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.
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.
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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Breuning, S. (1965a). Revision der 35. Gattung der Pteropliini der asiatischen Region (Col. Cerambycidae).Entomologischen Arbeiten aus dem Museum G. Frey 16 :161–472.
Breuning, S. (1965b). Neue Cerambyciden aus den Sammlungen des Zoologischen Museums der Humboldt–Universität zu Berlin (Coleoptera, Cerambycidae) (vierter teil).Mitteilungen aus dem Zoologischen Museum in Berlin 41 (1): 81–93.
Breuning, S. (1966a). Nouvelles formes de Lamiaires (dix–septième partie).Bulletin de l'Institut royal des Sciences naturelles de Belgique 42 (21): 1–22.
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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.
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.
2. With complete median longitudinal groove marking deep internal endocarina, which continues into occipital area.
3. Eyes almost always deeply emarginate or divided into upper and lower parts, finely to very coarsely facetted.
4. Antennal insertions always exposed and supported by raised tubercles.
5. Frontoclypeal suture distinctly impressed. Labrum visible and free.
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.
1. Lateral pronotal carinae always absent, with secondary development in some Tmesisternini.
2. Prosternal process complete.
3. procoxal cavities almost always closed externally. Procoxae often projecting well below prosternum almost always with lateral extension and exposed trochantin.
4. Scutellum usually visible; mesoscutum often with stridulatory file or rarely absent (e.g. Rhytiphora).
5. Mesocoxal cavities usually laterally open.
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.
1. usually long and flexible with short subapical styli.
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.
1. prognathous to moderately inclined head with frontal region rarely deflexed from behind eyes.
2. Frontoclypeus usually subquadrate, occasionally distinctly longer than wide, forming rostrum
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
4. Postocular constriction absent or weak, forming distinct neck in a few 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.
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.
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.
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.
1. Prosternal process variable, ranging from broad to absent
2. Procoxal cavities round or weakly transverse often with lateral extensions and exposed protrochantin, closed internally, closed or open externally.
3. Scutellum always visible; mesoscutum rarely with median endocarina, usually with undivided stridulatory file.
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.
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.
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couplet 1 of 20
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) . . . . . . . . . . . .
- Elytra without a net-like pattern (Figs. 1c, 1cc); size variable.
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couplet 2 of 20 (from 1)
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 →
- Protibia and mesotibia without antenna cleaners (Figs. 2c, 2cc); head prognathous or weakly deflexed (Figs. 2d, 2dd).
[subfamilies Cerambycinae and Spondylidinae]
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couplet 3 of 20 (from 2)
SUBFAMILY LAMINAE
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]
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- Elytra entirely setose and of various colours and combinations (Figs. 3c, 3cc); procoxal cavities at least narrowly open posteriorly (Figs. 3d, 3dd).
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couplet 4 of 20 (from 3)
Pronotal disc flat (Figs. 4a, 4aa); elytra glossy black, finely punctate, smooth at basal part (Figs. 4b, 4bb) . . . . . . . . . . . .
couplet 5 of 20 (from 3)
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 →
- 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) . . . . . . . . . . . .
couplet 6 of 20 (from 5)
Pronotum with two longitudinal orange stripes (Figs. 6a, 6aa); each elytron with 5 longitudinal bands of black and grey rectangular spots (6b, 6bb) . . . . . . . . . . . .
couplet 7 of 20 (from 2)
SUBFAMILIES CERAMBYCINAE AND SPONDYLIDINAE
Elytra very short, not extending over abdomen (Figs. 7a, 7aa); femora strongly clavate (club-like) (Figs. 7b, 7bb) . . . . . . . . . . . .
- Elytra either completely covering abdomen (Figs. 7c, 7cc) or with only the terminal segment exposed (Figs. 7d, 7dd).
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couplet 8 of 20 (from 7)
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]
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- 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
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) . . . . . . . . . . . .
- 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) . . . . . . . . . . . .
couplet 10 of 20 (from 8)
Eye deeply emarginate, two lobes only connected by 1 row of ommatidia (Figs. 10a, 10aa). [genus Tetropium]
. . . . . . . . . . . . Go to couplet 11 →
- 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)
Pronotum densely punctate, without any glabrous raised area (Figs. 11a, 11aa); elytra reddish or yellowish brown (Figs. 11b, 11bb) . . . . . . . . . . . .
- Pronotum weakly punctate, with 3 weak shiny, raised nodules (Figs. 11c, 11cc); elytra uniformly dark brown (Figs. 11d, 11dd) . . . . . . . . . . . .
couplet 12 of 20 (from 10)
Pronotum lateral edge with sharp median projection (Figs. 12a, 12aa) . . . . . . . . . . . .
- Pronotum with lateral edge rounded (Figs. 12b, 12bb).
. . . . . . . . . . . . Go to couplet 13 →
couplet 13 of 20 (from 12)
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) . . . . . . . . . . . .
- 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)
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 →
- 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)
Frons with triangular keel (Figs. 15a, 15aa); pronotum brown with sparse yellow recumbent setae; elytra black with yellow markings (Figs. 15b, 15bb) . . . . . . . . . . . .
couplet 16 of 20 (from 14)
Eye shallowly emarginate (Figs. 16a, 16aa); Protibial spur single (Figs. 16b, 16bb). [genus Arhophalus]
. . . . . . . . . . . . Go to couplet 17 →
- Eye distinctly emarginate (Figs. 16c, 15cc); Protibial spurs paired (Figs. 16d, 16dd).
. . . . . . . . . . . . Go to couplet 19 →
couplet 17 of 20 (from 16)
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) . . . . . . . . . . . .
- 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)
Terminal segment of maxillary palp strongly securifrom, with length 1-1.26 times its apical width (Figs. 18a, 18aa) . . . . . . . . . . . .
couplet 19 of 20 (from 16)
Prosternal process very narrow, incomplete between procoxae (Figs. 19a, 19aa); elytra with iridescent colours (Figs. 19b, 19bb) . . . . . . . . . . . .
- 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)
Elytra with orange markings (Figs. 20a, 20aa); body distinctly flattened (Figs. 20b, 20bb) . . . . . . . . . . . .
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