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 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.

exotic and iconic speciesexotic and iconic speciesexotic and iconic species

Exotic and iconic Cerambycidae

Shape variation in Cerambycidae

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 (Ślipiíski 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 1,300 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.

The vast world literature on cerambycid biology has been reviewed by Duffy (1953, 1963), Linsley (1959, 1961) and Švacha and Lawrence (in press). The information published so far is heavily biased towards a few economically important pest species and/or to generally better known species from the Palaearctic and Nearctic Regions.

Various aspects of biology of the Australian species were described in numerous early papers, mostly by D. Best (1881–1920), A.R. Brimblecombe (1943–1956) and W.W. Froggatt (1893–1930). These were summarised by Duffy (1963) who has described numerous larvae and provided new biological insight based on larval collecting data and unpublished information from his collaborators.

exotic and iconic speciesexotic and iconic species

Rhytiphora nigrovirens

Ancita sp.

Later contributions to the knowledge of biology of the Australian Cerambycidae include many papers on pollination and host plants by G.A. Webb (1985–1997) and T.J. Hawkeswood and their collaborators (1985–2011). Food preferences Cerambycidae are phytophagous beetles.

Their larvae, called round-headed borers, develop in tissues of woody or herbaceous plants in conditions ranging from healthy and alive to dead and decomposing due to fungi (Duffy 1953). In addition to the standard set of digestive enzymes, the gut of cerambycid larvae contains cellulotic enzymes and yeast-like symbionts (in the mid gut) help to extract sugars and critical particles like nitrogen from the wood material (Švacha & Lawrence, in press). Some Australian cerambycids apparently are specialists confined to a single or a few plant species suitable for larval development, but many show amazingly broad host ranges that include native and introduced gymnosperms and angiosperms (Duffy 1963; Hawkeswood & Dauber 1991).

Hanks (1999) classified cerambycids into four categories based on the condition of the larval host plant at the time of colonisation. The species that attack healthy and vigorous plants are able to develop in stems of herbaceous plants and in twigs of woody plants, often inducing gall-like structures on the host plants.

All serious pest species classified in Lamiinae (e.g. Anaplophora, Monochamus) belong to this first category. In addition, some Lamiinae girdle branches of their hosts before oviposition (Stride & Warwick 1962), or their larvae are able to girdle the branches or roots internally during feeding (Linsley 1961).

Many Lamiinae and Cerambycinae colonise the host that has been attacked previously by bark borers or has been weakened by poor growing conditions, fire or flood. These species often are able to complete several generations on an individual tree and only exceptionally kill the host. The species in the third category, are able to colonise woody plants that are severely stressed and dying because of intense fire, drought or attack by bark beetles or nematodes; these species are able to complete only one generation before killing the host. The dead wooden hosts, often also infested by fungi, are then available for many generations of cerambycid species of the fourth category that develop in dead hosts until they decompose and are no longer suitable for their larvae.

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photo of the authors

Adam Ślipiński did 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 and held a joint appointment as the professor of biology at the University of Zielona Gora teaching entomology and environmental biology. He has been a research scientist at the Australian National Insect Collection for the last 13 years. He is the author of over 160 research publications, editor of a two-volume book on the phylogeny and classification of beetles and an author of a book on Australian ladybird beetles. Adam’s research concentrates on the phylogeny and higher classification of beetles.
Contact: Adam.Slipinski@csiro.au

Hermes Escalona earned his PhD in Entomology from the Universidad Central de Venezuela in 2012. He is interested in systematics, evolution, and historical biogeography of Coleoptera, with a current focus on Longhorns beetles (Cerambycidae) and small beetle families within Cucujiformia. He is currently affiliated with the Museo del Instituto de Zoología Agrícola-UCV and is a visiting scientist at the Australian National Insect Collection working with Adam Ślipiński on the Australian Cerambycidae.
Contact: Hermes.Escalona@gmail.com

Anne Hastings has a degree in Fine Arts from the Tasmanian School of Art in Hobart and is currently working as an Entomological Artist in the Australian National Insect Collection in CSIRO. In 2005 she gained a Graduate Diploma in Information Technology from the University of Canberra and is currently working with Adam and Hermes on the Australian Cerambycidae project.
Contact: Anne.Hastings@csiro.au

Cate Lemann has a Science Degree from the University of New England, Armidale and a Biological Technicians Certificate. She is working as a research projects officer in the CSIRO Australian National Insect Collection supporting both the Australian Cerambycidae project with specimen preparation and imaging and more undertaking more general collection management activities.
Contact: Cate.Lemann@csiro.au

Mengjie Jin registered as a PhD student in 2016, from Sun Yat-sen University, in Guangzhou, China. She is currently involved in a postgraduate student project on Australian Cerambycidae supported by CSIRO and the Australian Department of Agriculture. She is working under the supervision of Adam Ślipiński. At the moment, she is researching the taxonomy and phylogeny of the subfamilies Cerambycinae and Prioninae.
Contact: jinmengj@mail2.sysu.edu.cn

Lamiinae is the biggest subfamily of Cerambycidae with about 20,000 species classified in hundreds of genera worldwide. Seventy-four genera and about 550 described species are know from Australia but the species-level taxonomy remains a highly challenging project for future research.

Head

1. The frontal region strongly deflexed from behind the eyes or vertical. This is the case in most taxa but there are exceptions in some of the Tmesisternini tribe.

Deflexed (common)

Vertical (common)

Obliquely forward (rare)

2. With complete median longitudinal groove marking deep internal endocarina, which continues into occipital area.

Groove marking internal endocarina

3. Eyes almost always deeply emarginate or divided into upper and lower parts, finely to very coarsely facetted.

Deeply emarginate

Divided into upper and lower parts

4. Antennal insertions always exposed and supported by raised tubercles.

Raised tubercles and exposed insertions

5. Frontoclypeal suture distinctly impressed. Labrum visible and free.

Visiible labrum and impressed suture

6. Maxilla always with distinct, setose galea and lacinia; apical maxillary palpomere fusiform (tapering at both ends; spindle-shaped) but triangular in males of some genera.

Palpomere 4 fusiform (broad in middle, narrowing at ends), distinct setose (hairy) galea and lacinia

Thorax

1. Lateral pronotal carinae always absent, with secondary development in some Tmesisternini.

Lateral pronotal carinae always absent

2. Prosternal process complete.

Complete prosternal process

3. procoxal cavities almost always closed externally. Procoxae often projecting well below prosternum almost always with lateral extension and exposed trochantin.

Exposed trochantin

4. Scutellum usually visible; mesoscutum often with stridulatory file or rarely absent (e.g. Rhytiphora). 

Visible scutellum

5. Mesocoxal cavities usually laterally open.

Open mesocoxal cavities

Legs

1. Tibial spurs usually 2-2-2, well developed, rarely reduced to 1 or 0 on some pairs of legs. Protibia with oblique band of dense setae usually within pubescent groove (antennal cleaner) on inner face, mesotibia with similar structure on outer face.

Mesotibia with structure on outer face

Ovipositor

1. usually long and flexible with short subapical styli.

Long ovipositor

Cerambycinae

Australia has been often cited as the only continent where the species of Cerambycinae outnumber the species of Lamiinae. Australian Cerambycinae include quite a variety of diurnal (day active) and often brightly coloured species with contrasting hues or brilliantly iridescent elytra.

Head

1. prognathous to moderately inclined head with frontal region rarely deflexed from behind eyes.

Head prognathous, example 1

Head prognathous, example 2

Head slightly inclined

Head moderately inclined

2. Frontoclypeus usually subquadrate, occasionally distinctly longer than wide, forming rostrum

Frontoclypeus distinctly longer than wide

3. median longitudinal groove and associated internal endocarina usually present but incomplete posteriorly. Glandular opening near mandibular bases and tongue-like dispenser present in few genera only

Tongue-like dispenser for glandular opening

4. Postocular constriction absent or weak, forming distinct neck in a few ant-like genera.

Distinct neck in ant-like genera

5. Frontoclypeal suture impressed or not, straight, arcuate or angulate.

6. Labrum visible externally, usually transverse.

7. Maxilla with distinct, setose galea and lacinia; apical labial and maxillary palpomeres fusiform to strongly triangular; ligula almost always membranous and expanded laterally.

Distinct, setose galea and lacinia

8. Eyes variable in size, mostly emarginate, forming larger lower and smaller upper lobes, rarely oval or divided into separate upper and lower parts; finely to very coarsely facetted.

image of state onediagram of state oneimage of state twodiagram of state two

Mostly emarginate, forming larger lower and smaller upper lobes

Rarely divided into separate upper and lower parts

9. Antenna usually 11-segmented, rarely 12-segmented, filiform or serrate, sometimes dilated toward apex, modified flabellate, biflabellate or clavate. Scape always distinctly longer than pedicel; pedicel short, usually transverse, shorter than antennomere 3.

image of state oneimage of state two

Antennae sometimes dilated toward apex

10. Antennal insertions have two major types of articulation: (a) insertion prominent supported by raised tubercle, the rim is thin, usually directed laterally with scape shallowly received so that the articulation point is on the rim; (b) insertion depressed, usually flat, often located in a shallow depression, the rim is thicker, reflexed with base of scape deeply received so that the point of articulation is located deep inside the insertion.

image of state oneimage of state twoimage of state threeimage of state four

Insertion type a: thin rim and raised tubercle

Insertion type a: prominent articulation point

image of state fiveimage of state siximage of state sevenimage of state eight

Insertion type b: thick rim

Insertion type b: deep articulation point

Thorax

1. Prosternal process variable, ranging from broad to absent

image of state onediagram of state oneimage of state twodiagram of state two

Broad prosternal process

Prosternal process absent

2. Procoxal cavities round or weakly transverse often with lateral extensions and exposed protrochantin, closed internally, closed or open externally.

image of state

Procoxal cavities round, closed internally and externally

Procoxal cavities weakly transverse with lateral extensions, exposed trochantin, open externally

3. Scutellum always visible; mesoscutum rarely with median endocarina, usually with undivided stridulatory file.

Scutellum removed from a specimen

4. Mesocoxal cavities open or closed laterally to mesepimeron; mesotrochantin visible or not. Mesocoxae flat or rarely projecting; in males of Syllitosimilis with inner spinose projections.

Mesocoxal cavities can be seen through the mesocoxa

5. Elytra usually completely covering abdomen but sometimes shortened or narrowing apically and dehiscent.

6. Wings always present, always without wedge cell and with 3 or 4 free veins in median field.

Free veins in median field of wing

wedge cell example

Key to exotic Cerambycidae

couplet 1 of 20

Fig. 1a

Fig. 1aa

Fig. 1b

Fig. 1bb

Large, 30-50 mm long brown beetle; elytra with complex net-like pattern of whitish veins (Figs. 1a, 1aa); pronotum bearing sharp middle projection and dense pubescence (Figs. 1b, 1bb) . . . . . . . . . . . .

[subfamily Prioninae, the Huhu Beetle; New Zealand]
Prionoplus reticularis White

Fig. 1c

Fig. 1cc

- Elytra without a net-like pattern (Figs. 1c, 1cc); size variable.

. . . . . . . . . . . . Go to couplet 2 →

couplet 2 of 20 (from 1)

Fig. 2a

Fig. 2aa

Fig. 2b

Fig. 2bb

Protibia with grooved antennal cleaner on inner surface (Figs. 2a, 2aa); mesotibia with sulcate or comb-like antennal cleaner on outer surface (Figs. 2a, 2aa); head strongly deflexed (Figs. 2b, 2bb).
[subfamily Lamiinae]

. . . . . . . . . . . . Go to couplet 3 →

Fig. 2c

Fig. 2cc

Fig. 2d

Fig. 2dd

- Protibia and mesotibia without antenna cleaners (Figs. 2c, 2cc); head prognathous or weakly deflexed (Figs. 2d, 2dd).
[subfamilies Cerambycinae and Spondylidinae]

. . . . . . . . . . . . Go to couplet 7 →

couplet 3 of 20 (from 2)

SUBFAMILY LAMINAE

Fig. 3a

Fig. 3aa

Fig. 3b

Fig. 3bb

Elytra bluish or black, mostly glabrous with white setae forming small isolated spots (Figs. 3a, 3aa); procoxal cavities closed posteriorly(Figs 3b, 3bb).
[genus Anoplophora]

. . . . . . . . . . . . Go to couplet 4 →

Fig. 3c

Fig. 3cc

Fig. 3d

Fig. 3dd

- Elytra entirely setose and of various colours and combinations (Figs. 3c, 3cc); procoxal cavities at least narrowly open posteriorly (Figs. 3d, 3dd).

. . . . . . . . . . . . Go to couplet 5 →

couplet 4 of 20 (from 3)

Fig. 4a

Fig. 4aa

Fig. 4b

Fig. 4bb

Pronotal disc flat (Figs. 4a, 4aa); elytra glossy black, finely punctate, smooth at basal part (Figs. 4b, 4bb) . . . . . . . . . . . .

[Asian longhorn beetle; USA and Canada, Europe and Northern Asia]
Anoplophora glabripennis (Motschulsky)

Fig. 4c

Fig. 4cc

Fig. 4d

Fig. 4dd

- Pronotal disc with dorsal nodules (Figs. 4c, 4cc); basal part of elytra with coarse granulation and nodules (Figs. 4d, 4dd) . . . . . . . . . . . .

[Citrus longhorn beetle;
Europe, Northern Asia, South and South-East Asia]
Anoplophora chinensis (Forster)

couplet 5 of 20 (from 3)

Fig. 5a

Fig. 5aa

Fig. 5b

Fig. 5bb

Fig. 5c

Fig. 5cc

Antennal scape with apical carina (Figs. 5a, 5aa); apices of elytra rounded or weakly angulate (Fig. 5b, 5bb); antennomere 3 very long, at least twice as long as antennal scape (Figs. 5c, 5cc).
[genus Monochamus]

. . . . . . . . . . . . Go to couplet 6 →

Fig. 5d

Fig. 5dd

Fig. 5e

Fig. 5ee

Fig. 5f

Fig. 5ff

- Antennal scape without apical carina (Figs. 5d, 5dd); apices of elytra bispinose (Figs. 5e, 5ee); antennomere 3 moderately long, about 1.5 times as antennal scape (Figs. 5f, 5ff) . . . . . . . . . . . .

[genus Apriona, Brown mulberry longhorn beetle;
Myanmar, China, India, Japan, Korea, Pakistan and Vietnam]
Apriona germari Hope

couplet 6 of 20 (from 5)

Fig. 6a

Fig. 6aa

Fig. 6b

Fig. 6bb

Pronotum with two longitudinal orange stripes (Figs. 6a, 6aa); each elytron with 5 longitudinal bands of black and grey rectangular spots (6b, 6bb) . . . . . . . . . . . .

[Pine Sawyer beetle;
Europe and Northern Asia, South and South-East Asia]
Monochamus alternatus Hope

Fig. 6c

Fig. 6cc

Fig. 6d

Fig. 6dd

- Pronotum (Figs. 6c, 6cc) and elytra (Figs. 6d, 6dd) black or with other colour pattern . . . . . . . . . . . .

[Holarctic and Oriental Regions]
Monochamus spp.

couplet 7 of 20 (from 2)

SUBFAMILIES CERAMBYCINAE AND SPONDYLIDINAE

Fig. 7a

Fig. 7aa

Fig. 7b

Fig. 7bb

Elytra very short, not extending over abdomen (Figs. 7a, 7aa); femora strongly clavate (club-like) (Figs. 7b, 7bb) . . . . . . . . . . . .

[Spruce Shortwing Beetle; Europe and Asia extending to China]
Molorchus minor (Linnaeus)

Fig. 7c

Fig. 7cc

Fig. 7d

Fig. 7dd

- Elytra either completely covering abdomen (Figs. 7c, 7cc) or with only the terminal segment exposed (Figs. 7d, 7dd).

. . . . . . . . . . . . Go to couplet 8 →

couplet 8 of 20 (from 7)

Fig. 8a

Fig. 8aa

Fig. 8b

Fig. 8bb

Antennomere 3 very long, about 1.5 times of antennal scape, reaching beyond the posterior margin of pronotum (Figs. 8a, 8aa); male with large patch of dense yellowish setae on prothoracic hypomeron (Figs. 8b, 8bb).
[genus Stromatium]

. . . . . . . . . . . . Go to couplet 9 →

Fig. 8c

Fig. 8cc

- Antennomere 3 at most slightly longer than scape and never reaching posterior margin of pronotum (Figs. 8c, 8cc).

. . . . . . . . . . . . Go to couplet 10 →

couplet 9 of 20 (from 8)

GENUS Stromatium

Fig. 9a

Fig. 9aa

Elytra densely and coarsely punctate (Figs. 9a, 9aa); elytral setae sparse, partially erect and not covering rather shiny body surface (Figs. 9a, 9aa); each elytron with at least two costae (ridges) (Figs. 9a, 9aa) . . . . . . . . . . . .

[Teak Trunk Borer; worldwide]
S. barbatum (Fabricius)

Fig. 9b

Fig. 9bb

- Elytral surfaces dull covered by dense and adpressed pubescence almost completely obscuring the body surface but leaving some sparse shiny granules (Figs. 9b, 9bb); elytral costae (ridges) absent or very weak (Figs. 9b, 9bb) . . . . . . . . . . . .

[Oriental Region, native to Christmas Island]
S. longicorne (Newman)

couplet 10 of 20 (from 8)

Fig. 10a

Fig. 10aa

Eye deeply emarginate, two lobes only connected by 1 row of ommatidia (Figs. 10a, 10aa). [genus Tetropium]

. . . . . . . . . . . . Go to couplet 11 →

Fig. 10b

Fig. 10bb

- Eye with two lobes connected by at least several rows of ommatidia (Figs. 10b, 10bb).

. . . . . . . . . . . . Go to couplet 12 →

couplet 11 of 20 (from 10)

Fig. 11a

Fig. 11aa

Fig. 11b

Fig. 11bb

Pronotum densely punctate, without any glabrous raised area (Figs. 11a, 11aa); elytra reddish or yellowish brown (Figs. 11b, 11bb) . . . . . . . . . . . .

[Brown spruce longhorn beetle;
Europe and Asia, now distributed in Northern Asia, USA and Canada]
Tetropium fuscum (Fabricius)

Fig. 11c

Fig. 11cc

Fig. 11d

Fig. 11dd

- Pronotum weakly punctate, with 3 weak shiny, raised nodules (Figs. 11c, 11cc); elytra uniformly dark brown (Figs. 11d, 11dd) . . . . . . . . . . . .

[Black Spruce Beetle;
Europe and Northern Asia, Mediterranean Basin, USA and Canada]
Tetropium castaneum (Linnaeus)

couplet 12 of 20 (from 10)

Fig. 12a

Fig. 12aa

Pronotum lateral edge with sharp median projection (Figs. 12a, 12aa) . . . . . . . . . . . .

[Two-toothed Longhorn; New Zealand]
Ambeodontus tristis (Fabricius)

Fig. 12b

Fig. 12bb

- Pronotum with lateral edge rounded (Figs. 12b, 12bb).

. . . . . . . . . . . . Go to couplet 13 →

couplet 13 of 20 (from 12)

Fig. 13a

Fig. 13aa

Fig. 13b

Fig. 13bb

Prosternal process very broad, at least as wide as procoxa (Figs. 13a, 13aa); pronotum covered with dense erect pubescence, with two glabrous nodules on pronotal disk (Figs. 13b, 13bb) . . . . . . . . . . . .

[European house borer; cosmopolitan, established in WA]
Hylotrupes bajulus (Linnaeus)

Fig. 13c

Fig. 13cc

Fig. 13d

Fig. 13dd

- Prosternal process distinctly narrower than width of procoxa (Figs. 13c, 13cc); pronotum not covered with dense erect pubescence, without two glabrous nodules on pronotal disk (Figs. 13d, 13dd).

. . . . . . . . . . . . Go to couplet 14 →

couplet 14 of 20 (from 13)

Fig. 14a

Fig. 14aa

Fig. 14b

Fig. 14bb

Procoxal cavities oval, without lateral projections, not exposing protrochantin (Figs. 14a, 14aa); elytra with coloured bands formed by short adpressed setae (Figs. 14b, 14bb).

. . . . . . . . . . . . Go to couplet 15 →

Fig. 14c

Fig. 14cc

Fig. 14d

Fig. 14dd

Fig. 14e

Fig. 14f

Fig. 14ff

- Procoxal cavities with lateral projections, at least partially exposed protrochantin (Figs. 14c, 14cc, 14d, 14dd); elytra either uniformly coloured (Figs. 14e) or with orange bands (not formed by adpressed setae) (Figs. 14f, 14ff).

. . . . . . . . . . . . Go to couplet 16 →

couplet 15 of 20 (from 14)

Fig. 15a

Fig. 15aa

Fig. 15b

Fig. 15bb

Frons with triangular keel (Figs. 15a, 15aa); pronotum brown with sparse yellow recumbent setae; elytra black with yellow markings (Figs. 15b, 15bb) . . . . . . . . . . . .

[Clytine longhorn beetle; South and South-East Asia]
Xylotrechus magnicollis (Fairmaire)

Fig. 15c

Fig. 15cc

- Frons without triangular keel; body black, with white markings on elytra (Figs. 15c, 15cc) . . . . . . . . . . . .

[Kokeshi longicorn beetle; South and South-East Asia]
Chlorophorus diadema (Motschulsky)

couplet 16 of 20 (from 14)

Fig. 16a

Fig. 16aa

Fig. 16b

Fig. 16bb

Eye shallowly emarginate (Figs. 16a, 16aa); Protibial spur single (Figs. 16b, 16bb). [genus Arhophalus]

. . . . . . . . . . . . Go to couplet 17 →

Fig. 16c

Fig. 16cc

Fig. 16d

Fig. 16dd

- Eye distinctly emarginate (Figs. 16c, 15cc); Protibial spurs paired (Figs. 16d, 16dd).

. . . . . . . . . . . . Go to couplet 19 →

couplet 17 of 20 (from 16)

Fig. 17a

Fig. 17aa

Articulation of tarsomere 4 with tarsomere 3 (or point of division into two lobes of tarsomere 3) is about halfway along the total length of tarsomere 3 (Figs. 17a, 17aa) . . . . . . . . . . . .

[Burnt pine longhorn beetle;
Europe, North Africa, established in New Zealand]
Arhophalus ferus (Mulsant)

Fig. 17b

Fig. 17bb

- Articulation of tarsomere 4 with tarsomere 3 (or point of division into two lobes of tarsomere 3) is fairly close to the base of tarsomere 3 (Figs. 17b, 17bb).

. . . . . . . . . . . . Go to couplet 18 →

couplet 18 of 20 (from 17)

Fig. 18a

Fig. 18aa

Terminal segment of maxillary palp strongly securifrom, with length 1-1.26 times its apical width (Figs. 18a, 18aa) . . . . . . . . . . . .

[Southern Europe along the Mediterranean region and Middle East. established in NSW around Sydney]
Arhophalus syriacus (Reitter)

Fig. 18b

Fig. 18bb

- Terminal segment of maxillary palp slightly widened apically, with length 1.34-1.39 times its apical width (Figs. 18b, 18bb) . . . . . . . . . . . .

[Europe, North Africa and Asia; found in Melbourne]
Arhophalus rusticus (Linnaeus)

couplet 19 of 20 (from 16)

Fig. 19a

Fig. 19aa

Fig. 19b

Fig. 19bb

Prosternal process very narrow, incomplete between procoxae (Figs. 19a, 19aa); elytra with iridescent colours (Figs. 19b, 19bb) . . . . . . . . . . . .

[Violet tanbark beetle; Europe, northern Asia, USA and Canada]
Callidium violaceum (Linnaeus)

Fig. 19c

Fig. 19cc

- Prosternal process narrow, but always complete between procoxae (Figs. 19c, 19cc); elytra without iridescent colours.

. . . . . . . . . . . . Go to couplet 20 →

couplet 20 of 20 (from 19)

Fig. 20a

Fig. 20aa

Fig. 20b

Fig. 20bb

Elytra with orange markings (Figs. 20a, 20aa); body distinctly flattened (Figs. 20b, 20bb) . . . . . . . . . . . .

[USA, Canada, Europe and Asia]
Semanotus spp.

Fig. 20c

Fig. 20cc

Fig. 20d

Fig. 20dd

- Elytra uniformly brown (Figs. 20c, 20cc); body not flattened (Figs. 20d, 20dd) . . . . . . . . . . . .

[Mulberry longhorn beetle]
Trichoferus campestris (Faldermann)

Key to the 5 Cerambycidae subfamilies

couplet 1 of 4

Labrum fused to clypeus and not visible externally.

Tarsomere 4 relatively long, not concealed by lobe on tarsomere 3.

Antennal insertions (antennal foramina) lateral and not visible from above.

Lateral pronotal carinae entire and simple.

. . . . . . Parandrinae

- Labrum separate from clypeus and visible externally.

Tarsomere 4 very short and concealed by lobe on tarsomere 3.

Antennal insertions (antennal foramina) variable but they are clearly visible from above.

Lateral pronotal carinae absent, incomplete or complete.

. . . . . . . . . . . . Go to couplet 2 →

couplet 2 of 4

Lateral pronotal carinae present, often incomplete and dentate.

Procoxae strongly transverse with large exposed trochantins.

Protibia often expanded and serrate along external edge.

. . . . . . Prioninae

- Lateral pronotal carinae absent (sometimes pronotum irregularly crenulate laterally.

Procoxae oval or weakly transverse with small trochantins visible in procoxal extensions.

Protibia never expanded and serrate along external edge.

. . . . . . . . . . . . Go to couplet 3 →

couplet 3 of 4

Protibia with inner grooved antennal cleaner.

mesotibia with sulcate or comb-like antennal cleaner.

Head usually strongly deflexed or vertical.

...............exception Tmesisternini. Head is obliquely forward.

Procoxal cavites externally closed (exceptions Batocera and Rosenbergia)

. . . . . . Lamiinae

- Protibia and mesotibia without antennal cleaners.

Head usually prognathous or weakly deflexed.

Procoxal cavities externally closed or open.

. . . . . . . . . . . . Go to couplet 4 →

couplet 4 of 4

Antennal pedicel 1.5 times longer than wide and usually about 0.5 times as long as scape or antennomere 3.

Antennae short, at most reaching middle of elytra.

Mandible externally densely setose, without distinct tuft of long setae.

. . . . . . Spondylidinae

- Antennal pedicel subquadrate or transverse, ≤ 0.3 times as long as scape or antennomere 3.

Antennae variable but usually extending beyond middle of elytra.

Mandible externally glabrous or with distinct tuft of long setae.

Lateral pronotal carinae absent, incomplete or complete.

. . . . . . Cerambycinae