Ladybird Beetles of the Australo-Pacific Region: Coleoptera: Coccinellidae: Coccinellini

By Adam Slipinski, Jiahui Li and Hong Pang

True ladybirds, classified in the tribe Coccinellini, are easily recognisable by their relatively large and shiny bodies and contrasting colour patterns. They are one of the most widely studied groups of beetles, being of economic importance and used as model organisms in biological and ecological research.

Ladybird Beetles of the Australo-Pacific Region covers 22 genera and 95 valid species, including 12 new species, of ladybird beetles from Australia, New Guinea and the Pacific area. For each species, descriptions, illustrations and keys will assist with the correct identification of ladybirds from this large but practically unknown fauna.

This book is a valuable contribution to the taxonomy of the ladybirds and to the knowledge of the biodiversity of this unique biogeographic region. It will be of use to entomologists, biologists, ecologists, quarantine officers, natural history museum curators, and students.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Mar 2, 2020
• ISBN: 9781486303892

Adam Slipinski

Adam Slipinski completed his PhD and DSc in Poland, where he worked for 20 years at the Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw. He is currently working as a senior principal research scientist and curator at the Australian National Insect Collection, CSIRO. He is the author of over 200 research publications and multiple book chapters, and author of six books on the phylogeny and classification of various beetles, including Australian Longhorn Beetles (Coleoptera: Cerambycidae) Volumes 1 and 2 (CSIRO Publishing, 2013 and 2016), Australian Beetles Volumes 1 and 2 (CSIRO Publishing, 2013 and 2019), and Ladybird Beetles of the Australo-Pacific Region (CSIRO Publishing, 2020).

Book preview

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-Up-Photolibrary); 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.

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 Zürcher-Pfander)

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 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, free-moving 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 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 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