Australian Longhorn Beetles (Coleoptera: Cerambycidae) Volume 1: Introduction and Subfamily Lamiinae

By Adam Slipinski and Hermes Escalona

Longhorn Beetles — Cerambycidae are one of the most easily recognised groups of beetles, a family that worldwide encompasses over 33,000 species in 5,200 genera. With over 1,400 species classified in 300 genera, this is the sixth largest among 117 beetle families in Australia.

These beetles often attack and kill living forest or orchard trees and develop in construction timber (like European House borer, introduced to WA), causing serious damages. Virtually all Cerambycidae feed on living or dead plant tissues and play a significant role in all terrestrial environments where plants are found. Larvae often utilise damaged or dead trees for their development, and through feeding on rotten wood form an important element of the saproxylic fauna, speeding energy circulation in these habitats. Many species are listed as quarantine pests because of their destructive role to the timber industry.

This volume provides a general introduction to the Australian Cerambycidae with sections on biology, phylogeny and morphology of adult and larvae, followed by the keys to the subfamilies and an overview of the 74 genera of the subfamily Lamiinae occurring in Australia. All Lamiinae genera are diagnosed, described and illustrated and an illustrated key to their identification is provided. A full listing of all included Australian species with synonymies and bibliographic citations is also included.

Biologists worldwide, curators and staff at natural history museums, quarantine/inspection services, entomologists and collectors - many of these beetles are collector's items.

Winner of the 2016 J.O. Westwood Medal
Winner of the 2014 Whitley Medal

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Sep 20, 2013
• ISBN: 9781486300051

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

Australian Longhorn Beetles

(Coleoptera: Cerambycidae)

Volume 1

Introduction and Subfamily Lamiinae

Adam Ślipiński

CSIRO Australian National Insect Collection

&

Hermes E. Escalona

Museo del Instituto de Zoologia Agricola

Universidad Central de Venezuela

© Commonwealth of Australia 2013

This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be reproduced by any process without prior written permission from the Commonwealth. Requests and inquiries concerning reproduction and rights should be addressed to:

Director

Australian Biological Resources Study

GPO Box 787

Canberra ACT 2601

Australia

Email [e-mail]

Disclaimer

The views and opinions expressed in this publication are those of the authors and do not necessarily reflect those of the Australian Government, the Minister for Sustainability, Environment, Water, Population and Communities, the Minister for Agriculture, Fisheries and Forestry, Commonwealth Scientific and Industrial Research Organisation (CSIRO) or CSIRO Publishing.

While reasonable efforts have been made to ensure that the contents of this publication are factually correct, the Commonwealth, CSIRO and CSIRO Publishing do not accept responsibility for the accuracy or completeness of the contents, and shall not be liable for any loss or damage that may be occasioned directly or indirectly through the use of, or reliance on, the contents of this publication.

This publication is available from

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National Library of Australia Cataloguing-in-Publication entry

Ślipiński, Adam, author.

Australian longhorn beetles (coleoptera: cerambycidae). Volume 1, Introduction and subfamily lamiinae / Adam Slipinski and Hermes E. Escalona.

978 1 486 30003 7 (hardback)

978 1 486 30004 4 (ePDF)

978 1 486 30005 1 (ePub)

Includes bibliographical references and index.

Cerambycidae – Australia. Beetles – Australia. Escalona, Hermes E., author.

595.76480994

Contents

Abstract

Acknowledgements

Material and methods

Family Cerambycidae

Introduction

Morphology of adult beetles

Morphology of larvae and pupae

Biology and ecology

Economic importance

Geographic distribution

Phylogeny and classification

Australian Cerambycidae

History of research

Higher classification of Australian Cerambycidae

Diagnosis of Family Cerambycidae

Keys to subfamilies of Australian Cerambycidae

Adults

Larvae

Subfamily Lamiinae

Classification of Australian Lamiinae

Diagnosis of Subfamily Lamiinae

Key to adults of genera of Lamiinae in Australia

Review of the Australian genera of Lamiinae

Lamiinae figures

Appendix 1: New synonymies

Appendix 2: New generic combinations proposed for Australian species

Appendix 3: Type specimens, Australian and extra-Australian

Bibliography

Index of scientific names

Dedication

We dedicate this book to our mentor DR JOHN F. LAWRENCE, as a tribute to his scientific achievements and to celebrate his upcoming 80th Birthday next year. Dr Lawrence’s work has informed and inspired entomologists worldwide; it will endure.

Abstract

Longhorn Beetles — Cerambycidae are one of the most easily recognised groups of beetles, a family that worldwide encompasses over 33,000 species in 5,200 genera. With over 1,400 species classified in 300 genera, this is the sixth largest among 117 beetle families in Australia.

This volume provides a general introduction to the Australian Cerambycidae with sections on biology, phylogeny and morphology of adult and larvae, followed by the keys to the subfamilies and an overview of the 74 genera of the subfamily Lamiinae occurring in Australia. All Lamiinae genera are diagnosed, described and illustrated and an illustrated key to their identification is provided. A full listing of all included Australian species with synonymies and bibliographic citations is also included.

Three new genera are proposed: Alice gen. nov. for Alice dekeyzeri sp. nov.; Annemabel gen. nov. for Zygocera elongata Breuning, 1939, and Carteridion gen. nov. for Carteridion wilsonensis sp. nov. Three genera: Australoleiopus Breuning, 1970, Temnolamia Breuning, 1961 and Tetroreopsis Breuning, 1940 are treated as genera incertae sedis, and a further two, Elongatocontoderus Breuning, 1977 and Pseudocalamobius Kraatz, 1879, probably do not occur in the Australian fauna.

Seventy-five genera and 13 species (Appendix 1) are synonymised, and 165 new generic combinations are proposed for the Australian species (Appendix 2). Twelve replacement names are proposed for the junior secondary homonyms: Microlamia viridis Ślipiński & Escalona, for Parapteridotelus norfolkensis Breuning, 1962; Rhytiphora callosa Ślipiński & Escalona for Menyllus maculicornis Pascoe, 1864; R. confusa Ślipiński & Escalona for Paradaxata spinosa Breuning, 1938; R. denisoniana Ślipiński & Escalona for Hathliodes moratus Pascoe, 1866; R. elongatissima Ślipiński & Escalona for Prosoplus elongata Breuning, 1938; R. molloiensis Ślipiński & Escalona for Prosoplus (Prosoplus) demarzi Breuning, 1963; R. pascoei Ślipiński & Escalona for Penthea (Melanopenthea) obscura Breuning, 1961; R. onychina Ślipiński & Escalona for Etaxalus marmoratus Breuning, 1950a; Sybra arnhemensis Ślipiński & Escalona for Mimosybra continentalis Breuning, 1961; S. gaindahensis Ślipiński & Escalona for Mimosybra uniformis Breuning, 1940; S. gressitti Ślipiński & Escalona for Sybra (Sybra) albovittata Breuning, 1943 and Temnosternus mosaicus Ślipiński & Escalona for Tuberothelais flavolineatus Breuning, 1963.

Neissa inconspicua Pascoe, 1866 is here designated as the type species of Neissa Pascoe, 1866. Sybra cinerascens Breuning, 1948 was described from the Tapapatauai Island and has been wrongly recorded from Australia.

Acknowledgements

The Australian Biological Resources Study (ABRS), through its National Taxonomy Research Grant Program (NTRGP), funded our research on the Australian Cerambycidae which was undertaken with generous support from the CSIRO Ecosystem Sciences. The Department of Agriculture, Fisheries and Forestry (DAFF), and the Australian Biological Resources Study (ABRS) provided resources and help towards publication of this volume. CSIRO Library services provided innumerable references through interlibrary loans and attended to our requests with patience and professionalism.

This book would have never been published in the present form without the help and very generous support of Alice Wells (ABRS) who provided constant editorial support and advice throughout the long period of completing the book. Alice critically read and corrected the entire text, greatly improving the language and clarity of our writing. She has been very understanding and resilient while carrying out the stressful duty of pushing us to the finish line. Brigitte Kuchlmayr (ABRS) expertly designed and set up the book, which is very much appreciated.

The critical help of many colleagues at CSIRO Ecosystem Sciences is gratefully acknowledged, foremost that of Anne Hastings for her artwork, photographs and help with graphics side of the book. Cate Lemann provided constant support in our day-today work at ANIC and helped with hundreds of photographs of beetles, dissections and keys to genera. John F. Lawrence critically read the morphology chapter, providing many corrections and suggestions. Tom Weir and Kim Pullen tested the key to the genera and shared their extensive knowledge on Australian beetles. Rolf Oberprieler has helped with translations from German and Latin. Marianne Horak has supported us with her friendship and stimulating discussions.

Andrew Calder during his tenure at CSIRO initiated studies on the Australian Cerambycidae and his notes, draft checklists, databases, references and his curatorial work in ANIC greatly facilitated our research.

We are very grateful to the following curators and their institutions for loan of critical material for our study and for their support and assistance during our visits to their collections: Derek Smith, Chris Reid (AM); Lee Herman (AMNH); David Kavanaugh, Jere Schweikert (CAS); Max Barclay, Roger Booth (BMNH); Shepherd Myers (BPBM); François Gernier (CMN); Patrice Bouchard (CNC); Peter Gillespie, Graham Goodyer, Christine Stone (DARI); Eduard Vives (EVC); Dirk Ahrens (FAK); Ottó Merkl, Aranka Grabant (HNHM); Phil Perkins (MCZ); Giulio Cuccodoro (MHNG); Azadeh Taghavian (MNHN); Roberto Poggi, Maria Tavano (MSNG); Ken Walker (MV); the late Michel Brancucci, Daniel Burckhardt, Eva Sprecher, Isabelle Zürcher (NHMB); Bert Viklund (NRM); Gavin Dally (NTMD); Haidee Brown (NTDA); Justin Bartlett (QDPIB); Geoff Monteith, Geoff Thompson (QMB); Alain Drumont (RBI); Peter Hudson (SAM); Steve Lingafelter (USNM); Joahim Willers (ZMB); Ralph Peters (Zoologishes Museum Hamburg).

Our heartfelt thanks go to our colleagues worldwide working on Cerambycidae: Steven Lingafelter (USDA, USA) supported HE’s visit to the USNM through the Smithsonian Institution grant and provided help and advice on the project; Petr Švácha (Czech Academy of Sciences, Czech Republic) generously shared his enormous knowledge of longhorns including drafts of his chapters for the Handbook of Zoology; Duane McKenna (University of Memphis, USA) shared his unpublished data on molecular phylogeny of Cerambycidae; Eugenio Nearns (USDA, USA) assisted with important references and contacts; Roger de Keyzer (Sydney) provided many important specimens; Nathan Lord (University of New Mexico, USA) helped to test the keys.

Geoff Monteith, Federica Turco (QM), Nicole Gunter (CSIRO), Nadine Guthrie (Department of Wildlife and Conservation, Wanneroo, WA), Sara Pinzon-Navarro (CSIRO), Jack Hasenpush, Natalie Banks, David Rentz and Alberto Venchi helped in many ways to collect Australian longhorn beetles in the field. Geoff Monteith has been instrumental to successful collecting of many critical Cerambycidae, always sharing his enthusiasm and enormous knowledge of the Australian insects and their natural histories. Penny Edwards and Max Whitten provided larvae, adults and interesting observations on biology of native Prionine Cacodacnus planicollis Blackburn damaging pine studs in a house in Maleny (Queensland).

Ben Boyd and Graham Teakle (DAFF Biosecurity — CSIRO Liaison Entomologists) supplied specimens of Cerambycidae of quarantine concern and provided other technical support. Mark Talbot and Rosemary White (CSIRO Plant Industry) are sincerely acknowledged for their help, patience and training with the SEM. Mark Hoddle, Steve Dreistadt, Cheryl Reynolds (University of California, Davis) helped us to obtain pictures of Phoracantha damage.

We are very grateful to the following photographers for providing photographs of living longhorn beetles: Kaisa and Stan Breeden; Peter Chew; Jack Kelly Clark (courtesy University of California Statewide IPM Program); Tony Daley; Jack Hasenpusch; Greg Harold; Jiří and Marie Lochman (Lochman Transparencies); Iain Macaulay; Ron May; David Rentz; Pavel Svihra (courtesy University of California Statewide IPM Program); Paul Zborowski (Close-Up Photolibrary) and Alberto Venchi.

The staff from the Museo del Instituto de Zoologia Agricola, Maracay, Venezuela, in particular Luis J. Joly, Vilma Savini, Jose Clavijo and John Lattke, are sincerely acknowledged for their permanent support to HE.

We would like to acknowledge the world class Coleoptera collection at the Australian National Insect Collection (CSIRO), build up over last 60 years by Ev Britton, John Lawrence, Tom Weir, Murray Upton, Andrew Calder and other ANIC colleagues, volunteers and amateurs. That superb and well-curated collection made this project possible and feasible in the limited time that was available.

And finally, we would like to thank our families and loved ones (Ślipiński, Escalona, Garcia and Banks) for understanding our passion for beetles, and for their gracious support and patience during the years it took to produce this book.

Material and methods

MATERIAL STUDIED. Our research is based on the study of about 12,000 specimens of more than 1,250 described and undescribed species of Australian Cerambycidae during the past four years. In addition, specimens representing most of the genera, tribes and subfamilies of the Australo-Pacific region and Asia, as well as other regions of the world, were examined for comparative purposes. Almost all of the necessary types of Australian Cerambycidae species were examined and, usually, photographed (Appendix 3); this resulted in several new generic placements and synonymies.

The following abbreviations and acronyms are used for the names of the institutions where the specimens used in the project are held:

PHOTOGRAPHS. The photographs of the whole beetles and larger details were executed as layers on a BK Plus Lab System (Visionary Digital, USA) and subsequently combined using stacking software, Auto-Montage version 4.00 (Syncroscopy), Zerene Stacker (Zerene Systems LLC) or Helicon Focus (Helicon Soft). Some structural illustrations of adults and larvae were made from dissected specimens using a Leica MZ20 microscope with various attached digital cameras. Scanning Electron Microscope (SEM) images were generated with a Zeiss Evo LS 15 (CSIRO) from gold coated specimens, previously softened and dissected, cleaned in cold 10% KOH and rinsed in water and 95% ethanol.

MEASUREMENTS. In generic and species descriptions, several specimens of extreme size and variations were measured using a micrometer attached to a dissecting microscope, always including the largest and the smallest available ones. The number presented is the mean number, or range of these measurements.

In the Lamiinae keys and descriptions we have used standard measurements and indices. The length was measured dorsally along the midline from the most anterior point of the head to the apex of the elytra, and compared to the maximum width across the broadest part of the body, usually the elytra. The frontoclypeus was always measured in frontal view, from the middle of antennal tubercles to the clypeus apex (excluding membranous anteclypeus) and compared to the narrowest distance between eyes. The interocular index was calculated as the minimum distance between the inner eye orbits compared to the width of an eye at the same level. The pronotal length was measured from the middle of the anterior margin to the margin of basal foramen and compared to the maximum width of the pronotum. The length of the prosternum was measured as shortest distance between anterior margin of the prosternum and the procoxal cavity and compared to the length of the coxal cavity along the same line. The lengths of meso- and metaventrites were measured along the midlines.

TERMINOLOGY used for adult and larval beetle morphology follow most recent treatments of Coleoptera by Lawrence et al. (2011) and of Australian Beetles by Lawrence and Ślipiński (in press).

DISSECTIONS. In order to examine characters that might be used for generic diagnoses and for further phylogenetic analysis in the Australian Cerambycidae, at least two adults of one species (male and female) of each genus, were cleared in cold 10% KOH solution, disarticulated and placed in glycerol in containers or on slides for further examination and photographs; sclerotised but membranous and transparent tissues were stained with Chlorazol Black to generate contrast. After completing the examinations these specimens were transferred to standard vials with ethanol for permanent storage in the ANIC wet collection.

TAXONOMIC CONVENTIONS. A diagnosis and complete description is presented for each of the genera known to occur in Australia. The diagnosis is a summary of critical character states of the adult longhorn beetle that should be checked, without going into a lengthy description, once a specimen has been successfully keyed out in the generic key. Identified beetle should possess all or most of the mentioned character states. Under Remarks there is always a statement about genera that are very similar that could be confused with the genus in question, and also a list of the critical distinguishing features. Extensive use was made of the DELTA software (Dallwitz et al. 2000). The initial data matrix used in the generic descriptions was produced directly from the DELTA files using DELTA editor and was modified subsequently in a word processor.

HIGHER CLASSIFICATION OF LAMIINAE. Based on our preliminary molecular- and morphology-based research on Australian Cerambycidae we believe that the current tribal classification of Lamiinae is artificial and should be abandoned. It was devised by S. Breuning based on a combination of a few adult characters without any phylogenetic signal and has been perpetuated and ‘adjusted’ by cerambycid workers ever since. It is also not a practical system because many genera cannot be placed in recognised tribes, thus creating serious obstacles for further research. Certainly, some of the currently recognised groups are monophyletic, but that can be only established from the worldwide perspective through comprehensive phylogenetic research. Some of these lineages will probably be established in a comprehensive longhorn molecular phylogeny now in preparation (McKenna et al., in preparation).

Nevertheless, in this book we have listed the genera under their respective tribes in the classification section and have provided the tribal assignment under each genus but have not discussed or provided keys to the tribes. In the taxonomy section the genera are arranged in alphabetical order.

Figure 1. Longhorn beetles in their natural habitats

Figure 2. Longhorn beetles in their natural habitats

Figure 3. Longhorn beetles in their natural habitats

Figure 4. Longhorn beetles in their natural habitats

Figure 5. Longhorn beetles in their natural habitats

Figure 6. Longhorn beetles in their natural habitats

Figure 7. Longhorn beetles in their natural habitats

Figure 8. Longhorn beetles in their natural habitats

Family Cerambycidae

Latreille, 1802

Introduction

The Cerambycidae or longhorn beetles are among the most popular and easily recognised beetles and, for people who are more familiar with insects, mention of cerambycids invokes images of the largest and most iconic species, Titanus giganteus or Macrotoma heros (Fig. 31C,D) from the Amazon forests in South America. Long antennae and rather elongate bodies are used as general diagnostic characteristics for these beetles. However, longhorn beetles come in all sizes, shapes and colours, even commonly mimicking unpalatable beetles, stinging ants or wasps and thus making scientific definition of the family rather difficult.

Larvae of longhorn beetles are mostly internal feeders, developing in living or dead plant tissues. They are capable of extracting nutrient compounds and microelements from that low energy source, aided by various symbiotic microorganisms and cellulolytic enzymes. They often utilise damaged or dead trees for their development and, through feeding on rotten wood form an important element of the saproxylic fauna, speeding wood decay and energy circulation in woodland habitats. Many species are pests of quarantine concern, attacking and killing forest or ornamental trees, spreading injurious nematodes or, by developing in seasoned wood, causing structural damage to timber constructions.

Evolutionarily, Cerambycidae are members of the Phytophaga, a highly derived group of polyphagous beetles that feed primarily on vascular plants as larvae and adults. This clade comprises Chrysomeloidea (longhorn beetles, seed beetles and leaf beetles) and Curculionoidea (weevils) and includes over 124,000 described species (Ślipiński et al. 2011). The Phytophaga constitutes the largest known radiation of beetles, a radiation usually attributed to the the co-evolution of these phytophagous beetles and the rapidly radiating Angiosperm plants in the Tertiary (e.g. Farrell 1998). However, this attractive scenario has not been confirmed, at least for Chrysomeloidea (e.g. Gomez-Zurita et al. 2007), suggesting independent radiations of plants and phytophagous beetles driven by as yet unknown forces.

The division of Cerambycidae into subfamilies and tribes was gradually developed along with the first beetle classifications in 18th Century Europe and North America. It was quickly followed by a dedicated monographic study on Cerambycidae by Audinet-Serville, Thompson, LeConte, Pascoe, Lacordaire and Lameere, resulting in division of Cerambycidae into 7 major groups, usually treated as subfamilies: Disteniinae, Parandrinae, Spondylidinae, Prioninae, Aseminae, Lepturinae, Cerambycinae and Lamiinae. The advance of morphological research and use of novel larval characters in cerambycid classification provided support for many traditional groupings but highlighted some new divisions. There is now general agreement that the former Cerambycidae comprises four families, Oxypeltidae, Disteniidae, Vesperidae and Cerambycidae, of which only the largest family — Cerambycidae — occurs in Australia.

The Cerambycidae, in this sense, includes approximately 33,000 described species. A large number of new taxa are being added constantly from all over the world but particularly from Asia and South America. The family is cosmopolitan but the largest subfamilies — Prioninae, Cerambycinae and Lamiinae — are most diverse in the tropical and subtropical parts of the world.

Subfamily level classification of Cerambycidae has been very unstable and proposed relationships between recognised subfamilies are mostly based on larval characters alone. Following Švácha and Lawrence (in press) we recognise eight subfamilies in Cerambycidae: Prioninae, Parandrinae, Dorcasominae, Cerambycinae, Spondylidinae, Necydalinae, Lepturinae, and Lamiinae. The Dorcasominae, Necydalinae and Lepturinae have restricted geographical distributions and do not occur in Australia, and the Spondylidinae are represented here only by two introduced species of Arhopalus.

Cerambycidae are one of the largest families of Australian beetles, with over 300 described genera and 1,400 species. Yet despite their economic importance and obvious appeal to professional and amateur entomologists, they have been neglected for a long time. Published work on the Australian Cerambycidae has been very fragmentary and mostly limited to isolated species or generic descriptions, often without illustrations or a reference to the tremendous taxonomic difficulties within the generic and tribal classifications of most of subfamilies. The number of genera and species to be treated and the problems in higher classification have been the main impediments to research on this group in Australia. While actively participating in the international team working on higher classification of Cerambycidae that will take many years to publish, we decided to produce an overview and identification keys to Australian genera to facilitate further research on this group.

This is the first in a series of three volumes dedicated to the genera of the Australian Cerambycidae. This book contains a general introduction to the Australian longhorn beetles, keys to identification of subfamilies based on adults and larvae and treatment of the family Lamiinae, including 74 genera and 536 described species. Volume two of this series will cover the genera of Cerambycinae, and the last volume will be devoted to the genera and species of Parandrinae, Spondylidinae and Prioninae.

Morphology of adult beetles

The Longhorn Beetles are a very large and diverse group that are of immense popularity with members of the taxonomic community. Yet rarely have they been studied morphologically in any depth. The work of Napp (1994) is a notable exception: in her cladistic analyses she compared morphologies of critical cerambycoid taxa. More recently, Švácha and Lawrence (in press) have treated the morphology of Cerambycidae in their entirety for the first time, in a chapter in the Handbook of Zoology. Lawrence et al. (2011) and Lawrence and Ślipiński (in press) provide general accounts of beetle morphology and the terminology employed in these books is followed here. The descriptions below are based solely on representatives in the Australian fauna.

Body

The body of the longhorn beetle is usually elongate and parallel-sided, dorsally convex, rarely broadly oval (e.g. Phaolus) or flattened. Most of the Australian species are medium-sized beetles with body length between 10 and 25 mm. The smallest Australian species are found among species of Lamiinae (e.g. Somatidia, Microlamia), measuring about 2 mm, while males of the prionine Xixuthrus microcerus (White) reach above 90 mm in length. The dorsal surfaces of Cerambycidae may be polished and apparently glabrous as in Parandrinae and some Prioninae, but are usually clothed with a single or double vestiture of hairs, bristles, thickened and adpressed setae or combinations of these. In some Prioninae (e.g. Sclerocantha) both venter and part of dorsum are covered by long tomentose pubescence, sometimes completely obscuring the integument. Body colour varies from black or brown in Spondylidinae, Parandrinae and Prioninae, with notable exceptions in Rhipidocerus (yellowish-green) and Phaolus (metallic blue or green), to brightly bi- or multicoloured in Cerambycinae and Lamiinae.

Head

The head capsule in Parandrinae, Spondylidinae and most Prioninae and Cerambycinae is prognathous or weakly declined with the mouthparts directed antero-ventrally (Fig. 11A,C,F) and clearly visible from above. However, in most Lamiinae (except for Enicodes or Tmesisternini) the frontal region of head capsule is strongly deflexed from behind the eyes, causing the mouthparts to be directed ventrally or even posteriorly and often not visible from above. The head is usually transverse and slightly narrower than the prothorax, and in some Lamiinae (e.g. many Rhytiphora) in repose is received in a concave area of the prosternum in front of strongly developed procoxae. The head is rarely strongly transverse (e.g. Enicodes), elongate or rostrate, with the frontoclypeal region forming a distinct muzzle in some flower-visiting Cerambycinae (e.g. Rhinophthalmus). The median longitudinal groove (Fig. 10C) marking the internal endocarina on the frontoclypeal region is usually well developed (absent in Parandrinae) and often continues into the occipital region in some Prioninae and all Lamiinae.

The frontoclypeal suture is often distinctly impressed and arcuate, rarely absent or strongly angulate (e.g. Arhopalus). Anterior tentorial pits are usually visible externally near dorsal mandibular articulations. The clypeus is often short and subdivided into the membranous anteclypeus (also called clypeolabral membrane) and the sclerotised postclypeus (clypeus) (Fig. 10C). In all Parandrinae, some Prioninae and some genera of Lamiinae (e.g. Apomecyna) the clypeus is entirely sclerotised and prominent anteriorly (Fig. 166B). Several genera of Cerambycinae (e.g. Stenoderus, Syllitus, Tricheops) have gland openings and variously developed dispensers of odoriferous substances, located laterally near the mandibular articulations (Fig. 12B).

The eyes of cerambycids provide a number of characters used for defining genera. In most species, the eyes are large and variously emarginate around antennal tubercles (Fig. 12A) with their facets convex and of moderate size. The eyes are reduced to a small number of coarse facets or completely divided into upper and lower lobes (Fig. 170C) in some Lamiinae (e.g. Microlamia) and Cerambycinae (e.g. Skeletodes, Nenenia). In Parandrinae, the eyes are often smaller and weakly emarginate (Fig. 11A) while in many Cerambycinae and in some Prioninae they are very large and extend well onto the ventral side (Figs 11F, 12A). Tricheops has the lower eye lobe deeply emarginate and surrounding a gland opening and dispenser (Fig. 12A).

Antennal insertions are almost always dorsal and supported by raised tubercles (Figs 12B, 45B). The antennal foramina (cavities) are clearly visible from above and distant from the mandibular articulations in most Lamiinae and Cerambycinae (Figs 12B, 45B) but are situated more laterally and closer to the mandibular articulations in Spondylidinae (Fig. 11C) and most Prioninae (Fig. 11F). In Parandrinae, the antennal insertions are distinctly lateral and the foramina are more or less concealed from above by bulging frontal projections (Fig. 11A). Transverse occipital carinae and subantennal grooves are always absent from Cerambycidae.

The labrum is usually clearly visible, free and transverse or about as long as wide; only rarely is it distinctly longer than wide. In some Prioninae, the labrum is heavily sclerotised and adpressed to the clypeus, and in Parandrinae it is completely fused to the clypeus and not visible externally (Fig. 11B).

The antennae are long, usually distinctly longer in males than in females, almost always 11-segmented (12-segmented in males of the prionine Phaolus, cerambycine Distichocera and in both sexes of the lamiine genus Phelipara). Antennae are usually filiform or serrate but sometimes pectinate or flabellate (e.g. in males of Rhipidocerus, Enneaphyllus), bi-flabellate (in the cerambycine tribe Distichocerini, Fig. 34E) or, exceptionally, capitate (e.g. in the cerambycine Telocera, Fig. 36B). Antennomeres are apparently glabrous or sparsely setose in Parandrinae and some Prioninae but usually hairy or setose in the remaining groups; and exhibit extreme development of the setal fringe in some species of Rhytiphora (Lamiinae). Antennal sensilla are spread over surfaces of the antennomeres but in Parandrinae appear to be concentrated in concave areas along the external edge of antennomeres (Fig. 14A). The antennal scape is longer than wide (subquadrate in Parandrinae), sometimes very long and reaching beyond the base of the pronotum (e.g. in the lamiines Phelipara, Acanista, Olenecamptus, and the cerambycines Macrones, Psilomorpha). In many Lamiinae the apex of the scape is provided with a sensory area surrounded by the apical carina (= cicatrix). The surface of the scape is asperate or rugose (e.g. Olenecamptus) or even tuberculate (some species of Microlamia). The pedicel is almost always very short and transverse; only in Arhopalus (Spondylidinae) and a few members of other genera it is distinctly longer than wide (Fig. 11D).

The mandibles are symmetrical, triangular and usually unidentate apically with one or more teeth on incisor edges. In many exotic Prioninae the mandibles are strongly dimorphic and in males they can be used to combat competing males. In Australia, the males of some Prioninae have moderately large and/or elongate mandibles but in males of Storeyandra frenchi (Parandrinae) the head and mandibles are particularly strongly developed in comparison to females (Figs 11B, 33E). In a peculiar genus of Cerambycinae — Australodon, the mandibles are very large and strongly bent downwards in the male (Fig. 12F). The mandible at base is usually broad and often forms a molar part that is less differentiated, microspinulose and apparently not homologous with the true mola (Švácha & Lawrence, in press, but see Napp 1994). Also the fringe of hairs along the desclerotised inner margin of the mandible is treated as not homologous with the prostheca of other Polyphaga (Švácha & Lawrence, in press, but see Napp 1994).

Figure 9. Adult morphology

Abbreviations

Figure 10. Adult morphology

[Anne Hastings]

Abbreviations

Maxillae are usually well developed in Cerambycidae (Fig. 10F) and bear a triangular, transverse cardo, short stipes and, almost always, apically setose galea and lacinia. Both maxillary lobes can variously be modified, especially in pollen- or nectar-feeding Cerambycinae. The lacinia is strongly reduced in most Prioninae and entirely absent in Parandrinae. Maxillary palps are usually long, 4-segmented, with the apical palpomere fusiform and pointed apically in most Lamiinae to triangular or securiform in other subfamilies.

In Cerambycidae, the labium (Fig. 10G) is clearly visible and exposed between short, ventral genal projections. The mentum is usually short and transverse and distinctly articulated with, at most, a weakly prominent submentum anteriorly. The ligula is often well developed and membranous or sclerotised (e.g. Parandrinae), usually emarginate or bilobed. Labial palps are 3-segmented, broadly separated at the bases with the apical palpomere fusiform to variously expanding apically. The shape of terminal maxillary and labial palpomeres is sexually dimorphic in some Lamiinae (e.g. Athemistus), with males having the segment triangular or securiform (Fig. 59B).

Gular sutures are usually short and broadly separated, providing support for the internal tentorial arms attached anteriorly to the frontoclypeal region and visible externally as tentorial pits. The transverse tentorial bridge (corpotentorium) is usually well developed and broad.

Prothorax

The prothorax is very variable in shape but usually transverse or subquadrate, rarely longer than wide (e.g. the lamiine Gnoma, Fig. 88D) with sides often slightly rounded or bearing a single lateral tooth or spine at or near middle. The pronotum is often constricted before its base and usually narrower than the basal elytral width.

A lateral pronotal carina separating notum from prothoracic hypomera is absent (Fig. 11C) in Cerambycinae, Lamiinae and Spondylidinae. The carina is complete and simple in Parandrinae (Fig. 11A) and variable and often serrate or dentate in most Prioninae (Fig. 11F). An apparently secondary lateral pronotal carina appears in many members of the Lamiine tribe Tmesisternini (mostly Papuan) and is located dorsad to its regular position (indicated by a placement of the lateral tubercle, Fig. 150A). The anterior pronotal angles are almost always blunt or rounded and the posterior angles are broadly rounded to right and rarely projecting posteriorly.

The pronotal disc is very variable and variously sculptured in most taxa, in most cases bearing characters commonly used in the generic classification. The surface is often convex and punctured or rugose, rarely smooth, and occasionally ornamented with impressions, grooves, or paired and/or unpaired nodules or tubercles. Depending on the shape of the prothorax, its venter (prosternum) in front of the procoxae can be very long or very short (e.g. Figs 12E, 71G) with its surface usually flat or weakly convex but in many Lamiinae it is strongly elevated and concave receiving the ventral side of the head in repose (Fig. 68E).

Figure 11. Adult morphology

The prosternal process between procoxae is always complete and extremely variable, also providing useful characters for the generic classification. It is usually flat but often arcuate or angulate and abruptly bent beyond the procoxae. In some Lamiinae, the prosternal process extends well beyond the procoxae (Fig. 156E) and articulates with a projection or a pit on the mesoventrite or is received in a deep fossa (e.g. Trigonoptera). Some genera of Cerambycinae and Lamiinae (e.g. Sybra) have an accessory mesal articulation developed as a knob on the prosternal process that fits into a procoxal fossa (Fig. 181B). The notosternal sutures are usually short and incomplete anteriorly but complete in some Prioninae.

The procoxae are usually flat and weakly to strongly transverse in Parandrinae, Prioninae and Spondylidinae (Fig. 11A,C,E) but are usually oval and project well below the prosternum in many Cerambycinae and Lamiinae. The protrochantin (Figs 11C, 164G) is exposed in all Parandrinae, Prioninae and Spondylidinae, but is sometimes concealed in Lamiinae (e.g. Stenellipsis, Fig. 178G) and Cerambycinae (e.g. Skeletodes). The procoxal cavities are variable but usually transverse and narrowly separated medially, with lateral extensions exposing the trochantins. The prosternal process is often expanded beyond the coxae and forms a narrow external closure by meeting narrow postcoxal projections in most Lamiinae (except Batocerini) but it does not form this external closure in most Prioninae and Cerambycinae. The internal closure (Fig. 45G) is usually broad and complete but it is absent in Parandrinae and Prioninae.

Pterothorax

The scutellar shield is almost always visible and not abruptly elevated (not visible in some wingless Lamiinae), anteriorly simple or separated from the mesoscutum by a transverse impression (Fig. 175G); in some genera of Tmesisternini (e.g Tmesisternus) the scutellum is large, projecting and anteriorly overlapping the posterior edge of pronotum (Fig. 155C). A stridulatory file is usually present on the mesoscutum, but always absent in Parandrinae and Prioninae; it is divided by a median mesoscutal carina in Arhopalus (Spondylidinae) and is nearly always complete in Cerambycinae and Lamiinae (strongly reduced or absent in some genera, e.g. Rhytiphora).

The elytra are very variable in Cerambycidae but are usually much longer than wide and longer than the pronotum, often covering the entire abdomen, and are apically rounded, spinose or truncate. Substantial variation in elytra occurs in wasp-mimicking Cerambycinae with their elytra often short and truncate (e.g. Hesthesis) or apically narrowed (e.g. Enchoptera, Macrones), exposing several abdominal tergites (Figs 34A, 35A,B). The dorsal surfaces of the elytra are often polished and variably punctate with a leathery appearance in most Prioninae and Parandrinae, to coarsely, and sometimes serially punctate in various Lamiinae and Cerambycinae. Elytral intervals sometimes form elongate tubercles, ridges, costae or a reticulate network. The elytral epipleuron is usually vertical or oblique and incomplete apically (the flat, complete epipleura in Phyxium are a notable exception, Fig. 113E). The epipleura are deeply emarginate at the level of the humeral tubercle, in flight exposing the wing bases in Tragocerus (Fig. 37G) and related taxa of Tragocerini that fly without opening the elytra.

The mesoventrite is large and almost always separated