EL/GD 1.75–2.02 (1.88 ± 0.08); TL/EW 1.12–1.21 (1.14 ± 0.03).

Types

Description Body strongly convex. Colour pattern (Figs 102A–E). Colour pattern highly variable, but elytra without a pattern of longitudinal fasciae Frons yellow or reddish; head appendages yellow. Prothoracic hypomeron yellow or reddish. Scutellum black or yellow, with black borders. Mesoventrite usually yellow; metaventrite largely black; mesepimeron pale; mesanepisternum, metanepisternum and metepimeron usually reddish. Elytral epipleuron mostly black with the interior part light coloured. Legs yellow or reddish. Morphology. Length 4–5 mm. Frons ~0.8–1.0 times as broad as the eye width. Eyes relatively very large, the interior orbits strongly divergent anteriorly. Antenna shorter than the head capsule, club loosely articulated; terminal antennomere rounded apically. Pronotum moderately convex, anterior border arcuate medially between the lateral angles, the lateral margin strongly arcuate. Prothoracic hypomeron with distinct foveae. Prosternal process triangular apically, surface without

Distribution of Phrynocaria astrolabiana

C. astrolabiana: N. Guinea, Biro, 1900/ Stephansort, Astrolabe Bay/ Holotypus 1902 Coelophora astrolabiana Weise/ Typus Coelophora astrolabiana Weise Term. Fuz. 25. 1902. 498/ Coelophora astrolabiana Weise (TMB, lectotype).

Specimens examined INDONESIA: Western New Guinea: Noemfor Island, ii.1945, H. H. Blakemore (1, CAS); Cyclops Mts., 600m, 4km nordi Sentani, 8–13.ix.1990, Balke & Hendrich (1, BERLIN); Cyclops Mts, 600m, 4km nordl. Sentani, 8–13.ix.1990, Bailke & Hendrich (1, MNB); Holandia, J. R. Helfer (1, CAS). PAPUA NEW GUINEA: Port Moresby, 13.i.1974, V. Parker (3, AM); Lae, 22.i.1971; R. E. Parrott (1, CNC); (NE) Lae, sea level, 13.vi.1961, J. H. Sedlacek (1, BPBM); NE Wau, Mt. Missim, 950–1300m, 8–9.i.1966 (1, BPBM); Dogura Papua, 25.xi.1955, E. L. Cassidy (1, BPBM); (NE) Sum-Sum, 580m, 64 km N. of Wau, 15.ii.1963, H. W. Clissold (1, BPBM); Misima Island, Louislade Arch., R. J. Andrew (1, SAM); New Guinea:

102

Ladybird Beetles of the Australo-Pacific Region

NE Madang District, Wanuma, 600–720m, viii.1968, N. L. H. Krauss (1, BPBM); Nadzab, Terr. New Guinea,28.v.1944 U.N. Lanham (1, CAS); Oro Bay, H. Chandler (1, CAS); NE Madang District, Wanuma, 600–720m, viii.1968, N. L. H. Krauss (1, BPBM); Admiralty Islands, Los Negros iv.1945, G. E. Bohart (2, CAS). SOLOMON ISLANDS: Guadalcanal, Tenaru River., i.1945, G.E. Bohart (1, CAS). AUSTRALIA: Queensland: Cairns (7, SAM); Cairns Dist., A. M. Lea (2, SAM); Brisbane, ix.1976 (1, ANIC).

Distribution New Guinea; Solomon Islands and north-eastern Australia.

Phrynocaria crotchi Li, Tomaszewska, Pang and Ślipiński, 2014 (Fig. 103)

Chilocorus wallacii Crotch, 1874: 187. Type locality: Batjian. Gordon 1987: 23 (lectotype). Chilocorus wallacei: Korschefsky 1932: 243. Phrynocaria crotchi Li et al., 2014: 597 (replacement name for Phrynocaria wallacei).

Diagnosis Phrynocaria crotchi is very similar to P. wallacii but can be easily distinguished by having smaller size, two black stripes on each elytron, and very distinctive male terminalia.

Description Body broadly oval, convex. Colour pattern (Figs 103A–C). Head and pronotum uniformly yellow or orange. Scutellum black, or brown with black margins. Elytron yellow or orange, with a broad black stripe along the suture that usually does not reach scutellum anteriorly, a median longitudinal stripe and a lateral stripe very

Distribution of Phrynocaria crotchi

close to, but separated from, the lateral margin (Fig. 103C). Background colour of ventral surfaces brown or yellowish brown, but the metaventrite and part of the elytral epipleuron are black; the mesepimeron and metepimeron are pale yellow; the legs and antennae yellow or brown. Morphology. Length 4.6–4.9 mm. Eyes large, finely facetted, the interior orbits strongly arcuate and divergent anteriorly. Frons 0.8–1.0 times the eye width; frontal punctures as large as the eye facets, 1–3 diameters apart; interspaces densely reticulate, weakly shiny. Pronotum convex, broadest shortly behind the middle, not laterally explanate; dorsal punctures about as large as the frontal ones, interspaces reticulate and faintly shiny. Prothoracic hypomeron with a shallow fovea anteriorly. Prosternal process extending beyond the procoxae with the apex triangular; prosternal carinae absent. Elytron much broader than the pronotum at the base; distinctly convex with external margins narrowly explanate; a marginal bead very narrow but complete. Discal punctures of two distinct sizes, the larger ones about as large as those on the pronotum, others distinctly smaller and apparently randomly arranged, interspaces smooth and shiny; punctures along the explanate external margins larger and more deeply impressed than the discal ones. Elytral epipleuron 2.4 times broader than the metanepisternum, with distinct foveae for receiving the mid and hind femorae. Anterior margin of the mesoventrite with a complete carina, deeply emarginate medially to receive the prosternal process; the posterior margin arcuate medially. Metaventrite with a complete discrimen. Abdominal postcoxal lines connected medially (Fig. 103K), reaching the posterior margin of the ventrite and almost reaching the lateral margin, not recurved. Ventrite 5 with a truncate posterior margin in both sexes. Ventrite 6 with posterior margin emarginate medially in males and arcuate in females. Male terminalia (Figs 103D–H). Female terminalia (Figs 103I,J). Spermatheca strongly bent, C-shaped; nodulus narrow and short; ramus indistinct; the sperm duct slim and short. Infundibulum very broad, lightly sclerotised.

VII. Review of the Australo-Pacific Coccinellini

Measurements (n = 4): TL 4.60–4.90 (4.75 ± 0.17); PL 0.70–0.90 (0.78 ± 0.10); EL 3.90–4.10 (3.98 ± 0.10); PW 2.30–2.70 (2.55 ± 0.19); EW 4.10–4.60 (4.35 ± 0.24); GD 2.10–2.50 (2.30 ± 0.18). Ratios: PL/PW 0.28–0.33 (0.30 ± 0.02); EL/EW 0.89–0.95 (0.91 ± 0.03); EL/PL 4.44–5.57 (5.18 ± 0.53); EL/GD 1.60–1.86 (1.73 ± 0.11); TL/EW 1.07– 1.12 (1.09 ± 0.02).

Types C. wallacii: TYPE / TYPE Wallacii Batch/ Coelophora Wallacei Cr (nec Chilocorus) det Korschefsky/ Lectotype Chilocorus wallacei Crotch, 1874 R. Gordon 1987 (UCCC, lectotype).

Specimens examined INDONESIA: Morotai Is, Mollucca Gap., 1945, N. E.I. (CAS); Misool (MNHN).

Distribution Known only from the Indonesian islands of Maluku (Bacan, Morotai, Ternate) and Misool.

Phrynocaria georgia sp. nov. (Fig. 104)

103

males; part of the prosternum and metaventrite dark in females. Legs reddish. Morphology. Length 4.2–4.6 mm. Frons much narrower than the eye width. Anterior border of the pronotum almost straight between lateral angles, lateral borders strongly and evenly arcuate. Surface of the prosternal process with short and weak carinae. Prothoracic hypomera with distinct foveae. Scutellum with a basal border as long as the lateral borders. Elytra strongly convex, weakly explanate laterally. Elytral epipleura with distinct foveae for the middle and hind legs. Anterior border of the mesoventrite deeply and triangularly excavate, the posterior border, postcoxal line on metaventrite with two curves medially. Abdominal postcoxal lines meet at the middle. Posterior margin of ventrite 5 almost truncate in both sexes, posterior margin of ventrite 6 truncate in males and arcuate in females. Male terminalia (Figs 104D–G). Female terminalia (Fig. 104H). Measurements (n = 4): TL 4.20–4.60 (4.40 ± 0.18); PL 0.70–0.80 (0.75 ± 0.06); EL 3.50–3.80 (3.65 ± 0.17); PW 2.20–2.50 (2.38 ± 0.17); EW 3.80–4.20 (3.93 ± 0.19); GD 2.00–2.30 (2.13 ± 0.13). Ratios: PL/PW 0.29–0.33 (0.320 ± 02); EL/EW 0.90–0.97 (0.93 ± 0.03); EL/PL 4.38–5.43 (4.89 ± 0.44); EL/GD 1.65–1.81 (1.72 ± 0.07); TL/EW 1.10–1.15 (1.12 ± 0.03).

Specimens examined

Diagnosis This species can be distinguished by the unique colour pattern on the elytra (Figs 104A–C) and the distinctive male terminalia.

Description Colour pattern (Figs 104A–C). Head and head appendages yellow in males, with darker medial area in females. Pronotum black in females, with a large yellow or orange lateral area in males. Scutellum black. Elytra black, with four large reddish markings, lateral borders and suture black. Ventral side reddish, except for the black external borders of elytral epipleura in

Distribution of Phrynocaria georgia

Holotype: SOLOMON ISLANDS: British Solomons: Kolombangara Isl., Ringi Cove. Main Road near R6 Road, 17.i.1979, James E. Tobler (CAS). Paratypes: same data as holotype (1, CAS); British Solomons: Kolombangara Isl., Ringi Cove. Main Road near R6 Road, 16.i.1979, James E. Tobler (2, CAS); Solomon Is. New Georgia: Monda 0–200 m, xi.1925, N.L.H. Krauss (2, BPBM).

Distribution Known only from the Solomon Islands.

104

Ladybird Beetles of the Australo-Pacific Region

Phrynocaria gratiosa (Mulsant, 1853) (Fig. 105)

Coelophora gratiosa Mulsant, 1853: 187. Type locality: Australia. Mulsant 1866: 276; Crotch 1874: 155; Blackburn 1894: 238. Phrynocaria gratiosa: Timberlake 1943: 35; Pope 1989: 670; Ślipiński 2007: 178. Lemnia (Phrynocaria) gratiosa: Iablokoff-Khnzorian 1984a: 211. Phrynocaria gratiosa var. flavoguttata Timberlake, 1943: 35. Type locality: Australia: Queensland, Bundaberg. Phrynocaria gratiosa var. nigrocincta Timberlake, 1943: 36. Type locality: Australia: Queensland, Bundaberg. Phrynocaria gratiosa var. palens Timberlake, 1943: 36. Type locality: Australia: Queensland, Brisbane. Phrynocaria gratiosa var. koebelei Timberlake, 1943: 37. Type locality: Australia: Queensland, Bundaberg. Coelophora elegans Crotch, 1874:156. Type locality: Gabun. Synonymised by Houston 1983: 19. Autotela elegans: Korschefsky 1932: 299. Coelophora nigrovittata Blackburn, 1895: 237. Type locality: Australia: Queensland, near Cairns. Synonymised by IablokoffKhnzorian 1979: 62. Phrynocaria gratiosa var. nigrovitata: Timberlake 1943: 37.

Diagnosis Phrynocaria gratiosa is a very variable and difficult to diagnose species. It is known from Australia only, where it forms colour patterns identical to species classified in other genera (e.g. Microcaria, Coelophora), but it can be easily distinguished by its narrowly separated eyes. However, it can be reliably differentiated from other Phrynocaria species by the male terminalia only.

Description Colour pattern (Figs 105A–E). Dorsal and ventral colour pattern variable, but legs usually with at least some parts darkcoloured, especially the femora and hind legs. Morphology. Length 4.5–5.4 mm. Head with eyes separated by about the eye width. Pronotum with lateral margins usually each with shallow foveae; disc convex, with punctures of varying size and separation. Elytra with discal punctures similar to those of the pronotum; punctures along the explanate external margins usually more deeply impressed than those elsewhere, and so appearing larger and more closely set, intervals smooth, shining. Pronotal hypomeron with distinct foveae. Abdomen

with apical margin of ventrite 6 shallowly emarginate medially in males, evenly arcuate in females. Male terminalia (Figs 105F–K). Female terminalia (Fig. 105L). Measurements (n = 5): TL 4.50–5.40 (4.95 ± 0.29); PL 0.70–0.90 (0.85 ± 0.08); EL 3.60–4.50 (4.10 ± 0.30); PW 2.40– 2.80 (2.60 ± 0.13); EW 4.10–4.60 (4.33 ± 0.18); GD 2.00–2.50 (2.18 ± 0.19). Ratios: PL/PW 0.27–0.38 (0.33 ± 0.04); EL/EW 0.88–1.02 (0.95 ± 0.05); EL/PL 4.00–6.14 (4.88 ± 0.72); EL/ GD 1.80–2.00 (1.88 ± 0.08); TL/EW 1.10–1.19 (1.14 ± 0.03).

Types C. gratiosa: TYPE / N. Holl./ A/ Type gratiosa Deyr (UCCC, lectotype). C. elegans: Type/ 57.71/ Coelophora elegans Crotch (Type)/ Elegans Gehin Gabon/ Elegans Crotch TYPE (BMNH, lectotype). C. nigrovittata: Type/ 5898 T N. Qu./ Coelophora nigrovittata, Blackb (BMNH, lectotype).

Specimens examined AUSTRALIA: Queensland: Bundaberg, 19.x.1904 (1, BPBM); Brisbane, 7.viii.1962, J. Cribb (1, UQIC); Nat. Park, 26.v.196 2(1, UQIC). New South Wales: ‘LORIEN’ 3km N Lansdowne Taree, ex r/f margin, Malaise trap, 27.xii-03.i.1988, G. Williams (1, ANIC); Swansea, xi.1950, J. Sedlacek (2, BPBM); Ashfield, 25.i.1980, D. A. Doolan (2, AM; 1,ANIC); Wiangaree S.F. 1050 m, 10–12.ii.1982, T. Weir & A. Calder (2, ANIC).

Distribution Known only from Queensland and New South Wales, Australia.

Phrynocaria kolombangara sp. nov. (Fig. 106) Diagnosis The dorsal colour pattern of this Phrynocaria species resembles some colour variations of Harmonia incognita and Coelophora jansoni. However, it can be distinguished from these two species by its very narrow frons. It can be distinguished from other species of Phrynocaria by its distinctive colour pattern (Figs 106A–C) and the male terminalia.

Description

Distribution of Phrynocaria gratiosa

Colour pattern (Figs 106A–C). Head and appendages orangereddish. Pronotum entirely orange-reddish. Scutellum orangereddish. Elytra background colour black, with two orange-red patches apically, and lateral borders usually orange-reddish. Ventral side completely orange-reddish, in some specimens the external borders of elytral epipleura and the metaventrite are dark. Morphology. Length 5.1–5.7 mm. Frons much narrower than the eye width. Antenna almost as long as the head capsule, terminal antennomere oval. Anterior border of the pronotum slightly arcuate medially, lateral borders strongly and evenly arcuate. Prothoracic hypomera with distinct foveae. Surface of the prosternal process with longitudinal carinae convergent anteriorly. Elytra strongly convex, narrowly explanate laterally, with large and deeply impressed punctures; elytral epipleura

VII. Review of the Australo-Pacific Coccinellini

105

Distribution of Phrynocaria kolombangara

shallowly foveate. Anterior border of the mesoventrite deeply and triangularly excavate; the postcoxal line of the metaventrite with two curves medially. Abdominal postcoxal line incomplete laterally (Fig. 106I). Posterior margin of ventrite 5 slightly arcuate in both sexes, the margin of ventrite 6 truncate in males and arcuate in males. Male terminalia (Figs 106D–G). Female terminalia (Fig. 106H). Measurements (n = 6): TL 5.10–5.70 (5.47 ± 0.23); PL 1.00–1.10 (1.02 ± 0.04); EL 4.10–4.70 (4.45 ± 0.23); PW 2.90– 3.20 (3.07 ± 0.10); EW 4.70–5.20 (5.02 ± 0.17); GD 2.20–2.60 (2.48 ± 0.16). Ratios: PL/PW 0.31–0.35 (0.33 ± 0.02); EL/EW 0.86–0.92 (0.89 ± 0.03); EL/PL 3.91–4.70 (4.38 ± 0.32); EL/ GD 1.73–1.86 (1.79 ± -0.05); TL/EW 1.06–1.12 (1.09 ± 0.02).

Specimens examined Holotype: SOLOMON ISLANDS: Kolombangara Isl., Rangi Cove, Main road near R6 road, 16.i.1979, James E. Tobler (CAS). Paratypes: same data as holotype (5, CAS; 2, ANIC); same locality and collector but 17.i.1979 (3, CAS); Southern coast of Kolombangara Is. New Georgia Group Solomon Islands I-X.1990, Fairlamb Family (1, ANIC); British Solomons: Malaita Isl., Auki, 24.i.1970, James E. Tobler (1, CAS); Solomon Is., New Georgia Group, Gizo I. 70 m, 18.vii.1964 (1, BPBM); Solomon Island, Guadalcanal, 24.viii.1984, R. McFairlane, Acc no. 07078, feeding on maize stem (1, BMNH); Solomon Is. Guadalcanal, Honiara Dist., Kukum, 29.x.1950, E.S. Brown (2, BMNH); same locality, 13.ii.1964, M.McQuillan (1, BMNH); same locality, 28.x.1963 and 26.i.1965, P. Greenslade (2, BMNH).

Distribution Known from the Solomon Islands only.

Phrynocaria moseri (Weise, 1902) comb. nov. (Fig. 107)

Coelophora moseri Weise, 1902: 500. Type locality: Misool. Korschefsky 1932: 294; Bielawski 1965: 225. Lemnia moseri: Iablokoff-Khnzorian 1982: 259 (lectotype).

Diagnosis The typical colour form of Phrynocaria moseri can be distinguished by the large black macula on reddish elytra (Fig. 107A), and the melanic form with extensive lateral yellow spots on an otherwise entirely black pronotum.

Description Colour pattern (Figs 107A–D). Typical form: body dorsally and ventrally rusty-brown; scutellum black or with black margins; elytra with a large central black marking along the suture extending forward to the scutellum and posteriorly to the elytral apices; sometimes the margins of the pronotum and elytra are more or less blackened. Melanic form: head, appendages and venter brown; pronotum black, with yellow anterolateral areas of various sizes; scutellum and elytra uniformly black. Morphology. Length 6.2–7.0 mm. Frons 1.0–1.3 times as broad as the eye width; eyes almost completely covered by the anterior, mostly transparent, part of the pronotum, interior orbits convergent. Anterior border of the pronotum slightly arcuate medially; lateral margins arcuate, slightly sinuate before broadly rounded anterior angles; disc weakly convex, rather densely punctate, interspaces reticulate. Prothoracic hypomeron with a well-defined fovea. Prosternal process with parallel carinae extending almost to the anterior margin of the prosternum. Anterior border of the mesoventrite strongly notched medially; metaventral postcoxal lines recurved and complete. Scutellum triangular, pointed apically, 1/12 of elytral width. Elytral epipleuron 1.8–2.0 times as broad as the metanepisternum with distinct foveae to receive the apices of the hind legs. Abdominal postcoxal line extending to the posterior margin of the ventrite, almost complete laterally. Posterior margin of ventrite 5 truncate in males and slightly arcuate in females. Male terminalia (Figs 107E–I). Female terminalia (Fig. 107J). Measurements (n = 4): TL 6.20–7.00 (6.68 ± 0.39); PL 1.00–1.45 (1.26 ± 0.21); EL 5.00–5.50 (5.38 ± 0.25); PW 3.50– 3.65 (3.54 ± 0.07); EW 5.60–6.10 (5.83 ± 0.22); GD 3.10–3.50 (3.26 ± 0.17). Ratios: PL/PW 0.29–0.40 (0.36 ± 0.05); EL/EW 0.89–0.96 (0.92 ± 0.03); EL/PL 3.79–5.50 (4.35 ± 0.78); EL/ GD 1.56–1.77 (1.65 ± 0.10); TL/EW 1.11–1.19 (1.15 ± 0.03).

106

Ladybird Beetles of the Australo-Pacific Region

Phrynocaria quadrivittata (Fauvel, 1903) comb. nov. (Fig. 108)

Coelophora quadrivittata Fauvel, 1903: 327. Type locality: New Caledonia, Kanala. Korschefsky 1932: 296; Chazeau 1978: 65 (lectotype); Chazeau 1980; Nattier et al. 2015: 306.

Diagnosis

Distribution of Phrynocaria moseri

Phrynocaria quadrivittata is similar to some colour variants of Coelophora mulsanti from New Caledonia, but it can be distinguished by its smaller body size and much narrower frons between the eyes.

Types

Description

C. moseri: Misool Insel/ moseri m. (ZMB, lectotype).

Specimens examined PAPUA NEW GUINEA: Daradae, nr. Java, Musgrove R. 100m, 4.x.1958, J. L. Gressitt (1, ANIC); Wau, L. Wagner (1, SAM); SE Popondetta Dist., Jumbora, 19.ix-15.x.1963, P. Shanahan (1, ANIC); Malahang near Lae, Morobe District, 3–5.xii.1972, G. A. Holloway (1, AM); Sattelberg, Wahnes. Franklin Muller (3, Berlin); Sapphire Creek, Port Moresby, 19.viii.1973, R. Mackay (1, AM); INDONESIA: KAI ISLANDS: Key Inseln, Schroder (2, DEI); Ins. Kei (Planten) (1, MIIZ);

Distribution New Guinea, Misool and Key Islands.

Remarks Coelophora varicolor Crotch, 1874: 155 described from ‘Batchian’ also belongs to Phrynocaria comb. nov., and is probably a senior synonym of this species. However, we were not able to confirm this synonymy because the holotype of C. varicolor is a badly damaged female without spermatheca and no matching male specimens from the type locality were available.

Distribution of Phrynocaria quadrivittata

Colour pattern (Figs 108A–C). Typical form: head and appendages yellow; pronotum yellow, with the posterior part of the lateral borders narrowly black. Scutellum yellow or with black borders. Elytra yellow, with four short longitudinal stripes at the base, and four additional small black spots, the external borders and suture black. Ventral side yellow, except for dark brown meso- and metaventrite and central part of abdomen. Melanic form: pronotum almost black, elytra with broad black external margins and suture, each elytron with two complete longitudinal black stripes. Morphology. Length 4.4–5.3 mm. Frons 0.8–1.0 times as wide as the eye width; eyes almost completely covered by the partially transparent anterior part of the pronotum. Pronotum with the anterior border almost straight medially between anterior angles; lateral borders more strongly arcuate posteriorly. Disc relatively flat and finely punctate; interspaces smooth and shiny. Prosternal process without carinae. Scutellum smooth, 1/12 of elytral width at the base. Elytra strongly convex, weakly explanate laterally, with weak beads; disc with punctures similar to these on the pronotum; interspaces smooth and shiny; elytral epipleura strongly descending, with shallow foveae. Anterior border of the mesoventrite moderately emarginate medially. Postcoxal line on the metaventrite with two lobes. Abdominal postcoxal line joined medially, almost meeting the posterior margin of the ventrite (Fig. 108J). Posterior margin of ventrite 5 almost

VII. Review of the Australo-Pacific Coccinellini

truncate in both sexes, the posterior margin of ventrite 6 truncate in males and arcuate in females. Male terminalia (Figs 108D–G). Female terminalia (Figs 108H,I). Measurements (n = 6): TL 4.40–5.30 (4.80 ± 0.40); PL 0.70–0.90 (0.82 ± 0.10); EL 3.70–4.40 (3.98 ± 0.33); PW 2.30–2.60 (2.48 ± 0.13); EW 3.80–4.60 (4.13 ± 0.33); GD 1.90–2.40 (2.17 ± 0.18). Ratios: PL/PW 0.29–0.35 (0.33 ± 0.02); EL/EW 0.93–0.98 (0.96 ± 0.02); EL/PL 4.11–5.43 (4.91 ± 0.46); EL/GD 1.68–1.95 (1.84 ± 0.09); TL/EW 1.15– 1.18 (1.16 ± 0.01).

Specimens examined NEW CALEDONIA: 22.20S/165.58E 80m Port Boise (Ca. 5 km N of) maquis ad lucem, 14.ii.2004, M. Wanat (2, ANIC); Thio, 11.xi.1958, C. R. Joycs (1, BPBM). FIJI: Fiji, ii.1960, N. L. H. Krauss (7, BPBM); Viti Levu Suva, xi.1957, N. L. H. Krauss (1, BPBM).

Distribution New Caledonia and Fiji.

Remarks Chazeau (1980) studied the inheritance model of polymorphism in the elytral colour pattern of this species in New Caledonia. He also recorded it as a predator of the soft scale, Coccus viridis (Green, 1889) (Hemiptera: Coccidae), feeding on frangipani (Plumeria sp.).

Phrynocaria wallacii (Crotch, 1874) (Fig. 109)

Coelophora wallacii Crotch, 1874: 156. Type locality: Batjan Island. Gordon 1987: 19 (lectotype). Coelophora wallacei: Korschefsky 1932: 298. Lemnia wallacei: Iablokoff-Khnzorian 1982: 260. Phrynocaria wallacii: Li et al. 2014: 593.

Distribution of Phrynocaria wallacii

107

Diagnosis Phrynocaria wallacii is easily distinguished from the externally similar P. crotchi by its superior size, a single longitudinal stripe on each elytron, and its very distinctive male terminalia.

Description Colour pattern (Figs 109A–C). Head and pronotum uniformly yellow or orange. Scutellum orange, with darker apical margins. Elytra yellow or orange, with a black narrow stripe along the suture and a much wider lateral stripe. Ventral surfaces brown or yellowish brown; mesepimeron pale; legs and antennae yellow or brown. Morphology. Length 5.8–6.3 mm. Frons between eyes about as broad as eye width; frontal punctures as large as the eye facets, 1–3 diameters apart; interspaces densely reticulate, faintly shiny. Antenna about twice as long as the frons width. Pronotum moderately convex, broadest at, or shortly behind, the middle, not laterally explanate; dorsal punctures as large as the frontal ones, interspaces reticulate and faintly shiny. Anterior pronotal margin arcuate medially; lateral margins arcuate with a very narrow but complete lateral bead, continuing along the anterior margin to the level of the inner margins of the eyes, and to the hind angles posteriorly. Prothoracic hypomeron with a distinct fovea anteriorly. Prosternal carinae converging weakly anteriorly, ending at ~2/3 of the way to the anterior margin. Scutellum forming an equilateral triangle, acute apically. Elytron moderately convex, much broader than the pronotum at the base; distinctly convex with the external margins narrowly explanate; the marginal bead very narrow and not very distinct, but complete. Discal punctures of two distinct sizes, larger ones about as large as those on the pronotum, the others much smaller and apparently randomly arranged, interspaces with distinct reticulation; punctures along explanate external margins larger and more deeply impressed than the discal ones. Elytral epipleuron with distinct foveae receiving the mid and hind femora. Anterior margin of the mesoventrite with a raised carina, deeply emarginate medially. Metaventral postcoxal line broadly emarginate medially, and complete laterally.

108

Ladybird Beetles of the Australo-Pacific Region

Abdominal postcoxal lines connected medially, reaching the posterior margin of the ventrite and almost reaching the lateral margin. Ventrite 5 with a posterior margin truncate in both sexes. Ventrite 6 with a posterior margin emarginate medially in males, rounded in females. Male terminalia (Figs 109D–H). Female terminalia (Figs 109I,J). Measurements (n = 6): TL 5.80–6.30 (6.08 ± 0.17); PL 1.00–1.20 (1.10 ± 0.06); EL 4.70–5.20 (4.98 ± 0.16); PW 3.30– 3.50 (3.42–0.10); EW 5.30–5.70 (5.50 ± 0.15); GD 2.60–3.00 (2.80 ± 0.17). Ratios: PL/PW 0.30–0.35 (0.32 ± 0.02); EL/EW 0.87–0.94 (0.91 ± 0.03); EL/PL 4.17–5.00 (4.54 ± 0.30); EL/ GD 1.67–1.93 (1.78 ± 0.10); TL/EW 1.07–1.15 (1.11 ± 0.03).

Types C. wallacii: TYPE / TYPE Wallacii Batc/ Batch/ Lectotype Coelophora wallacei Crotch, 1874 Gordon 1987’ (UCCC, lectotype).

Specimens examined INDONESIA: BACAN: Batjan, Doherty, C. E. Tottenham Collection, B. M. 1969–77 (BMNH); Batjan Is., Moluccas, 3.xii.1939, R. G. Wind (CAS). TERNATE: Ternate, Raffray & Maindron, 1878 (MNHN). HALMAHERA: Dodinga, Doherty, VIII, C. E. Tottenham Collection, B.M. 1969–77 (BMNH). KAI ISLANDS: Kai/ D/ Paralectotype Coelophora wallacii Crotch Gordon 1987 (UCCC, Paralectotype).

Distribution Indonesia: North Maluku and Kai Islands.

Subepilachna Bielawski 1963b gen. res. (Fig. 110)

Subepilachna Bielawski, 1963b: 445. Type species by original designation, Subepilachna latemarginata Bielawski, 1963b. Singhikalia: Miyatake 1972: 92; Jadwiszczak 1990: 56; Ren et al. 2009: 235.

Diagnostic combination Length 7.2–7.9 mm; body elongate-oval, moderately convex, distinctly pubescent. Elytra uniformly black. Antenna shorter than head width; antennal club distinct, loosely articulated. Anterior clypeal margin straight between lateral projections. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere weakly securiform. Prothoracic hypomeron with a shallow fovea near the anterior margins. Prosternal process with short carinae. Anterior border of mesoventrite deeply emarginate medially, with raised ridge. Tarsal claws with a subquadrate basal tooth. Abdominal postcoxal lines separate medially, incomplete laterally, not recurved; oblique dividing line absent. Penis consists of two sclerites. Infundibulum in female terminalia absent.

Description Body elongate-oval, moderately convex; dorsum distinctly pubescent (Fig. 110A). Head. Slightly withdrawn into prothorax; eyes largely exposed, the inner orbits slightly divergent anteriorly. Antenna shorter than the head width; scape slightly broadened interiorly, pedicel as long as, and slightly broader than, antennomere 3;

the antennal club three-segmented, loosely articulated; terminal antennomere rounded apically and longer than the penultimate. Clypeus straight between lateral projections. Mandible (Fig. 110K) bifid apically; the molar part with a basal tooth. Terminal maxillary palpomere weakly securiform (Fig. 110J). Prothorax. Pronotum weakly convex; anterior border shallowly emarginate; lateral border evenly arcuate, narrowly expanded without a marginal bead. Prothoracic hypomeron concave, but without a delimited fovea near the anterior angles. Prosternum raised anteriorly and forming a median knob; prosternal process flat, extending beyond the coxae and somewhat triangular apically; surface with indistinct lateral carinae. Pterothorax. Mesoventrite width 0.7 times as times the coxa width; surface weakly raised medially; the anterior margin deeply emarginate medially, with a raised but incomplete medial ridge, the posterior margin slightly arcuate. Discrimen almost complete. Metaventral postcoxal lines complete laterally. Scutellum triangular. Elytra slightly broader than the pronotum at the base; external borders narrowly explanate, without a marginal bead. Elytral epipleuron strongly inclined, without delimited foveae. Legs slim and long; tibia spur formula 0–2–2 but spurs very short and not well visible; outer margins of mid and hind trochanters smooth, without acute angle; tarsal claws with a subquadrate basal tooth. Abdomen with six ventrites in both sexes; intercoxal process narrow. Postcoxal line (Fig. 110L) short, incomplete laterally, not recurved; oblique postcoxal line absent. Posterior margin of ventrite 5 truncate in males and narrowly arcuate medially in females. Male terminalia. Penis consists of two sclerites (Fig. 110D); the terminal sclerite relatively long and spiral-shaped (Fig. 110F). Tegmen slim with a short phallobase; penis guide strongly curved apically, the parameres slim and long. Female terminalia. Spermatheca strongly curved, C-shaped and slim; the cornu deeply annulate (Fig. 110H); nodulus quite small, ramus distinct; sperm duct short attached to the apex of bursa copulatrix; infundibulum absent; coxites broad at the base, with distinct apical styli (Fig. 110I).

Biology The biology and immature stages of this genus are unknown, but adult specimens of Subepilachna and Singhikalia are mimicking co-occurring members of the herbivorous tribe Epilachnini.

Distribution This genus consists only of single species from Papua New Guinea.

Remarks Jadwiszczak (1990) synonymised Subepilachna with the Oriental genus Singhikalia Kapur, 1963 based on descriptions of both genera sharing many external similarities. However, after study of representatives of the Asian and Papuan species, we recognise both genera as valid based on pronounced differences in the male and female terminalia and external morphology. Externally, Subepilachna differs from Singhikalia by having a prosternum with a distinct median tubercle in the middle of the anterior margin, the mesoventrite is narrow and not bordered medially, the elytral margins are explanate but without a marginal bead and the elytral epipleuron is very broad and strongly descending. In contrast to Singhikalia,

VII. Review of the Australo-Pacific Coccinellini

the penis of Subepilachna consists of two distinct sclerites, and the spermatheca is strongly annulate. Subepilachna and Singhikalia are the most unusual genera in Coccinellini, with distinct dorsal pubescence and the colouration mimicking local members of the tribe Epilachnini. They can be distinguished from Epilachnini in having apically bifid instead of multidentate mandibles, tarsal claws with a basal tooth and lateral antennal insertions much closer to the eyes.

109

8–10.xi.1965 (1, BPBM); NE Wau, 1750–1900m, 19.viii.1965, J. & M. Sedlacek (1, BPBM); Bismark Gbg., Waghital, 5–7000 ft, 1963, Cl. Voss (1, NHMB). Tari subdistrict, Tigibi, native gardens, 1600m, 28.v-14.vi.1966. W. Vink (1, NBCL); Finsch Haven Dist, L. Waguei (1, SAM); Madang Distr. Finisterre Mts, 5500 ft., Moro, 30.x-15xi.1964, M.E. Bacchus (BMNH).

Distribution Endemic to Papua New Guinea.

Subepilachna latemarginata Bielawski, 1963b (Fig. 110)

Subepilachna latemarginata Bielawski, 1963b: 445. Type locality: Papua New Guinea, Hagen.

Description Colour pattern (Fig. 110A–C). Frons mostly yellow with median black area in males, mostly black with yellow area laterally in females. Labrum brown. Head appendages brown. Pronotum black, with two large yellow spots anterolaterally. Prosternum black, with a small yellow area close to the prothoracic hypomeron. Prothoracic hypomeron black, with a yellow area corresponsing to the pronotum spots. Scutellum, elytra and abdomen black. Dorsal side densely covered by white or yellowish hair. Morphology. Length 7.1–7.9 mm. Frons 1.8–2.0 times the eye width. Prosternal process triangular apically, the surface with weak but almost complete carinae extending forward to the medial projection. Mesoventrite with anterior margin deeply emarginated medially, the posterior margin straight. Abdominal postcoxal line (Fig. 110L) approaches the posterior margin of ventrite. Posterior margin of ventrite 5 almost truncate in males, narrowly arculate in females. Male terminalia (Figs 110D–G). Female terminalia (Figs 110H,I). Measurements (n = 5): TL 7.20–7.90 (7.56 ± 0.34); PL 1.10–1.20 (1.18 ± 0.04); EL 5.80–6.50 (6.12 ± 0.31); PW 3.20– 3.70 (3.40 ± 0.19); EW 5.80–6.50 (6.06 ± 0.26); GD 2.90–3.50 (3.16 ± 0.24). Ratios: PL/PW 0.32–0.36 (0.35 ± 0.01); EL/EW 0.98–1.07 (1.01 ± 0.03); EL/PL 4.83–5.42 (5.19 ± 0.26); EL/ GD 1.86–2.03 (1.94 ± 0.06); TL/EW 1.20–1.32 (1.25 ± 0.05).

Types S. latemarginata: New Guinea: (NE) W. Highlands: Hagen, SE of Korn Farm, 15 Oct. 1958, J.L. Gressitt (BPBM, paratype).

Specimens examined PAPUA NEW GUINEA: Miwaka, viii.1975, H. Ohlmus (1, ANIC); NE Iongai, 10km E of Mt. Albert Edward, 1800–1850m,

Distribution of Subepilachna latemarginata

Synona Pope, 1989 (Fig. 111)

Synia Mulsant, 1850: 375. Type species by subsequent designation of Crotch 1874: 177: Synia melanaria Mulsant, 1850 [junior homonym]. Lemnia (Synia): Iablokoff-Khnzorian 1982: 232. Synona Pope, 1989: 660 (replacement name).

References: Mulsant 1866: 248; Crotch 1874: 177; Korschefsky 1932: 276; Timberlake 1943: 56; Bielawski 1964c: 23; Iablokoff-Khnzorian, 1979: 74. Ślipiński, 2007: 179; Poorani et al. 2008. Diagnostic combination

Length 5.0–7.8 mm; body broadly oval, convex, glabrous. Elytra uniformly black. Antenna shorter than the head width; antennal club distinct, tightly articulated and with terminal segment truncate apically (Fig. 111J). Anterior clypeal margin emarginate between lateral projections. Mandible bifid apically, the molar part with a basal tooth. Terminal maxillary palpomere distinctly securiform. Prothoracic hypomeron with a distinct fovea near the anterior margins. Prosternal process with short carinae. Anterior border of the mesoventrite deeply emarginate medially, with a raised ridge. Tarsal claws with a subquadrate basal tooth. Abdominal postcoxal lines separate medially, incomplete laterally, not recurved; an oblique postcoxal line present. Tibial spur formula 0–2–2. Penis consists of a single sclerite. Infundibulum in female terminalia weakly sclerotised not tubular.

Description Body circular to broad oval, only slightly longer than broad, dorsum hemispherical and strongly convex, glabrous (Figs 111A–D). Head with anterior clypeal margin distinctly and arcuately emarginate between lateral projections (Fig. 13A). Terminal maxillary palpomere securiform. Antenna (Figs 13V, 111J) short, ~1.5 times as long as the head width, scape flattened and arcuate exteriorly, club three-segmented and compact, the apical margin of terminal antennomere obliquely transverse. Prothorax. Pronotal hypomera with conspicuous foveae on anterior inner corners; prosternal carinae present basally, reaching up to the middle or into the anterior half. Pterothorax. Scutellum large, triangular. Elytra laterally narrowly explanate, with a prominent marginal bead. Elytral epipleuron broad, moderately to strongly reflexed, deeply foveolate at the level of the middle and hind legs; inner carina apically incomplete, reaching up to the fifth abdominal ventrite. Anterior margin of the mesoventrite deeply emarginate medially, the posterior margin triangularly produced posteriorly, fitting into the medially emarginate anterior margin of the metaventrite.

110

Ladybird Beetles of the Australo-Pacific Region

Abdomen. Intercoxal process of the first abdominal ventrite with a distinct medial process, short or long and acutely pointed, rarely not prominent. Abdominal postcoxal lines joined in the middle, incomplete laterally, posteriorly very close to the posterior margin of the ventrite; oblique postcoxal line close to, or apparently fused with, the postcoxal line (Fig. 111K). Posterior margin of ventrite 6 weakly arcuate in females and subtruncate in males. Mid and hind tibiae with a pair of apical spurs; tarsal claws appendiculate, with a large basal tooth. Male terminalia. Parameres and phallobase symmetrical; penis guide symmetrical; parameres articulated with the phallobase. Penis stout, consisting of a single sclerite; basal capsule T-shaped. Female terminalia. Coxites shaped like a club handle; styli terminal, well developed, with apical setae; infundibulum lightly sclerotised, not forming a tube; sperm duct indistinct. Spermatheca distinctly curved, with cornu and basal ramus and nodulus; spermathecal accessory gland very large and adjacent to the sperm duct.

Synona melanaria (Mulsant, 1850) (Fig. 111)

Synia melanaria Mulsant, 1850: 375. Type locality: India. Crotch 1874: 178; Korschefsky 1932: 276. Lemnia (Synia) melanaria: Iablokoff-Khnzorian 1982: 232. Synona melanaria: Poorani et al. 2008: 589. Coelophora seminigra Weise, 1902: 501. Type locality: New Guinea. Korschefsky 1932: 296. Synonymised by Poorani et al. 2008: 589. Synia seminigra: Bielawski 1964c: 24. Lemnia (Synia) seminigra: Iablokoff-Khnzorian 1982: 232. Synona seminigra: Pope 1989: 661; Ślipiński 2007: 181. Coccinella cassidoides Montrouzier, 1857: 77. Type locality: New Guinea, Woodlark Island. Crotch 1874:48; Korschefsky 1932: 276. Synonymised by Poorani et al. 2008: 589. Harmonia anthracina Iablokoff-Khnzorian, 1982: 474. Type locality: New Guinea, Wau. Synonymised by Poorani et al. 2008: 589.

Description

Distribution Oriental and Australasian regions (India, Sri Lanka, Bhutan, Burma, Thailand, Vietnam, China, Philippines, Indonesia, Papua New Guinea, Solomon Islands, Australia).

Biology Synona melanaria is known to feed on plataspidid bugs (Hemiptera: Plataspididae) (Schilder and Schilder 1928; Afroze and Uddin 1998; Pope 1989). Monteith (2006) reported S. melanaria (as S. seminigra) preying upon nymphs of Cumare pallida Blote, 1945 (Hemiptera: Tessarotomidae). Ślipiński (2007) has described in detail and keyed the fourth instar larva of S. seminigra.

Remarks Synona is one of the more distinctive genera of Coccinellini, easily distinguished by its compact and convex body, deeply emarginate anterior clypeal margin, the short antenna with the scape expanded externally and the terminal antennomere truncate apically.

Distribution of Synona melanaria

Colour pattern. Frons yellow in males, and black except for the narrowly yellowish inner margins of eyes in females. Head appendages yellow. Pronotum black, except the anterolateral areas are yellow. Specimens from the Solomon Islands (Fig. 111B) have a pronotum that is reddish-testaceous, with a large black subtrapezoidal medial marking on the posterior margin, slightly short of touching the anterior margin in both sexes. Elytra and scutellum entirely black. Ventral side background yellowish, except for the prosternum and pronotal hypomera (except for anterolateral corners) are dark pitchy brown, elytral epipleuron black. Legs yellow. Morphology. Length 5–7 mm. Frons with punctures very shallowly impressed, larger than the eyes facets. Pronotum with finely punctatate, punctures separated by 2–6 diameters, much coarser and deeply impressed on lateral sides, particularly so on the anterolateral corners; interspaces strongly reticulate. Elytra with punctures slightly finer, but more deeply impressed, than those on the pronotum. Mesoventrite with the anterior margin medially deeply emarginate. Epipleura deeply impressed at the level of the middle and hind legs. Postcoxal line meets at the middle and deeply emarginated medially, with an oblique

VII. Review of the Australo-Pacific Coccinellini

lateral line (Fig. 111K). Posterior margin of ventrite 5 almost truncate in both sexes. Male terminalia (Figs 111F–H). Penis with membranous tissues apically; parameres curved, slightly shorter than the penis guide. Female terminalia (Fig. 111I). Spermatheca strongly curved, surface comparably smooth, cornu tapering apically, nodulus and ramus indistinct. Measurements (n = 9): TL 5.40–7.20 (6.34 ± 0.64); PL 0.90–1.10 (1.03 ± 0.07); EL 4.50–6.10 (5.26 ± 0.54); PW 3.20– 4.00 (3.54 ± 0.30); EW 4.70–6.80 (5.76 ± 0.71); GD 2.65–3.70 (3.06 ± 0.42). Ratios: PL/PW 0.28–0.32 (0.29 ± 0.02); EL/EW 0.87–0.96 (0.92 ± 0.03); EL/PL 4.64–5.55 (5.09 ± 0.28); EL/ GD 1.64–1.81 (1.73 ± 0.07); TL/EW 1.06–1.16 (1.10 ± 0.04).

Types Synia melanaria: ‘Ost. Ind., Schaum/ Coll. Haag/ Syntypus/ Synia Mls/ melanaria sp. Mls/ Coll. DEI Müncheberg/ Synia melanaria Muls., female’ (DEI, lectotype). Coccinella cassidoides: ‘Coll. R.I. Sc. N.B. C. cassidoides Synia melanaria, Colln? I. Woodlark/ Coll. R.I. Sc. N.B. Ile Woodlark Entrecasteaux 1847–1852 R.P. Montrouzier / Epilachna cassidoides Montr./ Synia seminigra (Wse) ♀, J. Chazeau det. 1976’ (IRSNB, lectotype). Coelophora seminigra: ‘N. Guinea Fenichel/ Austrolabe Bay, Stephansort/ Holotypus 1902 Coelophora seminigra Weise/ Term. Füz. XXV, 1902, p. 501/ Lemnia seminigra Wse. Khnzorian det.’, (TMB, lectotype). Harmonia anthracina: New Guinea, Wau, 1300 m, 7.III.1968/ J. & M. Sedlacek collectors, Bishop/ K-I. 144/ Holotypus Harmonia anthracina Khnz./ Harmonia anthracina Khn. Khnzorian det. (BPBM, holotype).

Specimens examined INDONESIA: Western New Guinea: Jayawijaya Prov. Samboka, 200m, 10–14.x.1996, A. Riedel (1, NHMS); Finschhafen, 21.iv.1944 (2, CAS); Nabire llaga, road km 54 Pusppenssat, i.1996 P. Hoyois, (6, EVFURZ); Japen Is. E Serui, Wawuti, 100–300m, 17.xii.2000, A. Riedel (1, NKME); Nabire Kali Bobo, 100 m, ix.1990, Balke & Hendrich (NHMW). PAPUA NEW GUINEA: Tapini, 1000–1100m, 18.v.1961, J. L. & M. Gressitt (1, BPBM); SE. Popondetta, 60m, 3–4.ix.1963, J. Sedlacek (1, BPBM); SE Weam, 9m, 20.v-6.vii.1964, H.

111

Clissold (1, BPBM); Wau, 1200m, Malaise trap, 3.xi.1966, J. & M. Sedlacek (1, BPBM); Wau, 1100–1200m,vi.1968, N.L.H. Krauss (1, BPBM); Wau, ii.1974 (1, CAS); NE Wau, Karinga Ck., 950–1150m, 30.ix.1964, J. Sedlacek (1, BPBM); Oriomo R., 3m, 5.viii.1964, H. W. Clissold (1, BPBM); Daru Island, 3m, 25.vii.1964, H. W. Clissold (1, BPBM); Kokoda, 1200 ft, ix.1933, L.E. Cheesman (BMNH); Kerema Govt. Stn, Gulf Distr., 6.v.1959, on Catanus catan, J. Szent-Ivany, A. Catley (BMNH); Magada PTN, Madang Dist., 18.x.1958, on Casava, J. Szent-Ivany (BMNH) Mt Lamington, 1300–1500 feet, C.T. McNamara (SAM). SOLOMON ISLANDS: Guadalcanal, 29 km SE Honiara, 5.vi.1960, Nalimbu R. (1, BPBM); Buka Agric. Sta., 6–10. xii.1959, T. C. Maa (1, BPBM); NW Malaita, Dala, 8.vi.1964, R. Sedlacek (1, BPBM); NW Malaita, Dala, 8.vi.1964, R. Staatman Collector (BPBM). Paratypes: Buka Agric. Sta., 6–10.xii.1959, T.C. Maa Collector (1, BPBM); Guadalcanal, 29 km SE Honiara, Nalimbu R., 5.vi.1960, C.W. O’Brien (1, BPBM); Guadalcanal, Langa River (Mootb.), 26.v.1944, M.F. Milliron (1, BPBM). Guadalcanal, 12–20 or 1–20, J.A. Kusche Coll., Collection of M.W. Giffard (BPBM); Guadalcanal, Kukum, 16.ii.1964, M. McQuillan (BMNH); Guadalcanal, Tenaru, 30.viii.1956, E.S. Brown (BMNH); N end Bougainville, 14.iv.1956, J.L. Gressitt (1, BPBM). AUSTRALIA: Queensland: Murray Isl., Torres Strait, H.L. Clark, 1 female (ANIC); Thursday Is., Torres Strait, 21.iv.1996, J.F. Grimshaw/ JFG 2848A, ex. Centrosema pubescens, 1 male (NAQSM). Auburn River NP via Mundubbera, 17–19.iv.1987, on Petalostigma pubescens, predator of Cumare nymphs Tessarotomidae, GB. Monteith (QDPIB); Lockhart River, Cape York, 8.iv.1992, J.F. Grimshaw/ JFG 1027 (NAQSM); Iron Range NP, 13.v.1974, M. Walford-Huggins (BMNH); Iron Range, 13.v.1971, J.G. Brooks, at light (BMNH); Southedge, via Mareeba, 7–8.iv.1976, R. Storey at light (QDPIM); Hammer Creek, Riverina forest, iv.1993, R. & J. Spencer (ANIC); 13 km W of Ravenshoe, MtGarnet Road, 2.v.1967, D. Colles (ANIC); 8 km SW Kuranda, 30.iii.1984, A. Calder, T. Weir (ANIC); Bowen, Townsville, 14.iii.1965, Exp. De F. Balogh (TMB). Northern Territory: Darwin East Pt. Res. 26.xi.1980, M.B. Malipatil (BMNH, NTM).

Distribution This species is known from the island of New Guinea, Solomon Islands and northern Australia.

BIBLIOGRAPHY Afroze S, Uddin S (1998) Bioecology of Synia melanaria Mulsant (Coleoptera: Coccinellidae) predating on Coptosoma ostensum Distant. Journal of Entomological Research 22, 329–336. Anderson JME (1981) Biology and distribution of Scymnodes lividigaster (Mulsant) and Leptothea galbula (Mulsant), Australian ladybirds (Coleoptera: Coccinellidae). Proceedings of the Linnean Society of New South Wales 105, 1–15. Anderson JME (1982) Seasonal habitat utilization and food of the ladybirds Scymnodes lividigaster (Mulsant) and Leptothea galbula (Mulsant) (Coleoptera: Coccinellidae). Australian Journal of Zoology 30, 59–70. doi:10.1071/ZO9820059 Anderson JME, Hales DF (1983) Micraspis lineata (Thunberg) (Coleoptera: Coccinellidae) – seasonality and food. General and Applied Entomology 15, 47–52. Arrow GJ (1927) Coccinellidae. Clavicornia and Lamellicornia. Insects of Samoa and other Samoan terrestrial Arthropoda Part IV. Coleoptera. Fasc. 1, pp. 1–66. British Museum (Natural History), London. Aruggoda AGB, Shunxiang R, Baoli Q (2010) Molecular phylogeny of ladybird beetles (Coccinellidae: Coleoptera) inferred from mitochondria 16S rDNA sequences. Tropical Agricultural Research 21(2), 209–217. doi:10.4038/tar.v21i2.2602 Balduf WV (1935) The Bionomics of Entomophagous Coleoptera. JS Swift Co. Inc., St Louis, Missouri. Barcos AK, Velasco LI, Medina C (2014) Foraging Behavior of the Yellow Netted Ladybeetle Heteroneda billardieri Crotch (Coleoptera: Coccinellidae). Philippine Agriculturist 97(2), 155–160. Bashford R (1999) Predation by ladybird beetles (Coccinellids) on immature stages of Eucalyptus leaf beetle Chrysophtharta bimaculata (Olivier). Tasforests 11, 77–86. Bielawski R (1957) Coccinellidae (Coleoptera) von Ceylon. Verhandlungen der Naturforschenden Gesellschaft in Basel 68, 72–96. Bielawski R (1961a) Bemerkungen über die männlichen Genitalien von Arten der Gattung Illeis Muls., nebst Beschreibung einer neuen Art und einer Unterart (Coleoptera: Coccinellidae). Annales Zoologici 19(8), 353–368. Bielawski R (1961b) Materialien zur Kenntnis der Coccinellidae (Coleoptera). II. Annales Zoologici 19(10), 383–415. Bielawski R (1961c) Ein neuer Marienkafer, Illeis gressitti sp. n., aus NeuGuinea (Coleoptera, Coccinellidae). Bulletin de l’Académie Polonaise des Sciences. Série des Sciences Biologiques 9, 517–520. Bielawski R (1962) Materialien zur Kenntnis der Coccinellidae (Coleoptera). III. Annales Zoologici 20, 193–205. Bielawski R (1963a) Arten der Gattung Neda Muls. der Australischen Region (Coleoptera, Coccinellidae). Annales Zoologici 20, 457–470. Bielawski R (1963b) Monographie der Epilachninae (Coleoptera, Coccinellidae) der Australischen Region. Annales Zoologici 21, 295–461. Bielawski R (1964a) Die Arten der Gattung Gyrocaria Timberlake (Coleoptera, Coccinellidae). Bulletin de l’Académie Polonaise des Sciences. Série des Sciences Biologiques 12, 83–90. Bielawski R (1964b) A review of the New Guinea species of the genus Harmonia Muls. (Coleoptera: Coccinellidae). Pacific Insects 6, 5–13. Bielawski R (1964c) Die Arten der Gattung Synia Mulsant (Coleoptera, Coccinellidae). Bulletin de l’Académie Polonaise des Sciences. Série des Sciences Biologiques 12, 23–27. Bielawski R (1965) Contribution to the knowledge of Coccinellidae (Coleoptera). IV. Annales Zoologici 23(8), 211–235. Bielawski R (1968) [new species]. In: Bielawski, R. & M. Chûjô Coleoptera from Southeast Asia arranged by M. Chûjô (VII). Coccinellid-beetle from E. Nepal, Thailand, Cambodia, Viet-Nam, S. Korea and Is. Ambon, with descriptions of 4 new species and 33 figures. Transactions of the Shikoku Entomological Society 9, 119–128. Billberg GJ (1808) [new names]. In: Schönherr C. J. Synonyma Insectorum, oder: Versuch einer Synonymie oiler bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum &c. geordnet. Erster B. Eleutherata oder Kafer. Zweiter Theil. Sprechus—Cryptocephalus. CF Marquard, Stockholm.

Blackburn T (1892) Further notes on Australian Coleoptera, with descriptions of new genera and species. XII. Transactions of the Royal Society of South Australia 15(2), 207–261. Blackburn T (1894) Further notes on Australian Coleoptera, with descriptions of new genera and species. XVI. Transactions of the Royal Society of South Australia 18, 59–240. Blackburn T (1895) Further notes on Australian Coleoptera, with descriptions of new genera and species. XVIII. Transactions of the Royal Society of South Australia 19, 201–258. Boisduval JBA (1835) Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, Pendant les Années 1826–1827–1828–1829, sous le Commandement de M. J. Dumont d’Urville. Faune Entomologique de l’Océan Pacifique, avec l’illustration des Insectes Nouveaux Recueillis Pendant le voyage, Deuxième Partie. Coléoptères et autres ordres. J Tastu, Paris. Booth RG, Pope RD (1989) A review of the type material of Coccinellidae (Coleoptera) described by F.W. Pope and by E. Mulsant in the Hope Entomological Collections, Oxford. Entomologica Scandinavica 20(3), 343–370. doi:10.1163/187631289X00366 Brèthes J (1923) Note sur un genre et une espèce de Coccinellides australiens passes inaperçus. Bulletin de la Société Entomologique de France 1923, 227–229. Broun T (1893) Manual of the New Zealand Coleoptera. Parts V, VI, VII. Command, Wellington. Brown PMJ, Thomas CE, Lombaert E, Jeffries DL, Estoup A, Lawson Handley LJ (2011) The global spread of Harmonia axyridis (Coleoptera: Coccinellidae): distribution, dispersal and routes of invasion. BioControl 56, 623–641. doi:10.1007/s10526-011-9379-1 Caltagirone LE, Doutt RL (1989) The history of the Vedalia beetle importation to California and its impact on the development of biological control. Annual Review of Entomology 34, 1–16. doi:10.1146/annurev. en.34.010189.000245 Camacho-Cervantes M, Ortega-Iturriaga A, del-Val E (2017) From effective biocontrol agent to successful invader: the harlequin ladybird (Harmonia axyridis) as an example of good ideas that could go wrong. PeerJ 5, e3296 doi:10.7717/peerj.3296. Casey TL (1899) A revision of the American Coccinellidae. Journal of the New York Entomological Society 7(2), 71–169. Chapin EA (1965) Coleoptera Coccinellidae. Insects of Micronesia 16, 189–254. Chazeau J (1978) Les espèces du genre Coelophora en Nouvelle-Calédonie et aux Nouvelles-Hébrides (Coléoptères, Coccinellidae). Cahiers de l’Office de la Recherche Scientifique et Technique Outre-Mer. Série Biologie 13, 59–69. Chazeau J (1980) On polymorphism in elytral coloration pattern in Coelophora quadrivittata (Coleoptera, Coccinellidae). Entomologia Experimentalis et Applicata 27, 194–198. doi:10.1111/j.1570-7458.1980. tb02964.x Chazeau J (1990) Compléments aus Coccinellini de Nouvelle-Guinée (Coleoptera: Coccinellidae). Revue France d’Entomologie (Nouvelle Serie) 11(1989), 157–168. Chevrolat LAA [new names]. In: Dejean PFMA (1836) Catalogue des Coléoptères de la collection de M. le Comte Dejean. [2 edition, Fascicle 5]. Clausen CP (1940) Entomophagous Insects. New York, McGraw-Hill. Colenso W (1888) A description of a new species of Coccinella found in New Zealand. Transactions and Proceedings of the New Zealand Institute 2(1887), 40–42. Coutanceau J (2008) Le genre Harmonia (Mulsant, 1846) (Coleoptera Coccinellidae). Harmonia 1, 4–16. Crotch GR (1871) List of Coccinellidae. Self-published by the author, Cambridge, UK. Crotch GR (1874) A Revision of the Coleopterous Family Coccinellidae. EW Janson, London. DeBach P (1964) Biological Control of Insect Pests and Weeds. Chapman and Hall, New York. Dejean A, Orivel J, Gibernau M (2002) Specialized predation on plataspid heteropterans in a coccinellid beetle: adaptive behavior and responses

Bibliography

of prey attended or not by ants. Behavioral Ecology 13, 154–159. doi:10.1093/beheco/13.2.154 Dejean PFMA (1836) Catalogue des Coléoptères de la collection de M. le Comte Dejean. [2 edition, Fascicle 5]. Dixon AFG (2000) Insect Predator-Prey Dynamics. Ladybird Beetles and Biological Control. Cambridge University Press, Cambridge. Erichson WF (1842) Beitrag zur Insecten-Fauna von Vandiemensland, mit besonderer Berücksichtigung der geographischen Verbreitung der Insecten. Archiv für Naturgeschichte 8, 83–287. Escalona HE, Zwick A, Li HS, Li JH, Wang XM, Pang H, Hartley D, Jermiin LS, Nedvěd O, Misof B, Niehuis O, Ślipiński A, Tomaszewska W (2017) Molecular phylogeny reveals tremendous food plasticity in the evolution of true ladybird beetles (Coleoptera: Coccinellidae: Coccinellini). BMC Evolutionary Biology 17, 151. doi:10.1186/s12862-0171002-3 Evans EW (2009) Lady beetles as predators of insects other than Hemiptera. Biological Control 51(2), 255–267. doi:10.1016/j.biocontrol.2009.05.011 Fabricius JC (1775) Systema Entomologiae, sistens Insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Officina Libraria Kortii, Flensburgi et Lipsiae. Fabricius JC (1781) Species Insectorum exhibitiens eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Vol. 1 Impensis Carol. Ernest, Bohnii, Hamburgi et Kiloonii. Fabricius JC (1787) Mantissa Insectorum sistens eorum species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Vol. 1 Impensis Christ. Gottl. Proft, Hafniae. Fabricius J Ch (1792) Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species Adjectis Synonimis, Locis, Observationibus, Descriptionibus. Tom I [Pars I]. Impensis Christ. Gottl. Proft., Hafniae. Fabricius JC (1801) Systema Eleutheratorum secundum ordines, genera species adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Kiliae, Impensis Bibliopolii Academici Novi. Fairmaire L (1888) Coléoptères des voyages de M. G. Revoil chez les Somalis et dans l’intérieur du Zanguebar. Annales de la Société entomologique de France (Serie 6) 7, 277–368. Fauvel MA (1867) Catalogue des Coléoptères de la Nouvelle-Calédonie et dépendances avec description, notes et synonymies nouvelles. Bulletin de la Société Linnéenne de Normandie 1, 172–290. doi:10.5962/bhl. part.9927 Fauvel A (1903) Faune analytique des Coléoptères de la Nouvelle Calédonie. Revue d’Entomologie 22, 203–378. Franzmann BA (2002) Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae), a predacious ladybird new in Australia. Australian Journal of Entomology 41, 375–377. doi:10.1046/j.1440-6055.2002.00318.x Froggatt WW (1902) Australian ladybird beetles. Agricultural Gazette of New South Wales 13, 895–911. Fürsch H (1964) Neue Gesichtpunkte zur Beurteilung des Gattungsnamens Micraspis Dejean (Col. Coccinellidae). Nachrichenblatt der Bayerischen Entomologie 13, 70–72. Fürsch H (1989) Discussion about Iablokoff-Khnzorian’s paper of 1986 about Old World Coccinellinae. Coccinella 2, 28–33. Fürsch H (1990) Valid genera and subgenera of Coccinellidae. Coccinella 2, 7–18. Gadeau de Kerville H (1884) Descriptions de quelques espèces nouvelles de la famille des Coccinellidae. Annales de la Société Entomologique de France 4(6e), 69–72. Gage JH (1920) The larvae of Coccinellidae. Illinois Biological Monographs 6, 23–294. Galbreath RA, Cameron PJ (2015) The introduction of the eleven-spotted ladybird Coccinella undecimpunctata L. (Coleoptera: Coccinellidae) to New Zealand in 1874: a spurious record created by cumulative misreporting. New Zealand Entomologist 38, 7–9. doi:10.1080/00779962.2 014.924467 Ganglbauer L (1899) Die Käfer von Mitteleuropa. Die Käfer der österreichisch-ungarischen Monarchie, Deutschlands, der Schweiz, sowie des franzözischen und italienischen Alpengebietes. Familienreihe Clavicornia. Sphaeritidae, Ostomidae, Byturidae, Nitidulidae, Cucujidae, Erotylidae, Phalacridae, Thorictidae, Lathridiidae, Mycetophagidae,

113

Colydiidae, Endomychidae, Coccinellidae. Volume III. C Gerald’s Sohn, Wien. Giorgi JA, Vandenberg NJ, McHugh JV, Forrester J, Ślipiński A, Miller KB, Shapiro LR, Whiting MF (2009) The evolution of food preferences in Coccinellidae. Biological Control 51, 215–231. doi:10.1016/j.biocontrol.2009.05.019 Gmelin JF (1790) Caroli a Linné Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima tertia, aucta, reformata. Tomus I Pars IV. Classis V. Insecta. Georg Enanuel Beer, Lipsiae. Goeze JAE (1777) Entomologische Beytraege zu des Ritters Linné zwölften Ausgabe des Natursystems. Erster Theil. Weidmanns Erben und Reich, Leipzig. Gordon RD (1985) The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Entomological Society 93, 1–912. Gordon RD (1987) A catalogue of the Crotch collection of Coccinellidae (Coleoptera). Occasional Papers on Systematic Entomology 3, 1–46. Guérin-Méneville FE (1835) Iconographie du Régne Animal de G. Cuvier, ou représentations d’apres de l’une espèces les plus remarquables, et souvent non encore figurées, de chaque genre d’animaux. Avec un texte descriptif mis au courant de la science. Ouvrage pouvant servir d’atlas a tous les traités de zoologie. Vol. 2, Insectes. J.B. Balliere, Libraire de l’Académie Royale de Médicine, Paris. Gyllenhal. In: Schönherr CJ (1808) Synonyma Insectorum, oder: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum &c. geordnet. Erster Band. Eleutherata oder Käfer. Zweiter Theil. CF Marquard, Stockholm. Hagen KB (1962) Biology and ecology of predaceous Coccinellidae. Annual Review of Entomology 7, 289–326. doi:10.1146/annurev. en.07.010162.001445 Hales DF (1976) Inheritance of stripe elytral patterm in Coelophora inaequalis (F.) (Coleoptera: Coccinellidae). Australian Journal of Zoology 24, 273–276. doi:10.1071/ZO9760273 Hales DF (1977) Coleophora veranioides Blackburn: a variety of Coleophora inaequalis (F.) (Coleoptera: Coccinellidae). Australian Entomological Magazine 4, 55–56. Hawkeswood TJ, Turner JR (2002) Observations on the adults of Micraspis frenata (Erichson, 1842) (Coleoptera: Coccinellidae) feeding on the pollen of native and non-native grasses (Poaceae) in eastern New South Wales, Australia. Journal of Entomological Research Society 4, 21–29. Hinton HE (1946) The “gin-traps” of some beetle pupae; a protective device which appears to be unknown. Transactions of the Royal Entomological Society of London 97, 473–496. doi:10.1111/j.1365-2311.1946. tb00273.x Hoang DN (1982) Bo Rua Coccinellidae o Viet Nam (Insecta, Coleoptera) Tap I. Nha Xuat Ban Khoa Hoc Va Ky Thuat, Hanoi (in Vietnamese). Hoang DN (1983) Bo Rua Coccinellidae o Viet Nam (Insecta, Coleoptera) TapII. Nha Xuat Ban Khoa Hoc Va Ky Thuat, Hanoi. (in Vietnamese). Hodek I (1967) Bionomics and ecology of predaceous Coccinellidae. Annual Review of Entomology 12, 79–104. doi:10.1146/annurev. en.12.010167.000455 Hodek I (1973) Biology of Coccinellidae. Academia Publishing and W. Junk, Prague and The Hague. Hodek I (1996a) Food relationships. In Ecology of Coccinellidae. (Eds I Hodek and A. Honek) pp. 143–237 Kluwer Academic Publishers, Dordrecht Boston London. Hodek I (1996b) Dormancy. In Ecology of Coccinellidae (Eds I Hodek and A Honek) pp. 239–318. Kluwer Academic Publishers, Dordrecht Boston London. Hodek I, Honěk A (1996) Ecology of Coccinellidae. Series Entomologica, 54. Kluwer Academic Publishers, Dordrecht. Hodek I, Michaud JP (2008) Why is Coccinella septempunctata so successful? (A point-of-view). European Journal of Entomology 105, 1–12 doi:10.14411/eje.2008.001. Hodek I, van Emden HF, Honěk A (2012) Ecology and behaviour of the ladybird beetles (Coccinellidae). Wiley-Blackwell, Chichester. Hope FW (1840) The Coleopterist’s manual, part the third, contains various families, genera, and species, recorded by Linnaeus and Fabricius. Also descriptions of newly discovered and unpublished insects. JC Bridgewater, London.

114

Ladybird Beetles of the Australo-Pacific Region

Houston KJ (1979) Mosaic dominance in the inheritance of the colour patterns of Coelophora inaequalis (F.) (Coleoptera: Coccinellidae). Journal of the Australian Entomological Society 18, 45–51. doi:10.1111/j.1440-6055.1979.tb00810.x Houston KJ (1983) Inheritance of colour pattern in Phrynocaria gratiosa (Mulsant) (Coleoptera: Coccinellidae). General and Applied Entomology 15, 19–30. Houston KJ (1988) Larvae of Coleophora inaequalis (F.), Phrynocaria gratiosa (Mulsant) and P. astrolabiana (Weise) (Coleoptera: Coccinellidae) with notes on their relationships and prey records. Journal of the Australian Entomological Society 27, 199–211. doi:10.1111/j.1440-6055.1988.tb01525.x Houston KJ, Hales DF (1980) Allelic frequencies and inheritance of colour pattern in Coelophora inaequalis (F.) (Coleoptera: Coccinellidae). Australian Journal of Zoology 28, 669–677. doi:10.1071/ZO9800669 Iablokoff-Khnzorian SM (1979) Genera der paläarktischen Coccinellini (Coleoptera, Coccinellidae). Entomologische Blätter 75(1–2), 37–75. Iablokoff-Khnzorian SM (1982) Les Coccinelles. Coleoptères-Coccinellidae. Société nouvelle des editions Boubée, Paris. Iablokoff-Khnzorian SM (1984a) Notes sur la tribu des Coccinellini (Coleoptera, Coccinellidae). Nouvelle Revue d’Entomologie (NS) 1(2), 203–222. Iablokoff-Khnzorian SM (1984b) Synopsis von zwei Marienkäfergattungen aus der Australischen Region (Coleoptera, Coccinellidae). Entomologische Blätter 80(2/3), 107–122. Iablokoff-Khnzorian SM [as Jablokov-Khnzorjan] (1986) Bemerkungen über einige Coccinellini der Alten Welt (Coleoptera, Coccinellidae). Entomologische Abhandlungen Staatliches Museum für Tierkunde Dresden 50(3), 59–80. Iperti G (1999) Biodiversity of predaceous Coccinellidae in relation to bioindication and economic importance. Agriculture, Ecosystems & Environment 74, 323–342. doi:10.1016/S0167-8809(99)00041-9 Jadwiszczak A (1990) On the systematic position of the genus Subepilachna Bielawski (Coeloptera, Coccinellidae). Coccinella 2(2), 56–57. Kamiya H (1965a) Comparative morphology of larvae of the Japanese Coccinellidae, with special reference to the tribal phylogeny of the family (Coleoptera). The Memoirs of the Faculty of Education, Fukui University Series II. Nature and Science 14, 83–100. Kamiya H (1965b) A revision of the tribe Coccinellini of Japan and the Ryukyus (Coleoptera: Coccinellidae). Memoirs. Faculty of Liberal Arts. Fukui University. (2, Nat. Sci.) 15(3), 27–71. Kapur AP (1963) The Coccinellidae of the third Mount Everest Expedition, 1924 (Coleoptera). Bulletin of the British Museum (Natural History). Entomology 14(1), 1–48. Kirby W, Spence W (1822) An Introduction to Entomology or Elements of the Natural History of Insects. Vol. 1. (4th ed.). Longman, London. Kitano T (2014) Taxonomic notes on the genus Egleis Mulsant, 1850 and description of a new subgenus in the genus Illeis Mulsant, 1850 (Coleoptera: Coccinellidae). Studies and Reports. Taxonomical Series 10(2), 489–494. Korschefsky R (1931) Pars 118: Coccinellidae. I. In Coleopterorum Catalogus. (Eds W. Junk and S Schenkling) pp. 1–224. W. Junk, Berlin. Korschefsky R (1932) Pars 120: Coccinellidae. II. In Coleopterorum Catalogus. (Eds W Junk and S Schenkling) pp. 225–659. W. Junk, Berlin. Korschefsky R (1934a) Prof. Dr. E. Handschin, Studienreise auf den Sundainseln und Nordaustralien, 1930–32. 3. Drei neue Coccinelliden der Indo-malayischen und Papuanischen Region. Mitteilungen der Schweizerische Entomologische Gesellschaft 16(2), 107–109. Korschefsky R (1934b) Bemerkungen über Coccinelliden und Beschreibung einer neuen Neda-Art. Arbeiten über Morphologische und Taxonomische Entomologie aus Berlin-Dahlem 1(4), 267–270. Korschefsky R (1940) Vier neue Coccinelliden der Alten Welt. Arbeiten über Morphologische und Entomologische Blätter 36, 1–3. Korschefsky R (1943) Ueber Coccinelliden von den Neuen Hebriden aus dem National-Museum, Paris. Entomologische Blätter 39, 51–52. Korschefsky R (1944) Neue altweltliche Coccinelliden (Coleoptera: Coccinellidae). Arbeiten über Morphologische und Taxonomische Entomologie aus Berlin-Dahlem 11(1), 47–56. Kovář I (1996a) Morphology and anatomy. In Ecology of Coccinellidae. (Eds I Hodek and A Honek) pp. 1–18. Kluwer Academic Publishers, Dordrecht.

Kovář I (1996b) Phylogeny. In Ecology of Coccinellidae. (Eds I Hodek and A Honek) pp. 19–31. Kluwer Academic Publishers, Dordrecht. Kovář I (2007) Family Coccinellidae Latreille, 1807. In: Catalogue of Palaearctic Coleoptera, Volume 4. (Eds I Löbl and A Smetana) pp. 568–631. Apollo Books, Stenstrup. Kugelann JG (1794) Verzeichniss der in einigen Gegenden Preussens bis jetzt entdeckten Käfer-Arten, nebst kurzen Nachrichten von denselben. Neuestes Magazin für die Liebhaber der Entomologie 1(5), 513–582. Kuznetsov VN (1997) Lady beetles of the Russian Far East. Center for Systematic Entomology Memoir 1, 1–248. Latreille PA (1807) Genera Crustaceorum et Insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata. Tomus Tertius. A Koenig, Parisis et Argentorati. Latreille PA (1810) Considérations générales sur l’ordre naturel des animaux composant les classes des Crustacés, des Arachnides, et des Insectes; avec un tableau méthodique de leurs genres, disposés en familles. F Schoell, Paris. Latreille PA (1825) Familles Naturelles du Règne Animal, Exposées Succinctement et dans un Ordre Analytique, avec l’Indication de leurs Genres. JB Baillière, Paris. Lawrence JF, Ślipiński SA, Seago AE, Thayer MK, Newton AF, Marvaldi AE (2011) Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici 61, 1–217. doi:10.3161/000345411X576725 Lea AM (1902) Descriptions of new species of Australian Coleoptera. Proceedings of the Linnean Society of New South Wales 26 (1901), 481–513. Li JH, Pang H, Slipinski A (2009) Revision of the Genus Australoneda Iablokoff-Khnzorian, 1984 (Coleoptera: Coccinellidae: Coccinellini). Annales Zoologici 59(2), 223–235. doi:10.3161/000345409X464047 Li J, Tomaszewska W, Pang H, Ślipiński A (2014) Ladies in stripes: taxonomic confusion in a potential mimicry complex among Wallacean Coccinellidae (Coleoptera: Coccinellidae). Zootaxa 3900(4), 592–600. doi:10.11646/zootaxa.3900.4.9 Linnaeus C (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species cum characteribus, differentiis, synonymis, locis. Vol. 1 10 (reformata). Laurentii Salvii, Holmiae. Lundgren JG (2009) Nutritional aspects of non-prey foods and the life histories of predaceous Coccinellidae. Biological Control 51, 294–305. doi:10.1016/j.biocontrol.2009.05.016 Lundgren JG, Wiedenmann RN (2004) Nutritional suitability of corn pollen for the predator, Coleomegilla maculata (Coleoptera: Coccinellidae). Journal of Insect Physiology 50, 567–575. doi:10.1016/j.jinsphys.2004.04.003 Lundgren JG, Huber A, Wiedenmann RN (2005) Quantification of the consumption of corn pollen by the predator Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae) in an Illinois cornfield. Agricultural and Forest Entomology 7, 53–60. doi:10.1111/j.1461-9555.2005.00246.x Lydekker R (1896) A geographical history of mammals. Cambridge University Press, Cambridge. Macleay WS. In: King PP (ed.) (1826) Narrative of a survey of the intertropical and western coasts of Australia: performed between the years 1818 and 1822. Vol. 2. Murray, London. Mader L (1954) Coccinellidae III Teil. Exploration du Parc National Albert, Mission G.F. de Witte (1933–1935) 80, 1–106. Magro A, Lecompte E, Magne F, Hemptinne J-L, Crouau-Roy B (2010) Phylogeny of ladybirds (Coleoptera: Coccinellidae): are the subfamilies monophyletic. Molecular Phylogenetics and Evolution 54, 833–848. doi:10.1016/j.ympev.2009.10.022 Majerus MEN (1994) Ladybirds. The New Naturalist Library. Harper Collins Publishers, London. Majerus MEN (2016) A Natural History of Ladybird Beetles. Cambridge University Press, Cambridge. Martin NA (2016a) Two-spotted ladybird – Adalia bipunctata. Interesting insects and other invertebrates. New Zealand Arthropod Factsheet Series Number 37. . Martin NA (2016b) Southern ladybird – Cleobora mellyi. Interesting insects and other invertebrates. New Zealand Arthropod Factsheet Series Number 57. .

Bibliography

Martin NA (2016c) Large spotted ladybird – Harmonia conformis. Interesting insects and other invertebrates. New Zealand Arthropod Factsheet Series Number 39. . Martin NA (2017) Harlequin ladybird – Harmonia axyridis. Interesting insects and other invertebrates. New Zealand Arthropod Factsheet Series Number 51. . Masters G (1888) Catalogue of the described Coleoptera of Australia. Part VII. Proceedings of the Linnean Society of New South Wales 2(2), 1–82. Michaud JP (2002) Biological control of Asian citrus psyllid, Diaphorina citri (Hemiptera: Psyllidae), in Florida: a preliminary report. Entomological News 113, 216–222. Miyatake M (1965) Some Coccinellidae (Excluding Scymnini) of Formosa (Coleoptera). Special Bulletin of the Lepidopterists’ Society of Japan 1, 50–74. Miyatake M (1972) A new Formosan species belonging to the genus Singhikalia Kapur, with proposal of a new tribe (Coleoptera: Coccinellidae). Transactions of the Shikoku Entomological Society 11(3), 92–98. Monteith GB (2006) Maternal care in Australian oncomerine shield bugs (Insecta, Heteroptera, Tessaratomidae). Denisia 19, 1135–1152. Montrouzier X (1857) Essai sur la Faune l’île de Woodlark ou Moiou. Annales des Sciences Physiques et Naturalles d’Agriculture et d’Industrie (Serie 2) 7, 1–114. Montrouzier X (1861) Essai sur la faune entomologique de la NouvelleCalédonie (Balade) Coléoptères (Fin). Annales de la Société Entomologique de France (Serie 4) 1, 265–306. Mulsant E (1846) Histoire Naturelle des Coléoptères de France. Sulcicolles-Sécuripalpes. Maison, Paris. Mulsant E (1850) Species des Coléoptères Trimères Sécuripalpes. Annales des Sciencies Physiques et Naturelles, d’Agriculture et d’Industrie, publiées par la Société nationale d’Agriculture, etc., de Lyon, Deuxième Série. Mulsant E (1853) Supplément a la Monographie des Coléoptères Trimerès Sécuripapes. Annales de la Société Linnéenne de Lyon (Nouvelle Série) (2)2(1852–1853), 129–333. Mulsant E (1866) Monographie des Coccinellides. Savy et Deyrolle, Paris. Nakane T, Araki M (1959) The Coleoptera of Yakushima Islands. Coccinellidae. Scientific Report Saikyo University 2A, 291–293[In Japanese with English summary]. Nattier R, Jourdan H, Mille C, Chazeau J (2015) An annotated checklist of the Coccinellidae (Coleoptera) from New Caledonia. Zootaxa 4058(3), 301–331. doi:10.11646/zootaxa.4058.3.1 Nedvěd O (2014) Slunéčko východní (Harmonia axyridis) – pomocník v biologické ochraně nebo ohrožení biodiverzity? Jihočeská univerzita, České Budějovice. Oliff S (1895) Entomological notes – a new friendly ladybird. Agricultural Gazette of New South Wales 6, 30–31. Pallas PS (1773) Reise durch verschiedene Provinzen des Rußischen Reichs. Zweyter Theil, zweytes Buch vom Jahr 1771. Kayserliche Academie der Wißenschaften, St. Petersburg. Pang X-F, Mao J-L (1979) Economic Insect Fauna of China (Coleoptera: Coccinellidae) – II. Science Press, Beijing (in Chinese). Phuoc DT, Stehr FW (1974) Morphology and taxonomy of the known pupae of Coccinellidae (Coleoptera) of North America, with discussion of phylogenetic relationships. Contributions of the American Entomological Institute 10, 1–125. Poorani J (2002a) An annotated checklist of the Coccinellidae (Coleoptera) (excluding Epilachninae) of the Indian subregion. Oriental Insects 36, 307–383. doi:10.1080/00305316.2002.10417335 Poorani J (2002b) A review of the genus Oenopia Mulsant (Coleoptera: Coccinellidae) of the Indian subcontinent, with description of a new species. Oriental Insects 36, 97–116. doi:10.1080/00305316.2002.104 17328 Poorani J, Booth RG (2005) On the identity of Dysis excellens Crotch (Coleoptera: Coccinellidae: Coccinellini), a little known Oriental lady beetle. Zootaxa 875, 1–5. doi:10.11646/zootaxa.875.1.1 Poorani J, Booth RG (2016) Harmonia manillana (Mulsant), a new addition to Indian Coccinellidae, with changes in synonymy. Biodiversity Data Journal 4, e8030 doi:10.3897/BDJ.4.e8030.

115

Poorani J, Ślipiński A, Booth RG (2008) A revision of the genus Synona Pope, 1989 (Coleoptera: Coccinellinae: Coccinellini). Annales Zoologici 58(3), 579–594. doi:10.3161/000345408X364427 Pope RD (1989) A revision of the Australian Coccinellidae (Coleoptera). Part 1. Subfamily Coccinellinae. Invertebrate Taxonomy 2, 633–735. doi:10.1071/IT9880633 Rees BE, Anderson DM, Bouk D, Gordon RD (1994) Larval key to genera and selected species of North American Coccinellidae (Coleoptera). Proceedings of the Entomological Society of Washington 96, 387–412. Ren S, Wang X, Pang H, Peng Z, Tao Z (2009) Colored Pictorial Handbook of Ladybird Beetles in China. Science Press, Beijing. Richards AM (1980) Sexual selection, guarding sexual conflict in a species of Coccinellidae (Coleoptera). Journal of the Australian Entomological Society 19, 26. doi:10.1111/j.1440-6055.1980.tb00956.x Roy HE, Brown PMJ, Adriaens T, Berkvens N, Borges I, Clusella–Trullas S, Comont RF, De Clercq P, Eschen R, Estoup A, Evans EW, Facon B, Gardiner MM, Gil A, Grez AA, Guillemaud T, Haelewaters D, Herz A, Honěk A, Howe AG, Hui C, Hutchison WD, Kenis M, Koch RL, Kulfan J, Lawson Handley L, Lombaert E, Loomans A, Losey J, Lukashuk AO, Maes D, Magro A, Murray KM, San Martin G, Martinkova Z, Minnaar IA, Nedved O, Orlova–Bienkowskaja MJ, Osawa N, Rabitsch W, Ravn HP, Rondoni G, Rorke SL, Ryndevich SK, Saethre M, Sloggett JJ, Soares AO, Stals R, Tinsley MC, Vandereycken A, van Wielink P, Viglasova S, Zach P, Zakharov IA, Zaviezo T, Zhao Z (2016) The harlequin ladybird, Harmonia axyridis: global perspectives on invasion history and ecology. Biological Invasions 18, 997–1044. doi:10.1007/ s10530-016-1077-6 Rye EC (1873) Zoological Record 1871. Vol. 8. John van Voorst, London. Sasaji H (1968) Descriptions of the Coccinellid larvae of Japan and Ryukyus (Coleoptera). The Memoirs of the Faculty of Education, Fukui University Series II. Nature and Science 18, 93–135. Sasaji H (1971) Coccinellidae (Insecta: Coleoptera). Fauna Japonica. Academic Press of Japan, Tokyo. Sasaji H (1981) Biosystematics on Harmonia axyridis-complex (Coleoptera: Coccinellidae). The Memoirs of the Faculty of Education, Fukui University Series II. Nature and Science 30(3), 59–79. Sasaji H, Akamatsu M (1979) Reproductive continuity and genetic relationships in the forms of the genus Menochilus (Coleoptera: Coccinellidae). The Memoirs of the Faculty of Education, Fukui University Series II. Nature and Science 29(1), 1–18. Savoiskaya GI (1969) Novyy rod i vid kokcinellid iz Kazakhstana. Vestnik Sel’skokhazyaystvenoy Nauki (Alma-Ata) 6, 37–43. Savoiskaya GI (1983) Larvae of ladybird beetles (Coleoptera, Coccinellidae) of Soviet Fauna. Opredelileli Po Faunie SSSR, 137. Nauka, Leningrad. Say T (1824) Descriptions of coleopterous insects collected in the late expedition to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary of War, under the command of Major Long. Journal of the Academy of Natural Sciences of Philadelphia 4, 83–99. Schilder FA, Schilder M (1928) Die Nachrung der Coccinelliden und ihre Beziehung zur Vervandtschaft der Arten. Arbeiten aus der Biologischen Reichsanstalt für Land- und Forstwirtschaft 16, 213–282. Schönherr CJ (1808) Synonyma Insectorum, oder: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum &c. geordnet. Erster Band. Eleutherata oder Käfer. Zweiter Theil. CF Marquard, Stockholm. Scriba LG (1791) Beitrage zu der Insecten-Geschichte. Zweites Heft. Varrentrapp & Wenner, Frankfurt. Seago AE, Giorgi JA, Li J, Ślipiński A (2011) Phylogeny, classification and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution 60(1), 137–151. Shepard BM, Rapusas HR (1989) Life cycle of Micraspis sp. on brown planthopper (BPH) and rice pollen. International Rice Research Newsletter 14, 40. Sicard A (1907) Notes sur quelques Coccinellides Paléarctiques (Col.). Bulletin de la Société Entomologique de France 1907, 132–136. doi:10.5962/bhl.part.4570 Sicard A (1929a) Description d’espèces nouvelles de Coccinellidae. Annals & Magazine of Natural History (Series 10) 4, 515–524. doi:10.1080/00222932908673088

116

Ladybird Beetles of the Australo-Pacific Region

Sicard A (1929b) Descriptions d’espèces varietes nouvelles de Coccinellides appartenant au genre Caria Mls. Annales de la Société Entomologique de France 48, 188–192. Ślipiński A (2007) Australian ladybird beetles (Coleoptera: Coccinellidae): their biology and classification. Australian Biological Resources Study, Canberra. Sutherland AM, Parrella MP (2009) Mycophagy in Coccinellidae: Review and synthesis. Biological Control 51, 284–293. doi:10.1016/j.biocontrol.2009.05.012 Thunberg CP (1781) Dissertatio Entomologica Novas Insectorum Species, sistens cujus partem primam, Cons. Exper. Facul. Med. Upsal., publice ventilandam exhibent praeses Carol. P. Thunberg, et respondens Samuel Nicol. Casström. Joh. Edman, Upsaliae. Timberlake PH (1943) The Coccinellidae or ladybeetles of the Koebele collection – part I. The Hawaiian Planters’. Record 47(1), 7–67. Vandenberg NJ (1992) Revision of the New World Lady Beetles of the genus Olla and description of a new allied genus (Coleoptera: Coccinellidae). Annals of the Entomological Society of America 85(4), 370–392. doi:10.1093/aesa/85.4.370 Vandenberg NJ (1996) Notes on Oenopia Mulsant (Coleoptera: Coccinellidae). Proceedings of the Entomological Society of Washington 98, 385–389. Vandenberg NJ (2002) Family 93. Coccinellidae Latreille 1807. In: American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. (Eds RH Arnett Jr, MC Thomas, PE Skelley and JH Frank) CRC Press LLC, Boca Raton, Florida. Watt JC (1982) Terrestrial arthropods from the Poor Knights Islands, New Zealand. Journal of the Royal Society of New Zealand 12, 283–320. do i:10.1080/03036758.1982.10415350 Watt JC (1986) Beetles (Coleoptera) of the offshore islands of northern New Zealand. In: The Offshore Islands of Northern New Zealand. Information series, (16). (Eds AE Wright and RE Beever) pp. 221–228. Department of Lands and Survey, Wellington, New Zealand. Weise J (1892a) Neue Coccinelliden. Deutsche Entomologische Zeitschrift 1891(2), 282–288. Weise J (1892b) Kleine Beiträge zur Coccinelliden-Fauna Ost-Afrika’s. Deutsche Entomologische Zeitschrift 1892(1), 15–16.

Weise J (1895) Neue Coccinelliden, sowie bemerkungen zu bekannten Arten. Annales de la Société Entomologique de Belgique 39, 120–146. Weise J (1898) Ueber bekannte und neue Coccinelliden. Archiv fuer Naturgeschichte 64/1(2), 225–238. Weise J (1902) Coccinelliden aus der Sammlung des Ungarischen National-Museums. Természetrajzi Füzetek 25, 489–520. Weise J (1903) Neue Coccinelliden. Deutsche Entomologische Zeitschrift 1903(2), 229–232. Weise J (1908) Die Fauna Südwest-Australiens. Ergebnisse der Hamburger südwest-australischen Forschungsreise 1905. Herausgegeben von Prof. Dr. W. Michaelsen und Dr. R. Hartmeyer. Chrysomelidae und Coccinellidae. Vol. II(I). G Fisher, Jena. Weise J (1910) Verzeichnis von Coleopteren aus den Philippinen, nebst zwei neuen Arten aus Niederländisch Ostindien. Philippine Journal of Science 5, 139–148. Weise J (1912) Über Hispinen und Coccinelliden. Archiv für Naturgeschichte A 78(2), 100–120. Weise J (1923) Results of Dr. E. Mjöberg’s Swedish Scientific Expedition to Australia 1910–1913. 31. Chrysomeliden und Coccinelliden aus Queensland. Arkiv för Zoologi 15(12), 1–150. Weise J (1927) Uber bekannte und neue Chrysomeliden und Coccinelliden aus dem Reichsmuseum zu Stockholm. Arkiv för Zoologi 18A(34), 23–34. White A (1846) Insects of New Zealand. In: The zoology of the voyage of H.M.S. Erebus & Terror, under the command of Captain Sir James Clark Ross, R.N., F.R.S., during the years 1839 to 1843. Vol. II. Reptiles, fishes, Crustacea, insects, Mollusca. (Eds J Richardson and JE Gray) Pp. 1–27, plates 1–7. EW Janson, London. White A (1848) Description of a new species of Coccinella from New Zealand by M. Mulsant of Lyon, author of the Histoire Naturelle des Coléoptères de France. Annals and Magazine of Natural History. Second Series 1, 66–67. Yu G (1994) Cladistic analyses of the Coccinellidae (Coleopera). Entomologia Sinica 1(1), 1–30. Yu G (2010) Chinese Lady Beetles (The Subfamily Coccinellinae). Chemical Industry Press, Beijing.

Colour plates

A

D

B

E

C

F

117

Fig. 1.  Coccinellini in their natural habitats: A–B, Australoneda ruitong Li et al.; C, Australoneda karubakana (Bielawski); D, Australoneda bourgeoisi (Kerville); E, Docimocaria princeps (Mulsant); F, Microcaria mulsanti (Montrouzier). D © Nick Monaghan, remaining © Paul Zborowski.

118

Ladybird Beetles of the Australo-Pacific Region

A

D

B

E

C

F

Fig. 2.  Coccinellini in their natural habitats: A–B, Micraspis frenata (Erichson); C, Coelophora inaequalis (Fabricius); D, Illeis galbula (Mulsant); E, Harmonia conformis (Boisduval); F, Hippodamia variegata (Goeze). A–C © Paul Zborowski; D © Steve Axford; E © Andrew Bonnitcha; F © Nick Monaghan.

Colour plates

A

D

B

E

C

F

119

Fig. 3.  Coccinellini in their natural habitats: A–B, Cleobora mellyi (Mulsant); C, Harmonia conformis (Boisduval); D, Coccinella leonina Fabricius; E, Coelophora inaequalis (Fabricius); F, Micraspis flavovittata (Crotch). A © Jiri Lochman; B, C, E © Nick Monaghan; D © Jon Sullivan; F © Reiner Richter.

120

A

Ladybird Beetles of the Australo-Pacific Region

C

D B

E

Fig. 4.  Coccinellini in their natural habitats: A, winter gathering of Illeis galbula (Mulsant); B, overwintering Harmonia axyridis (Pallas) near Auckland; C–D, Harmonia axyridis (Pallas); E, Harmonia conformis (Boisduval) overwintering within open fruit of Brachychiton populneus (Schott & Endl.) R.Br. A © Nick Monaghan; B © Caleb Hutchinson-Brown and Samantha Brown; C–D © Yun Li; E © David McClenaghan, CSIRO.

Colour plates

A

121

D

E B

F

C G

Fig. 5.  Coccinellini in their natural habitats: A, Coccinella transversalis Fabricius, adult; B, same species, adult cannibalising pupa; C, Coelophora inaequalis (Fabricius); D, Harmonia octomaculata (Fabricius); E, Assasin bug (Reduviidae) feeding on ladybird; F, Harmonia testudinaria (Mulsant); G, Coccinella transversalis Fabricius, larva ready to pupate. A, B, G © Marie Lochman; C–D © David McClenaghan, CSIRO; E © Jiri Lochman; F © Nick Monaghan.

122

Ladybird Beetles of the Australo-Pacific Region

A

D

B

E

C

F

Fig. 6.  Coccinellini in their natural habitats: A, Cleobora mellyi (Mulsant), larva feeding on larvae of chrysomelid beetle; B, Coccinella undecimpunctata Linnaeus; C, Cheilomenes sexmaculata (Fabricius); D, Harmonia conformis (Boisduval), larva; E, Illeis galbula (Mulsant), larva; F, Subepilachna latemarginata Bielawski. A © Andrew Bonnitcha; B © Kristi Ellingsen; C © Jiri Lochman; D, E © Nick Monaghan; F © Paul Zborowski.

Colour plates

A

B

Ey

PB

123

Fr Cl LB

Oc

EB Sca Pd

Mx

C

ST Mnt CA Smt Gu

Pl

D Head Prothorax Pterothorax

Abdomen

Fig. 7.  Adult morphology: A, frontal view; B, ventral view; C, dorsal view; D, lateral view (from Ślipiński 2007). Abbreviations: A1–A5 – abdominal ventrites; An – antenna; APl – abdominal postcoxal lines; CA – cardo; Cl – clypeus; CX1–CX3 – coxal cavities/coxae; Ds – median discrimen; EB – elytral bead; EF – epipleural fovea; EL – elytron; EP – elytral epipleuron; Em2 – mesepimeron; Em3 – metepimeron; Es2 – mesanepisternum; Es3 – metanepisternum; Ey – compound eye; Fr – frons; Gu – gula; HP – prothoracic hypomeron; L1–L3 – legs; LB – labrum; Mnt – mentum; MPl – metaventral postcoxal line; MS – mesoventrite; MT – metaventrite; Mx – terminal maxillary palpomere; NPs – notopleural suture; Oc – ocular canthus; PB – pronotal bead; Pd – antennal pedicel; Pl – prosternal lines; PN – pronotum; PP – prosternal process; PR – prosternum; Sc – scutellum; Sca – antennal scape; Smt – submentum; ST – stipes. Design by A. Hastings

124

Ladybird Beetles of the Australo-Pacific Region

A

C

B

D

E

F

Fig. 8.  Adult morphology: A, female terminalia and genital track; B, spermatheca and apical part of bursa copulatrix; C, male terminal abdominal segments, dorsal view; D, penis, lateral view; E, tegmen, lateral; F, tegmen, ventral (from Ślipiński 2007). Abbreviations: BC – bursa copulatrix; BP – phallobase; c – cornu; CG – collateral glands; CO – common oviduct; Cx – coxite (genital plate); in – infundibulum; n – nodulus; Pc – penis capsule; PG – penis guide; PM – parameres; Pp – paraprocts (T9); r – ramus; Sd – sperm duct; Sg – spermathecal gland; SP – spermatheca; S8–9 – sternites 8–9; TS – tegminal strut (trabes); T8–10 – tergites 7–10; VG – vagina.

Colour plates

A

B

C

G

D

E

I

125

F

J

H

K

Fig. 9.  Larva and pupa morphology: A, Archegleis edwardsi (Weise), mature larva, dorsal; B, Micraspis frenata (Erichson), antenna; C, Harmonia conformis (Boisduval), early instar larva, lateral; D, Harmonia conformis (Boisduval), thorax, lateral; E, Docimocaria princeps (Mulsant), claw; F, Harmonia octomaculata, ventral mouth parts; G, H, Illeis galbula Mulsant, mandible; I, Docimocaria princeps (Mulsant), pupa ventral; J, Illeis galbula Mulsant, pupa dorsal; K, Archegleis kingi (MacLeay), pupa lateral (GT – gin traps).

126

Ladybird Beetles of the Australo-Pacific Region

Fig. 10.  Map of the study area.

Colour plates

127

Fig. 11.  Key generic characters: A, Subepilachna, body lateral; B, Coccinella, body lateral; C, Adalia, body dorsal; D, Hysia, body dorsal; E, Illeis, maxillary palp; F, Adalia, maxillary palp; G, Docimocaria, antenna; H, Harmonia, antenna; I, Adalia, recurved abdominal postcoxal line; J, Microcaria, not recurved abdominal postcoxal line; K, Harmonia, abdominal postcoxal line (PL) and associated oblique postcoxal line (OPL); L, Synona, abdominal ventrites; M, Illeis, mandibular apex; N, Hippodamia, tibial spurs; O, Synona, pretarsal claws; P, Hippodamia, pretarsal claws; Q, Illeis, head and pronotum.

128

Ladybird Beetles of the Australo-Pacific Region

Fig. 12.  Key generic characters: A, Adalia, prothorax, ventral; B, Coelophora, same; C, Archegleis, same; D, Synona, abdominal ventrites; E, Microcaria, apex of elytral epipleuron; F, Megalocaria, same; G, Archegleis, spermatheca; H, Microcaria, spermatheca; I, Docimocaria, antenna; J, Australoneda, antenna; K, Megalocaria, penis; L, Docimocaria, penis; M, Archegleis, mesoventrite; N, Docimocaria, mesoventrite; O, Microcaria, head; P, Phrynocaria, head; Q, Heterocaria, abdominal postcoxal line; R, Archegleis, same.

Colour plates

129

Fig. 13.  Key generic characters: A, Synona, head; B, Micraspis, same; C, Cheilomenes, same; D, Adalia, prothorax, ventral; E, Phrynocaria, same; F, Coelophora, mesoventrite; G, Oenopia, mesoventrite; H, M, Cheilomenes sexmaculata, colour patterns; I, Synona melanaria, colour pattern; J, Microcaria salomonensis, colour pattern; K, Coccinella transversalis, colour pattern; L, Oenopia, colour pattern; N, Coelophora, penis; O, Microcaria, penis; P, Coelophora, spermatheca; Q, Microcaria, spermatheca; R, Coccinella leonina, colour pattern; S, Olla, colour pattern; T, Microcaria salomonensis, antenna; U, W, Cheilomenes, antenna; V, Synona, antenna.

130

Ladybird Beetles of the Australo-Pacific Region

Fig. 14.  Key to Australian species: A, F, H, Synona seminigra; B, E, G, Harmonia bicolor; C, Phrynocaria astrolabiana; D, R, Coelophora inaequalis; I, Q, Australoneda bourgeoisi; J, Docimocaria princeps; K, Archegleis kingi; L, Archegleis duplicata; M, Hippodamia variegata: N, Coccinella undecimpunctata; O, Harmonia conformis; P, Adalia bipunctata; S, Microcaria mulsanti; T, Illeis flava; U, Illeis galbula.

Colour plates

131

Fig. 15.  Key to Australian species: A, Micraspis confusa; B, Archegleis edwardsi; C, Archegleis interrupta; D, Heterocaria delta; E, Micraspis flavovittata; F, Micraspis lineola; G, Micraspis frenata; H, Micraspis furcifera; I, Coelophora inaequalis; J, Phrynocaria astrolabiana; K, Oenopia guttata; L, Microcaria jansoni; M, N, Harmonia testudinaria; O, Phrynocaria gratiosa; P, Microcaria mulsanti.

132

Ladybird Beetles of the Australo-Pacific Region

Fig. 16.  Key to Australian species: A, Micraspis flavovittata; B, Micraspis confusa; C, Micraspis furcifera; D, I, J, Cheilomenes sexmaculata; E, R, Micraspis aphidectoides; F, Micraspis frenata; G, Micraspis lineola; H, T, Coccinella transversalis; K, L, Harmonia octomaculata; M, Heterocaria barronensis; N–P, Phrynocaria gratiosa; Q, S, Coelophora inaequalis; U, Heterocaria bella; V, Cleobora mellyi.

Colour plates

133

Fig. 17.  Adalia bipunctata (Linnaeus): A, dorsal view; B, frontal view; C, abdomen; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, female genitalia; I, infundibulum and spermatheca; J, maxilla; K, antenna.

134

Ladybird Beetles of the Australo-Pacific Region

Fig. 18.  Archegleis duplicata (Crotch): A, dorsal view; B, frontal view; C, lateral view; D, abdomen; E, tegmen lateral; F, tegmen ventral; G, female genitalia; H, coxite; I, penis; J, spermatheca.

Colour plates

135

Fig. 19.  Archegleis edwardsi (Mulsant): A, dorsal view; B, lateral view; C, frontal view; D, abdomen; E, female last abdominal ventrite; F, male last abdominal ventrite; G, penis; H, tegmen ventral; I, tegmen lateral; J, female genitalia; K, spermatheca; L, ovipositor.

136

Ladybird Beetles of the Australo-Pacific Region

Fig. 20.  Archegleis interrupta (Weise): A, frontal view; B, dorsal view; C, lateral view; D, abdomen; E, penis; F, tegmen lateral; G, tegmen ventral; H, female genitalia; I, spermatheca; J, ovipositor.

Colour plates

137

Fig. 21.  Archegleis kingi (Macleay): A, dorsal view; B, frontal view; C, lateral view; D, abdomen; E, tegmen ventral; F, tegmen lateral; G, penis; H, female genitalia; I, bursa copulatrix and spermatheca; J, coxite.

138

Ladybird Beetles of the Australo-Pacific Region

Fig. 22.  Australoneda bielawskii Li, Ślipiński & Pang: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, female genitalia.

Colour plates

139

Fig. 23.  Australoneda bourgeoisi (Kerville): A, dorsal view; B, lateral view; C, frontal view; D, spermatheca; E, tegmen ventral; F, tegmen lateral; G, penis; H, ovipositor; I, abdominal postcoxal line.

140

Ladybird Beetles of the Australo-Pacific Region

Fig. 24.  Australoneda fuerschi (Bielawski): A, dorsal view; B, lateral view; C, frontal view; D, spermatheca; E, ovipositor; F, penis; G, penis apex; H, tegmen lateral; I, tegmen ventral.

Colour plates

141

Fig. 25.  Australoneda horni (Korschefsky): A, dorsal view; B, frontal view; C, lateral view; D, spermatheca; E, female genitalia; F, penis; G, tegmen lateral; H, tegmen ventral; I, abdominal postcoxal line.

142

Ladybird Beetles of the Australo-Pacific Region

Fig. 26.  Australoneda karubakana (Bielawski): A–C, variation of colour pattern in dorsal view; D, spermatheca; E, tegmen lateral; F, tegmen ventral; G, penis apex; H, penis; I, abdominal postcoxal line.

Colour plates

Fig. 27.  Australoneda maai (Bielawski): A, lateral view; B, dorsal view; C, coxite; D, spermatheca; E, female genitalia; F, abdominal postcoxal line.

143

144

Ladybird Beetles of the Australo-Pacific Region

Fig. 28.  Australoneda ruitong Li, Slipinski & Pang: A, dorsal view; B, lateral view; C, spermatheca; D, tegmen ventral; E, tegmen lateral; F, penis; G, penis apex; H, female genitalia; I, abdominal postcoxal line.

Colour plates

145

Fig. 29.  Australoneda taengana (Bielawski): A, lateral view; B, dorsal view; C, tegmen lateral; D, tegmen ventral; F, penis; G, spermatheca; H, abdominal postcoxal line.

146

Ladybird Beetles of the Australo-Pacific Region

Fig. 30.  Cheilomenes polynesiae Crotch: A, C, variation of dorsal colour pattern; B, lateral view; D, frontal view; E, female genitalia; F, bursa copulatrix and spermatheca; G, penis; H, penis apex; I, tegmen ventral; J, tegmen lateral; K, abdominal postcoxal line.

Colour plates

147

Fig. 31.  Cheilomenes samoensis Arrow: A, dorsal view; B, lateral view; C, frontal view; D, tegmen lateral; E, tegmen ventral; F, penis apex; G, penis; H, bursa copulatrix and spermatheca; I, ovipositor; J, abdominal postcoxal line.

148

Ladybird Beetles of the Australo-Pacific Region

Fig. 32.  Cheilomenes sexmaculata (Fabricius): A–C, variation of colour pattern in dorsal view; D, lateral view; E, bursa copulatrix and spermatheca; F, ovipositor; G, abdominal postcoxal line; H, tegmen ventral; I, tegmen lateral; J, penis; K, penis apex.

Colour plates

149

Fig. 33.  Cleobora mellyi (Mulsant): A, dorsal view; B, lateral view; C, frontal view; D, spermatheca; E, female genitalia; F, penis; G, tegmen ventral; H, tegmen lateral; I, penis apex; J, coxite.

150

Ladybird Beetles of the Australo-Pacific Region

Fig. 34.  Coccinella leonina Fabricius: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, spermatheca; G, bursa copulatrix; H, ovipositor; I, tegmen ventral; J, tegmen lateral.

Colour plates

151

Fig. 35.  Coccinella transversalis Fabricius: A, dorsal view; B, frontal view; C, lateral view; D, penis apex; E, penis; F, paramere; G, I, tegmen ventral; H, tegmen lateral; J, spermatheca.

152

Ladybird Beetles of the Australo-Pacific Region

Fig. 36.  Coccinella undecimpunctata Linnaeus: A, dorsal view; B, frontal view; C, lateral view; D, ovipositor; E, bursa copulatrix and spermatheca; F, penis; G, penis apex; H, tegmen lateral; I, tegmen ventral; J, abdominal postcoxal line.

Colour plates

153

Fig. 37.  Coelophora bisellata Mulsant: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, paramere; G, tegmen ventral; H, tegmen lateral; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

154

Ladybird Beetles of the Australo-Pacific Region

Fig. 38.  Coelophora inaequalis (Fabricius): A–I, variation of colour pattern in dorsal view; J, penis; K, penis apex; L, tegmen ventral; M, tegmen lateral; N, bursa copulatrix and spermatheca; O, female genitalia.

Colour plates

155

Fig. 39.  Coelophora vesiculata Chazeau: A–B, variation of colour pattern in dorsal view; C, frontal view; D, lateral view; E, spermatheca; F, penis; G, penis apex; H, tegmen ventral; I, tegmen lateral; J, abdominal postcoxal line.

156

Ladybird Beetles of the Australo-Pacific Region

Fig. 40.  Docimocaria aruensis (Crotch): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, paramere; G, tegmen ventral; H, tegmen lateral; I, abdominal postcoxal line.

Colour plates

157

Fig. 41.  Docimocaria princeps (Mulsant): A, dorsal view; B, frontal view; C, lateral view; D, female genitalia; E, spermatheca; F, penis; G, tegmen ventral; H, tegmen lateral; I, penis apex; J, abdominal postcoxal line.

158

Ladybird Beetles of the Australo-Pacific Region

Fig. 42.  Docimocaria pulchra (Crotch): A, dorsal view; B, frontal view; C, lateral view; D, spermatheca; E, female genitalia; F, penis apex; G, penis; H, tegmen lateral; I, tegmen ventral; J, abdominal postcoxal line.

Colour plates

159

Fig. 43.  Harmonia antipodum (Mulsant): A–C, variation of colour pattern in dorsal view; D, frontal view; E, lateral view; F, coxite; G, bursa copulatrix and spermatheca; H, penis; I, penis apex; J, tegmen lateral; K, tegmen ventral; L, abdominal postcoxal line.

160

Ladybird Beetles of the Australo-Pacific Region

Fig. 44.  Harmonia areolata Bielawski: A–B, variation of colour pattern in dorsal view; C, lateral view; D, frontal view; E, bursa copulatrix and spermatheca; F, penis; G, penis apex; H, tegmen lateral; I, tegmen ventral; J, abdominal postcoxal line.

Colour plates

161

Fig. 45.  Harmonia axyridis (Pallas): A–D, variation of colour pattern in dorsal view; E, frontal view; F, lateral view; G, spermatheca; H, penis; I, penis apex; J, tegmen ventral; K, tegmen lateral; L, abdominal postcoxal line.

162

Ladybird Beetles of the Australo-Pacific Region

Fig. 46.  Harmonia basinotata Bielawski: A–B, variation of colour pattern in dorsal view; C, lateral view; D, frontal view; E, penis; F, penis apex; G, tegmen ventral; H, female genitalia; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

163

Fig. 47.  Harmonia bicolor (Blackburn): A–B, variation of colour pattern in dorsal view; C, lateral view; D, frontal view; E, coxite; F, bursa copulatrix and spermatheca; G, penis; H, penis apex; I, tegmen ventral; J, tegmen lateral; K, abdominal postcoxal line.

164

Ladybird Beetles of the Australo-Pacific Region

Fig. 48.  Harmonia boothi sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, bursa copulatrix and spermatheca; E, penis; F, penis apex; G, paramere; H, tegmen lateral; I, tegmen ventral.

Colour plates

165

Fig. 49.  Harmonia conformis (Boisduval): A, dorsal view; B, frontal view; C, lateral view; D, bursa copulatrix and spermatheca; E, female genitalia; F, penis; G, penis apex; H–I, tegmen ventral; J, tegmen lateral; K, abdominal postcoxal line.

166

Ladybird Beetles of the Australo-Pacific Region

Fig. 50.  Harmonia decussata (Crotch): A–D, variation of colour pattern in dorsal view; E, lateral view; F, female genitalia; G, bursa copulatrix and spermatheca; H, penis; I, penis apex; J, tegmen ventral; K, tegmen lateral.

Colour plates

167

Fig. 51.  Harmonia fijiensis (Sicard): A–B, variation of colour pattern in dorsal view; C, lateral view; D, coxite; E, bursa copulatrix and spermatheca; F, abdominal postcoxal line.

168

Ladybird Beetles of the Australo-Pacific Region

Fig. 52.  Harmonia manillana (Mulsant): A–L, variation of colour pattern in dorsal view; M, penis; N, penis apex; O, tegmen ventral; P, tegmen lateral; Q, female genitalia; R, bursa copulatrix and spermatheca; S, abdominal postcoxal line.

Colour plates

169

Fig. 53.  Harmonia micra sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, female genitalia; I, abdominal postcoxal line.

170

Ladybird Beetles of the Australo-Pacific Region

Fig. 54.  Harmonia nigromarginata Bielawski: A–C, variation of colour pattern in dorsal view; D, lateral view; E, frontal view; F, bursa copulatrix and spermatheca; G, female genitalia; H, penis; I, penis apex; J, tegmen ventral; K, tegmen lateral; L, abdominal postcoxal line.

Colour plates

171

Fig. 55.  Harmonia novaehebridensis (Korschefsky): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, bursa copulatrix and spermatheca; I, ovipositor; J, abdominal postcoxal line.

172

Ladybird Beetles of the Australo-Pacific Region

Fig. 56.  Harmonia octomaculata (Fabricius): A–D, variation of colour pattern in dorsal view; E, frontal view; F, penis; G, penis apex; H, tegmen ventral; I, tegmen lateral; J, female genitalia; K, bursa copulatrix and spermatheca; L, abdominal postcoxal line.

Colour plates

173

Fig. 57.  Harmonia ohlmusi sp. nov.: A, dorsal view; B, frontal view; C–D, variation of colour pattern in lateral view; E, penis; F–G, penis apex; H, tegmen ventral; I, tegmen lateral; J, paramere; K, spermatheca.

174

Ladybird Beetles of the Australo-Pacific Region

Fig. 58.  Harmonia tricolor (Korschefsky): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, paramere; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

175

Fig. 59.  Harmonia testudinaria (Mulsant): A–C, variation of colour pattern in dorsal view; D, lateral view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, ovipositor; J, bursa copulatrix and spermatheca; K, abdominal postcoxal line.

176

Ladybird Beetles of the Australo-Pacific Region

Fig. 60.  Harmonia tobleri sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, bursa copulatrix and spermatheca; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, abdominal postcoxal line.

Colour plates

177

Fig. 61.  Heterocaria barronensis (Blackburn) comb. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen lateral; G, tegmen ventral; H, coxite; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

178

Ladybird Beetles of the Australo-Pacific Region

Fig. 62.  Heterocaria bella sp. nov.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, abdominal postcoxal line.

Colour plates

179

Fig. 63.  Heterocaria cinerea (Chazeau) comb. nov.: A–C, variation of colour pattern in dorsal view; D, lateral view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, ovipositor; J, spermatheca; K, abdominal postcoxal line.

180

Ladybird Beetles of the Australo-Pacific Region

Fig. 64.  Heterocaria decemmaculata sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, paramere; H, tegmen lateral; I, abdominal postcoxal line.

Colour plates

Fig. 65.  Heterocaria delta (Weise): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral.

181

182

Ladybird Beetles of the Australo-Pacific Region

Fig. 66.  Heterocaria funerea (Crotch): A–C, variation of colour pattern in dorsal view; D, penis; E, tegmen ventral; F, penis apex; G, tegmen lateral; H, spermatheca; I, coxite; J, abdominal postcoxal line.

Colour plates

183

Fig. 67.  Heterocaria kaszabi (Iablokoff-Khnzorian): A–C, variation of colour pattern in dorsal view; D, lateral view; E, frontal view; F, penis; G, penis apex; H, tegmen lateral; I, spermatheca; J, abdominal postcoxal line.

184

Ladybird Beetles of the Australo-Pacific Region

Fig. 68.  Heterocaria papuana Timberlake: A, dorsal view; B, lateral view; C, frontal view; D, coxite; E, spermatheca; F, abdominal postcoxal line.

Colour plates

185

Fig. 69.  Heterocaria riedeli sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, bursa copulatrix and spermatheca; I, abdominal postcoxal line.

186

Ladybird Beetles of the Australo-Pacific Region

Fig. 70.  Heterocaria samuelsoni (Iablokoff-Khnzorian) comb. nov.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, bursa copulatrix and spermatheca; I, abdominal postcoxal line.

Colour plates

187

Fig. 71.  Heterocaria transversoguttata sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, paramere; H, tegmen lateral; I, abdominal postcoxal line.

188

Ladybird Beetles of the Australo-Pacific Region

Fig. 72.  Heteroneda grata (Iablokoff-Khnzorian) comb. nov.: A–C, variation of colour pattern in dorsal view; D, lateral view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, ovipositor; J, bursa copulatrix and spermatheca; K, abdominal postcoxal line.

Colour plates

189

Fig. 73.  Heteroneda mimetica Chazeau: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen lateral; G, tegmen ventral; H, female genitalia; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

190

Ladybird Beetles of the Australo-Pacific Region

Fig. 74.  Heteroneda principalis (Weise): A–D, variation of colour pattern in dorsal view; E, frontal view; F, lateral view; G, penis; H–I, penis apex; J, tegmen ventral; K, tegmen lateral; L, bursa copulatrix and spermatheca; M, abdominal postcoxal line.

Colour plates

191

Fig. 75.   Hippodamia variegata (Goeze): A, dorsal view; B, D, variation of colour pattern in frontal view; C, lateral view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, bursa copulatrix and spermatheca; J, coxite; K, abdominal postcoxal line.

192

Ladybird Beetles of the Australo-Pacific Region

Fig. 76.  Hysia endomycina (Boisduval): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, ovipositor; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

193

Fig. 77.  Illeis flava Pope: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, antenna; I, maxillary palp; J, abdominal postcoxal line.

194

Ladybird Beetles of the Australo-Pacific Region

Fig. 78.  Illeis galbula (Mulsant): A–D, variation of colour pattern in dorsal view; E, penis; F, penis apex; G, tegmen lateral; H, ovipositor; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

195

Fig. 79.  Illeis gressitti Bielawski: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, abdominal postcoxal line.

196

Ladybird Beetles of the Australo-Pacific Region

Fig. 80.  Illeis wyliei sp. nov.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen lateral; G, tegmen ventral; H, bursa copulatrix and spermatheca.

Colour plates

197

Fig. 81.  Megalocaria rufitarsis (Sicard): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, ovipositor; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

198

Ladybird Beetles of the Australo-Pacific Region

Fig. 82.  Megalocaria singularis (Weise): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, penis guide apex lateral; H, ovipositor; I, bursa copulatrix and spermatheca.

Colour plates

199

Fig. 83.  Megalocaria tricolor (Fabricius): A–B, variation of colour pattern in dorsal view; C, lateral view; D, frontal view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, ovipositor; J, spermatheca; K, abdominal postcoxal line.

200

Ladybird Beetles of the Australo-Pacific Region

Fig. 84.  Micraspis allardi (Mulsant): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, ovipositor; I, spermatheca; J, abdominal postcoxal line.

Colour plates

201

Fig. 85.  Micraspis aphidectoides (Korschefsky): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, paramere; H, tegmen lateral; I, coxite; J, spermatheca; K, abdominal postcoxal line.

202

Ladybird Beetles of the Australo-Pacific Region

Fig. 86.  Micraspis bennigseni (Weise): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, female genitalia; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

203

Fig. 87.  Micraspis confusa Pope: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, abdominal postcoxal line.

204

Ladybird Beetles of the Australo-Pacific Region

Fig. 88.  Micraspis flavovittata (Crotch): A–B, variation of colour pattern in dorsal view; C, frontal view; D, lateral view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, bursa copulatrix and spermatheca.

Colour plates

205

Fig. 89.  Micraspis frenata (Erichson): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, female genitalia; H, bursa copulatrix and spermatheca.

206

Ladybird Beetles of the Australo-Pacific Region

Fig. 90.  Micraspis furcifera (Guérin-Méneville): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, female genitalia; I, spermatheca, J, abdominal postcoxal line.

Colour plates

207

Fig. 91.  Micraspis lineata (Thunberg): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, spermatheca; I, ovipositor; J, abdominal postcoxal line.

208

Ladybird Beetles of the Australo-Pacific Region

Fig. 92.  Micraspis lineola (Fabricius): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen lateral; G, tegmen ventral; H, coxite.

Colour plates

209

Fig. 93.  Microcaria jansoni (Crotch): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, penis middle section; G, tegmen ventral; H–I, tegmen lateral; J, coxite; K, bursa copulatrix and spermatheca; L, abdominal postcoxal line.

210

Ladybird Beetles of the Australo-Pacific Region

Fig. 94.  Microcaria mulsanti (Montrouzier): A–D, variation of colour pattern in dorsal view; E–F, variation of colour pattern in frontal view; G, lateral view; H, penis; I–K, penis apex; L–M, tegmen ventral; N, tegmen lateral; O, spermatheca; P, abdominal postcoxal line.

Colour plates

211

Fig. 95.  Microcaria salomonensis (Korschefsky): A, dorsal view; B, frontal view; C, lateral view; D, spermatheca; E, coxite; F, penis; G, penis apex; H, tegmen ventral; I, tegmen lateral; J, antenna; K, maxillary palp; L, abdominal postcoxal line.

212

Ladybird Beetles of the Australo-Pacific Region

Fig. 96.  Microcaria versipellis (Mulsant): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis middle section; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, spermatheca; J, ovipositor; K, abdominal postcoxal line.

Colour plates

213

Fig. 97.  Microcaria vivida (Crotch): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, spermatheca; I, abdominal postcoxal line.

214

Ladybird Beetles of the Australo-Pacific Region

Fig. 98.  Oenopia guttata (Blackburn): A–B, variation of colour pattern in dorsal view; C, lateral view; D, frontal view; E, penis; F, penis apex; G, tegmen ventral; H, tegmen lateral; I, female genitalia; J, bursa copulatrix and spermatheca; K, abdominal postcoxal line.

Colour plates

215

Fig. 99.  Olla v-nigrum (Mulsant): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, coxite; H, spermatheca; I, abdominal postcoxal line.

216

Ladybird Beetles of the Australo-Pacific Region

Fig. 100.  Phrynocaria ambonina sp. nov.: A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, paramere; H, spermatheca; I, abdominal postcoxal line.

Colour plates

217

Fig. 101.  Phrynocaria aurita (Iablokoff-Khnzorian): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, ovipositor; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

218

Ladybird Beetles of the Australo-Pacific Region

Fig. 102.  Phrynocaria astrolabiana (Weise): A–E, variation of colour pattern in dorsal view; F, penis; G, penis middle section; H, penis apex; I, tegmen lateral; J, tegmen ventral; K, ovipositor; L, bursa copulatrix and spermatheca; M, abdominal postcoxal line.

Colour plates

219

Fig. 103.  Phrynocaria crotchi Li et al.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, paramere; I, ovipositor; J, bursa copulatrix and spermatheca; K, abdominal postcoxal line.

220

Ladybird Beetles of the Australo-Pacific Region

Fig. 104.  Phrynocaria georgia sp. nov.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, spermatheca; I, abdominal postcoxal line.

Colour plates

221

Fig. 105.  Phrynocaria gratiosa (Mulsant): A–C, variation of colour pattern in dorsal view; D, frontal view; E, lateral view; F, penis; G, penis apex; H, penis middle section; I, K, tegmen ventral; J, tegmen lateral; L, bursa copulatrix and spermatheca; M, abdominal postcoxal line.

222

Ladybird Beetles of the Australo-Pacific Region

Fig. 106.  Phrynocaria kolombangara sp. nov.: A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, bursa copulatrix and spermatheca; I, abdominal postcoxal line.

Colour plates

223

Fig. 107.  Phrynocaria moseri (Weise): A–B, variation of colour pattern in dorsal view; C–D, variation of colour pattern in lateral view; E, penis; F, penis apex; G, tegmen ventral; H, paramere; I, tegmen lateral; J, bursa copulatrix and spermatheca; K, abdominal postcoxal line.

224

Ladybird Beetles of the Australo-Pacific Region

Fig. 108.  Phrynocaria quadrivittata (Fauvel): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, tegmen ventral; F, tegmen lateral; G, paramere; H, ovipositor; I, bursa copulatrix and spermatheca; J, abdominal postcoxal line.

Colour plates

225

Fig. 109.  Phrynocaria wallacii (Crotch): A, dorsal view; B, frontal view; C, lateral view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, paramere; I, bursa copulatrix and spermatheca; J, ovipositor; K, abdominal postcoxal line.

226

Ladybird Beetles of the Australo-Pacific Region

Fig. 110.  Subepilachna latemarginata (Bielawski): A, dorsal view; B, lateral view; C, frontal view; D, penis; E, penis apex; F, tegmen ventral; G, tegmen lateral; H, spermatheca; I, coxite; J, maxillary palp; K, mandible; L, abdominal postcoxal line.

Colour plates

227

Fig. 111.  Synona melanaria (Mulsant): A–B, variation of colour pattern in dorsal view; C–D, variation of colour pattern in lateral view; E, frontal view; F, penis; G, tegmen lateral; H, tegmen ventral; I, female genitalia; J, antenna; K, abdominal postcoxal line.

This page intentionally left blank

INDEX OF SCIENTIFIC NAMES abdominalis  97 Acacia  17, 48, 61 Adalia  6, 8, 10, 12, 14, 15, 31, 114, 127, 128, 129, 130, 133 Adonia  71, 72 albolineata  89 Alesia  81, 85, 86, 87 allardi  8, 82, 83, 200 ambonina  6, 9, 99, 100 Anisolemnia  41, 79, 80 anthracina  110, 111 Antineda  6, 38, 39, 40 antipodum  8, 43, 44, 159 aphidectoides  8, 13, 82, 83, 132, 201 Archaeoneda  78 Archegleis  8, 10, 12, 15, 16, 17, 18, 19, 60, 61, 63, 125, 128, 130, 131, 134, 135, 136, 137 arcuata  54 areolata  8, 43, 44, 45, 49, 160 Artemis  93 artensis  37 aruensis  6, 8, 39, 40, 156 astrolabiana  9, 12, 13, 99, 101, 114, 130, 131, 218 aterrima  51 atrocincta  51 auberti  6, 94, 95 aurita  6, 9, 99, 100, 217 australis  28 Australoneda  8, 10, 12, 19, 20, 21, 22, 23, 24, 25, 114, 117, 128, 130, 138, 139, 140, 141, 142, 143, 144, 145 Autotela  104 axyridis  7, 6, 7, 8, 42, 43, 45, 112, 115, 120, 161 Ballia  42 barronensis  6, 8, 13, 16, 60, 61, 66, 177 basinotata  6, 8, 43, 45, 46, 162 bella  6, 8, 13, 60, 61, 62, 132, 178 bennigseni  8, 82, 84, 202 bicolor  8, 12, 43, 46, 47, 130, 163 bicruciata  86 bielawskii  8, 20, 21, 138 billardieri  68, 69, 70, 112 bimaculata  30, 112 bipunctata  6, 8, 12, 14, 15, 114, 130, 133 bisellata  8, 35, 36, 37, 153 boothi  6, 8, 43, 47, 164 Bothrocalvia  6, 89 bourgeoisi  8, 12, 19, 20, 21, 117, 130, 139 Brachychiton  120 brassicae  15 Brevicoryne  15 Callineda  6, 16, 17, 40, 42, 48, 61, 70, 71 cardinalis  6 Caria  6, 41, 51, 79, 80, 116 cassidoides  110, 111 Castanopsis  67 catan  111 Catanus  111 Centrosema  111 Ceolophora  86 cerasicolor  51 charybdis  6, 31 Cheilomenes  10, 13, 25, 26, 27, 28, 29, 122, 129, 132, 146, 147, 148 christophori  42 Chrysophtharta  30, 112 cincta  74 cinctella  95 cinerea  6, 8, 60, 62, 63, 179 Cisseis  81, 87

Cissella  81, 87 citri  115 Cleobora  iv, 6, 11, 13, 29, 30, 114, 119, 122, 132, 149 Cleodora  29, 30 Coccinella  iv, 3, 6, 7, 10, 12, 13, 14, 15, 18, 19, 25, 28, 31, 32, 33, 34, 35, 36, 37, 38, 42, 43, 45, 47, 50, 54, 68, 71, 72, 73, 74, 78, 80, 81, 86, 87, 88, 89, 91, 95, 97, 98, 110, 111, 112, 113, 114, 116, 119, 121, 122, 127, 129, 130, 132, 150, 151, 152 Coccus  99, 107 Coelophora  6, 11, 12, 13, 35, 36, 37, 38, 39, 41, 53, 56, 60, 64, 90, 91, 92, 93, 94, 95, 96, 99, 101, 103, 104, 105, 106, 107, 108, 110, 111, 112, 113, 114, 118, 119, 121, 128, 129, 130, 131, 132, 153, 154, 155 Coleomegilla  6, 114 comma  81 conformis  4, 6, 8, 12, 43, 47, 48, 115, 118, 119, 120, 122, 125, 130, 165 confusa  8, 13, 82, 84, 85, 131, 132, 203 congener  98 conglobata  95 Coptosoma  6, 112 coriacea  32 coryphaea  42 crotchi  9, 63, 64, 99, 102, 107, 219 Cryptocephalus  112 Cryptolaemus  6 csikii  76 Cumare  110, 111 Cycloneda  6, 61 Cydonia  26 Daulis  57, 58, 89, 91 decemmaculata  6, 8, 60, 63, 180 decussata  6, 8, 42, 43, 44, 46, 48, 49, 51, 52, 70, 166 delta  6, 8, 13, 16, 17, 18, 60, 63, 64, 131, 181 desolata  37, 38 Diaphorina  115 dilatata  79 dimidiata  42 Docimocaria  6, 8, 10, 12, 38, 39, 40, 41, 42, 51, 60, 66, 79, 117, 125, 127, 128, 130, 156, 157, 158 dunlopi  51 duodecimpunctata  28, 29 duplicata  8, 12, 16, 17, 18, 130, 134 Dysis  115 edwardsi  8, 12, 16, 17, 18, 125, 131, 135 Egleis  6, 17, 18, 61, 63, 64, 74, 114 elegans  104 Elpis  26 endomycina  8, 73, 74, 192 Eucalyptus  112 excellens  115 extensa  6, 64 fijiensis  6, 8, 43, 50, 80, 167 flava  8, 12, 75, 77, 130, 193 flavofasciata  28 flavoguttata  104 flavolineata  87 flavomaculata  51 flavomarginata  48, 49 flavovittata  9, 13, 82, 85, 119, 131, 132, 204 formosa  42 fragrantissima  75 frenata  4, 9, 13, 82, 86, 88, 113, 118, 125, 131, 132, 205 fuerschi  8, 20, 22, 140 funerea  6, 8, 60, 64, 65, 182 furcifera  9, 12, 81, 82, 87, 131, 132, 206

230

Ladybird Beetles of the Australo-Pacific Region

galbula  4, 8, 12, 74, 75, 76, 112, 118, 120, 122, 125, 130, 194 gauthardi  87 georgia  6, 9, 99, 103, 220 grata  6, 8, 69, 70, 71, 188 gratiosa  9, 13, 99, 104, 114, 131, 132, 221 gressitti  8, 70, 71, 75, 77, 112, 195 guttata  9, 13, 95, 96, 131, 214 Gynocaria  96 Halyzia  6, 17, 18, 30, 64, 76 Harmonia  iv, vi, vii, 6, 7, 10, 12, 13, 16, 39, 42, 43, 44, 45, 46, 47, 48, 49, 50, 51, 52, 53, 54, 55, 56, 57, 58, 59, 61, 69, 70, 94, 97, 104, 110, 111, 112, 115, 119, 120, 121, 122, 125, 127, 130, 131, 132, 159, 160, 161, 162, 163, 164, 165, 166, 167, 168, 169, 170, 171, 172, 173, 174, 175, 176 Heterocaria  6, 10, 13, 16, 17, 18, 59, 60, 61, 62, 63, 64, 65, 66, 67, 68, 128, 131, 132, 177, 178, 179, 180, 181, 182, 183, 184, 185, 186, 187 Heteroneda  6, 11, 68, 69, 70, 71, 112, 188, 189, 190 Hibbertia  98 Hippodamia  7, 10, 12, 71, 72, 113, 118, 127, 130, 191 horni  8, 20, 22, 23, 24, 141 Hyperomyzus  16, 20, 60, 99 Hysia  10, 73, 74, 127, 192 Icerya  6, 56 Idalia  14, 15 Illeis  iv, vi, 4, 5, 6, 8, 10, 12, 74, 75, 76, 77, 78, 112, 114, 118, 120, 122, 125, 127, 130, 193, 194, 195, 196 inaequalis  8, 12, 13, 35, 36, 37, 38, 113, 114, 118, 119, 121, 130, 131, 132, 154 incognita  6, 48, 49, 104 incompleta  51 insignis  38 interrupta  8, 12, 16, 17, 18, 131, 136 jansoni  9, 13, 46, 90, 91, 104, 131, 209 karubakana  8, 20, 23, 25, 117, 142 kaszabi  6, 8, 60, 62, 63, 65, 183 kingi  8, 12, 15, 16, 17, 18, 19, 125, 130, 137 koebelei  104 kolombangara  6, 9, 99, 104, 105, 222 korschefskyi  79 Lagerstromia  75 latemarginata  9, 108, 109, 226 Leaucaena  98 Lecanium  99 Leis  42, 43, 45, 47, 48, 51 Lemnia  6, 35, 37, 38, 60, 61, 62, 64, 65, 69, 70, 71, 82, 90, 91, 99, 100, 104, 105, 107, 109, 110, 111 leonina  7, 8, 31, 32, 119, 150 Leptospermum  18 Leptothea  74, 76, 112 Leucaena  98 leucocephala  98 lineata  9, 82, 86, 88, 112, 207 lineola  9, 13, 82, 88, 89, 131, 132, 208 lividigaster  112 longifolia  48 Lonicera  75 Lophomytus  48 lucens  98 lunata  25 maai  8, 20, 24, 143 maculata  114 malaccensis  82, 83 manillana  8, 39, 43, 48, 51, 115, 168 marginepunctata  42 mastersi  37, 38 Megalocaria  10, 39, 78, 79, 80, 81, 128, 197, 198, 199 melanaria  6, 9, 12, 109, 110, 111, 112, 129, 227

melanoxylon  18 mellyi  4, 6, 8, 13, 29, 30, 114, 119, 122, 132, 149 Menevillidia  87 Menochilus  25, 26, 28, 115 micra  6, 8, 43, 51, 52, 169 Micraspis  iv, 4, 6, 10, 12, 13, 81, 82, 83, 84, 85, 86, 87, 88, 89, 112, 113, 115, 118, 119, 125, 129, 131, 132, 200, 201, 202, 203, 204, 205, 206, 207, 208 Microcaria  6, 4, 11, 12, 13, 35, 46, 53, 89, 90, 91, 92, 93, 94, 95, 104, 117, 127, 128, 129, 130, 131, 209, 210, 211, 212, 213 Microlemnia  6, 35, 70, 71, 100 mimetica  8, 69, 70, 189 minuta  96 mniszechii  51 mollipes  28 montana  95 montrouzieri  6 moseri  6, 9, 76, 99, 105, 106, 223 Muehlenbeckia  88 mulsanti  9, 12, 13, 89, 90, 91, 92, 106, 117, 130, 131, 210 mutabilis  71 mysolensis  6, 64 Neda  6, 19, 21, 22, 23, 24, 25, 38, 40, 46, 47, 51, 57, 61, 78, 80, 112, 114 nigrocincta  104 nigromarginata  6, 8, 43, 44, 70, 49, 52, 53, 170 nigrovittata  104 novaehebridensis  8, 43, 53, 54, 94, 171 novaezealandiae  34 novemmaculata  37 novempunctata  37 occidentalis  48 ochracea  91, 92 octomaculata  4, 8, 13, 43, 54, 55, 121, 125, 132, 172 Oenocaria  96 Oenopia  11, 13, 50, 95, 96, 115, 116, 129, 131, 214 ohlmusi  6, 8, 43, 55, 56, 173 Oidium  75 Oiocaria  6, 16, 59, 60, 67 Olla  10, 12, 97, 98, 116, 129, 215 ostensum  112 palens  104 pallida  110 papuana  8, 60, 66, 184 papuensis  51 Paropsis  6, 31 pascoei  17 paulinae  51 Petalostigma  111 Phrynocaria  6, 10, 12, 13, 98, 99, 100, 101, 102, 103, 104, 105, 106, 107, 114, 128, 129, 130, 131, 132, 216, 217, 218, 219, 220, 221, 222, 223, 224, 225 Phrynolemnia  98 picturata  46 Plumeria  107 polynesiae  8, 26, 27, 28, 146 populneus  120 princeps  6, 8, 12, 38, 39, 40, 41, 117, 125, 130, 157 principalis  6, 8, 69, 70, 71, 190 Protocaria  95 Pseudoharmonia  95 Psyllobora  74, 76 Ptychanatis  42, 45, 54 pubescens  111 pulchra  6, 8, 39, 41, 42, 158 punctata  37 purchasi  6 quadriplagiata  28 quadrivittata  6, 9, 99, 106, 112, 224

Index of scientific names

reichei  78, 79 religiosa  37, 38 repanda  33 reticulata  68 rhopalocera  6, 52, 53 Rhopaloneda  42 riedeli  6, 8, 60, 66, 67, 185 ripponi  37, 38 Rodolia  6 rufitarsis  8, 79, 197 ruitong  8, 20, 24, 25, 117, 144 salomonensis  6, 4, 9, 90, 92, 93, 129, 211 samoensis  8, 26, 27, 28, 147 samuelsoni  6, 8, 59, 60, 67, 186 scalaris  95 schultzei  60 Scymnodes  112 sedecimnotata  42 seminigra  110, 111, 130 septempunctata  6, 31, 113 sexmaculata  8, 13, 26, 28, 29, 122, 129, 132, 148 sexmaculatus  28 Singhikalia  108 singularis  8, 79, 80, 198 solomonensis  58 Sospita  87 Spilocaria  35, 36 Stictoleis  42 striata  81 strigula  88 Subepilachna  4, 10, 108, 109, 122, 127, 226 Synharmonia  6, 50, 95 Synia  109, 110, 111, 112 Synona  4, 6, 10, 12, 109, 110, 115, 127, 128, 129, 130, 227

taengana  8, 20, 23, 25, 145 tasmanii  32 testudinaria  4, 8, 13, 43, 46, 47, 57, 58, 70, 121, 131, 175 tetrasticta  79 Thea  76 thoracica  6, 41, 42 tibialis  79 tobleri  6, 8, 43, 56, 58, 59, 176 transversalis  4, 8, 13, 31, 33, 121, 129, 132, 151 transversoguttata  6, 8, 60, 68, 187 tredecimpunctata  71 tricolor  6, 8, 10, 43, 56, 57, 78, 79, 80, 81, 174, 199 tutuilensis  27 undecimpunctata  6, 7, 8, 12, 31, 32, 34, 72, 113, 122, 152 uninotata  6, 45, 46 varicolor  18, 19, 106 variegata  7, 8, 12, 72, 113, 118, 130, 191 Verania  81, 82, 83, 84, 85, 86, 87, 88 veranioides  37, 38, 113 versipellis  6, 9, 90, 93, 94, 212 vesiculata  8, 36, 37, 38, 155 vidua  37, 38 viridis  99, 107 vittipennis  86 vivida  6, 9, 53, 90, 94, 95, 213 v-nigrum  9, 12, 97, 98, 215 wallacei  102, 103, 107, 108 wallacii  9, 99, 102, 103, 107, 108, 225 wyliei  6, 8, 75, 77, 196

231