Australian Longhorn Beetles (Coleoptera: Cerambycidae) Volume 3: Subfamily Prioninae of the Australo-Pacific Region

By Adam Slipinski, Roger de Keyzer and Mengjie Jin

Longhorn beetles — Cerambycidae — are one of the most easily recognised groups of beetles, a cosmopolitan family that encompasses more than 33,000 species in 5,200 genera worldwide. Out of the 117 beetle families occurring in Australia, Cerambycidae is the sixth largest, comprising more than 1,400 species classified in 300 genera.

Virtually all Cerambycidae feed on living or dead plant tissue and play a significant role in all terrestrial environments. 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 nutrient and energy circulation in these habitats. Longhorn beetles can cause serious damage by sometimes feeding on and eventually killing living forest or orchard trees. Many species are listed as quarantine pests because of their destructive role to the timber industry, such as the European house borer introduced into Western Australia.

This third volume in the series on Australian longhorn beetles extends to include the taxonomy of genera and species of the subfamily Prioninae of the Australo-Pacific Region. Seven tribes, 50 genera and 166 species are included. All genera and most species are diagnosed, described, illustrated and included in keys to their identification.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Dec 1, 2023
• ISBN: 9781486317325

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

Collecting and rearing longhorn beetles

Collecting of longhorn beetles can involve many different techniques such as netting, sweeping and various trapping techniques. However, for collection of Australian prionines sweeping and netting techniques generally are not productive for finding prionines. Few species are day active (Papunya picta, Sceleocantha gigas and Phaolus metallicus) and most adult prionines tend to hide during the day under bark of fallen or standing trees, under logs and billets or within timber and ground debris, and occasionally on vegetation (Rhipidocerus spp.).

Finding most prionines within their native habitat can be very difficult without the aid of specific trapping techniques which either attract or trap adults as they are moving about. The preferred techniques are light-trapping or use of flight intercept traps or a combination of the two.

Light-trapping for insects is always challenging as the weather is the controlling factor and while most prionines will come readily to lights at night, certain conditions need to be met in order to have a productive night. For most Australian prionines the optimum conditions are warm, moonless nights during late spring to summer (November to February). In dry and hot years light-trapping can also be quite productive several days after rain as the more humid conditions tend to favour insect activity. Warm nights when the moon rises very late in the evening or early in the morning can also be quite good, regardless of the moon phase. Adults of Eboraphyllus middletoni which lives in the cool temperate Nothofagus forest of New England National Park in NSW will even fly into lights on quite cold moonless nights (RDK pers obs.) but for most prionines, warm nights are favoured.

Lights that are used to attract insects are those that emit ultraviolet light — self-ballasted mercury vapour and metal halide globes or black light fluorescent tubes. Quite a diversity of mercury vapour or metal halide globes are available commercially and while some people prefer using the brightest globe they can find, 160 or 250 watt frosted globes generally work well. The globe or fluorescent tube once connected to an appropriate power source can be mounted on an aluminium pole in front of or over the top of a light sheet.

While some manufactured entomological light sheets with frames are available, a cheaper and more versatile option is to make your own light sheet. Choosing the right type of fabric for the light sheet comes down to personal preferences. Calico tends to be heavy when wet and rots readily if not properly dried. Most fabric stores will have a range of finely woven synthetic fabrics that can be used for your light sheet. Most people choose a finely woven white lace curtain, but organza is more lightweight and durable. The light sheet should ideally be 2–3 metres in length and about 1.8 metres in height and have a hem sewn along the top edge so that a rope can be inserted along its length. Always use a much longer rope so that you don’t limit your options when securing the light sheet to nearby trees. Reinforce the bottom hem of the sheet so that the sheet can be spread tightly and secured to the ground with metal pegs. On rocky ground you can secure the bottom of the light sheet with rocks or billets of timber.

Many options are available for power systems to run your lights; if you want to use a portable generator that is not too heavy, quiet and reliable then Honda make some excellent generators. Black lights can be run off car batteries or a portable battery pack and are certainly an option if you need much lighter gear for more remote field work.

Some prionines, although attracted to lights, will not always fly onto light sheets, especially if the light used is very bright. A head torch and regular inspections of the surrounding areas will often pick up species that shun the brighter light. Having a large light sheet is also advantageous for prionine beetles, giving light-sensitive species an area to land on that is away from the central light. It is always best practice to pick a relatively open area in which to set up your light trap so you can readily see the beetles as they come in; remember, beetles can approach from multiple directions. The edges of the summit of hilltops or mountains are ideal places to set up your light trap as beetles can be attracted to the light from considerable distances away.

Flight intercept traps are also useful if set up along insect flight corridors within forested areas. Longer term light traps powered by solar panels and utilising either a large funnel trap or clear Perspex or plastic flight intercept panel that empties into a trough or container with killing and preservative fluid can also be useful for collection of prionines, especially if time in the field is limited. Malaise traps are also useful when field time is limited and when longer-term biodiversity investigations warrant their use. Pitfall traps are not very useful; considerable time and effort is required to set up and maintain them and they rarely pick up prionines.

Some prionines such as Hermerius howei are readily taken at light on warm summer nights but can also be found during daylight by turning over logs or billets of timber in eucalyptus logging areas, during the seasons of adult emergence. Some species such as Agrianome spinicollis and Cnemoplites australis can also be found under bark of standing and fallen trees or, like many other Australian prionines, can be collected at bright lights at service stations and other building adjoining native bushland.

Finding prionine larvae within the forest is often not an easy process and it takes time and experience to pick timber that is likely to be infested. Large ovoid emergence holes are usually signs of longhorn activity but may not only be from prionines. Quite fresh coarse frass and long fibrous, coarsely shredded pieces of timber emanating from attacked sections of logs are usually good indicators of larval activity. Many prionine larvae are found in moist and decaying logs, some quite soft (e.g. Rhipidocerus spp. and Agrianome spinicollis); other larvae can be found in rotting logs that are structurally quite hard (e.g. Eboraphyllus middletoni and Catypnodes pascoei).

Tools required to extract larvae from a variety of timber types are generally a short-handled axe, a log splitter and a jimmy bar (also called a wrecking bar or pinch bar), and a chain saw. If use of a chainsaw is permitted it certainly minimises the risk of damage to larvae within the timber and the timber can easily be cut to appropriate lengths for ongoing storage in containers for rearing. As the biology of most of the Australo-Pacific Prioninae is poorly known it is worthwhile taking the time to identify the host plant and to record any biological attributes that you observe while the larvae are being reared.

Rearing of prionine larvae is generally not particularly difficult. It is important that you take note of the environmental conditions under which the larvae were found. If larvae were collected in cool temperate rainforest or tropical rainforest then timber should be kept moist and at temperatures similar to where the larvae were found. If the timber is quite dry or has been found in quite dry forest then less moisture is required for development of the larvae. If you have needed to extract larvae from the timber they were in, you should collect a good quantity of frass and the coarsely shredded bits of undigested timber from the workings they were in, along with enough timber in which to rear them for several years. Each larva should be transported in a separate container or in a separate compartment in a partitioned container as larvae will often attack each other if kept together (RDK has observed this happen in many cerambycid species he has reared).

Plastic tubs can be used to rear prionine larvae as long as they are relatively smooth, strong and thick as prionine larvae will often chew into the walls in search of food. It is important also that you make sure all your containers always have ventilation holes in the lids. Always store your rearing containers in a cool dark place and as the wood dries out use a clean spray bottle to maintain moisture levels. Although many Australian prionine species develop in quite moist rotting timber, others require drier conditions: RDK has reared Phaolus metallicus larvae which prefer drier conditions and usually die if the timber is kept too moist.

Rogers et al. (2002) successfully reared Prionoplus reticularis larvae on an artificial diet containing pine sawdust in a laboratory at 20°C and reduced the larval period to about 250 days compared with the normal development period of two or more years in the field. They had limited success with getting mature diapause larvae to pupate under laboratory conditions but found that all diapausing larvae exposed to field conditions from April to October successfully pupated in October and November. They also found that larvae reared at a higher temperature (25°C) had not only a higher mortality rate but also had a reduced larval weight.

Although currently we know relatively little about the biology of most of the Australo-Pacific prionine species, but you can assume that raising mature larvae of many of the species usually takes one to several years; it is important to maintain optimal conditions over that period. Some larvae will take longer to develop and you should always avoid disturbing them unnecessarily during this time. While RDK successfully reared an Eboraphyllus middletoni larva that took ten years from egg to adult, it is not known whether it would normally take a similar length of time in its natural cool temperate environment. Development may be much slower in such an environment where ambient daily temperatures are much lower than in Sydney where the larva was reared.

Throughout this volume we have recorded available information on the biology of the Australo-Pacific prionines and we encourage people to undertake further investigation to better understand the biology of these fascinating beetles.

Figure 1. Prioninae longhorn beetles in their natural habitats.

AAgrianome spinicollis (Macleay), female © Roger de Keyzer

BSceleocantha cuneata McKeown, © Roger de Keyzer

CBrephilydia jejuna (Pascoe), © Roger de Keyzer

DBifidoprionus rufus Komiya & de Keyzer, male, © Roger de Keyzer

EAnalophus parallelus Waterhouse, © Roger de Keyzer

FCnemoplitodes cephalotes (Pascoe), © Roger de Keyzer

Figure 2. Longhorn beetles in their natural habitats

AToxeutes arcuatus (Fabricius), male, © Roger de Keyzer

BToxeutes arcuatus (Fabricius), female, © Roger de Keyzer

CHermerius howei (Thomson), male, © Roger de Keyzer

DSceleocantha glabricollis , male © Roger de Keyzer

ESceleocantha carteri McKeown, © Roger de Keyzer

Figure 3. Longhorn beetles in their natural habitats

A,B Eboraphyllus middletoni McKeown, male, © Roger de Keyzer

CMethylethelius kozlovantoni (Zubov & Titarenko), female, © Roger de Keyzer

DEboraphyllus middletoni McKeown, female, © Roger de Keyzer

E,F Cacodacnus planicollis (Blackburn), © Roger de Keyzer

Figure 4. Longhorn beetles in their natural habitats

A,B Enneaphyllus aeneipennis Waterhouse, male, © Roger de Keyzer

CEnneaphyllus aeneipennis Waterhouse, female, © Roger de Keyzer

DPhaolus metallicus (Newman), female, © Roger de Keyzer

ERhipidocerus australasiae Westwood, female, © Roger de Keyzer

Figure 5. Longhorn beetles in their natural habitats

ARhipidocerus weiri Jin, de Keyzer & Ślipiński, male, © Roger de Keyzer

BRhipidocerus australasiae Westwood, female, © Roger de Keyzer

CPapunya picta Jin, de Keyzer and Ślipiński, male, © Glenda Walters

DRhipidocerus weiri Jin, de Keyzer & Ślipiński, male, © Roger de Keyzer

ECryptipus frenchi (Blackburn), male, © Jiří Lochman

FCryptipus frenchi (Blackburn), male, © Kaisa & Stan Breeden

Figure 6. Longhorn beetles in their natural habitats

AHagrides princeps (Gahan), female, © Roger de Keyzer

BHermerius howei (Thomson), female, © Roger de Keyzer

CCnemoplites edulis Newman, male, © Roger de Keyzer

DGeoffmonteithia aurivillii (Lameere), male, © Roger de Keyzer

EHermerius howei (Thomson), female, © Roger de Keyzer

FHagrides princeps (Gahan), female, © Roger de Keyzer

Figure 7. Longhorn beetles in their natural habitats

AAnalophus parallelus Waterhouse, male, © Roger de Keyzer

BBrephilydia jejuna (Pascoe), female, © Roger de Keyzer

CElaptus brevicornis Pascoe, female, © Roger de Keyzer

DPrionoplus reticularis White, male, © Arthur Anker

EEurynassa sp., female, © Roger de Keyzer

FElaptus brevicornis Pascoe, male?, © Roger de Keyzer

Subfamily Prioninae

Latreille, 1802

Introduction

The longhorns of the subfamily Prioninae are usually recognised for their heavy build and sclerotised, sombre brown or black bodies that often reach a large size, making them very popular subjects of research and collection by entomologists and various beetle enthusiasts. Over 300 genera and 1200 species of Prioninae have been described worldwide (Tavakilian & Chevillote 2022), but it is believed a large number of unknown or cryptic species are yet to be discovered (Jin et al. 2020a).

This volume departs from the convention of the series (Ślipiński & Escalona 2013, 2016) by including taxa at species level and by the expansion of the geographical cover beyond Australia to the Australo-Pacific Region (Fig. 8). In total we treat 50 genera and 166 recognised species. Due to the lack of available material and/or the complex and unresolved taxonomy within some genera, not all species are thoroughly described and illustrated.

Figure 8. Study area.

Biology and ecology

The biology of many of the Australo-Pacific Prioninae species is poorly known. From the little information available, we do know most species are active nocturnally: all species of genera in the Macrotomini, Osphryonini, Rhipidocerini, Tereticini and Parandrini have been taken at light.

Most species in the Sceleocanthini are also nocturnal except for Sceleocantha gigas. Males of S. gigas are very active day fliers and have highly modified net-like maxillary and labial terminal palpomeres which are likely to be used to detect pheromones released by the females waiting on the ground. The only known fully wingless species in the Australo-Pacific region is Psalidocoptus scaber, a large-bodied and heavily sclerotised species of unknown biology, endemic to the volcanic island of Tanna in the Vanuatu Archipelago. Švácha and Lawrence (2014) suggested that the use of pheromones by flightless female prionines is quite likely since the males are fully winged and quite mobile. Females of the Australo-Pacific genus Psalidosphryon are brachypterous and may release a pheromone to attract their males. The male of Storeyandra frenchi is brachypterous and, like the fully winged female, is nocturnal. It is not known if a pheromone is used to attract mates in this species, but if so, it may be the male which produces a pheromone to attract the fully winged female.

Barbour et al. (2006) found that females of the European prionine species, Prionus coriarius, have an eversible gland on the dorsal surface of the ovipositor that is everted when the female elevates her abdomen and extrudes ovipositor to release pheromone from a gland to attract males. Males of North American, Prionus spp. (Barbour et al. 2011) and Tragosoma spp. (Ray et al. 2012) have also been found to be attracted to female-synthesized pheromones.

The iridescent and bright blue, green and purple Phaolus metallicus in the tribe Catypnini is a diurnal species that is quite widespread but is not commonly collected. It is quite surprising that this strikingly beautiful species is not often found, since it is day active; however, that they are rapid fliers may explain why adults are not commonly seen or collected. Froggatt (1902) recorded adults of this species on leaves of Acacia decurrens and grass stalks in open forest. Another day active species in the tribe Catypnini is Papunya picta which, currently, is known only from the type series (2 specimens) and a specimen that was photographed live (Fig. 5C) near Toowoomba in 2016 (Jin et al. 2020a), but was not collected.

The larvae of many Australo-Pacific prionines are saproxylic feeders, utilising timber that is in the process of being decomposed by either white or brown rot fungi. Adults of some species such as Agrianome also attack living trees, and A. spinicollis, recorded to attack many different native and exotic trees (Hawkeswood 2005), has also been recorded as a minor pest of Pecan trees (Coombs & Crouch 1999). Cnemoplites australis is another common Australian east coast species which attacks both living and dying trees of Banksia spp. (Froggatt 1893, 1907; Fearn 1989) and a diverse range of both native and introduced trees in the genera Angophora, Betula, Casuarina, Eucalyptus, Quercus, Salix and Ulmus (Froggatt 1923; Duffy 1963; Webb 1987; Fearn 1989; Hawkeswood 1992b; Hawkeswood & Turner 2003).

Duffy (1963) stated that although Prionoplus reticularis attacks dead wood, the wood is not necessarily decayed and this may also be true for other Australo-Pacific prionines. Mature larvae of Sceleocantha carteri have been found below the bark of a newly fallen tree of a Eucalyptus sp. (Ironbark species), feeding on heartwood timber which was quite sound and only in the very early stages of decay (RDK pers. obs.). Larvae of other species like Eboraphyllus middletoni and Catypnodes pascoei utilise timber that, on the surface has begun to decay, but inside is still quite solid and only in early stages of decay (RDK pers. obs.). The larvae when tunnelling through this timber, facilitate further decay of the log and eventually after several or many years help reduce the logs to frass and other compostable materials.

Cerambycid larvae are a primary coloniser of timber and in some instances can rapidly reduce logs to frass and, in doing so, produce habitats for secondary colonisers and allow more water to enter logs (Duffy 1963). Emergence holes and tunnels of cerambycid larvae are often used as shelter by other insects and vertebrates. Fearn and Maynard (2019) state that Toxeutes arcuatus appears to be an important species in the breakdown of timber in Tasmanian forests and that the large emergence holes, pupal chambers and extensive larval galleries provide entry points and refuges for other species of saproxylic Coleoptera, especially tenebrionids. Even small vertebrates such as two species of skink, a brown tree frog and a juvenile tiger snake have been recorded using old larval galleries as retreats and winter torpor sites (Fearn 1993). The once common Lord Howe Island stick insect was reported to use the galleries of Agrianome howei as hiding places (Lea 1916). Other invertebrates sometimes encountered seeking shelter in prionine galleries are land snails, king crickets (family Stenopelmatidae), earwigs (Dermaptera), velvet worms (family Peripatopsidae) and a variety of other saproxylic beetle larvae and adults (RDK pers. obs.).

Prionine larvae are able to delay their development if conditions are unfavourable. Duffy (1963) stated that larvae of Prionoplus reticularis survived for over two and quarter years in the absence of food and during this time underwent four moults that substantially reduced their size; they resumed feeding when food was offered again. Prionine larvae also reduce in size if their timber dries out and can delay their development until such times as appropriate moisture becomes available (Duffy 1953). Cox (1906) thought this to be the case for a larva of Eurynassa australis that he reared from dry timber of Eucalyptus squamosa for two years and seven months and the same has been found by RDK when rearing a variety of Australian prionine larvae. This appears to be an adaptation that enables larvae to survive under harsh drought conditions until rains provide the moisture required for the larvae to continue their development. Unfortunately, not all larvae have the same ability to survive under such conditions and some die in their galleries before moister conditions develop (Duffy 1953; RDK pers. obs.).

In general, duration of the life cycle of prionines takes several years or much longer (Duffy 1953, 1963; Fearn & Maynard 2019). Most of the life cycle is spent in larval stages; and the vast majority of prionines overwinter as larvae (Duffy 1963; Haack et al. 2017). Oviposition involves the female simply pushing her ovipositor onto the substrate to lay eggs on, in or near the larval host plant, either singly or in batches (Duffy 1953, 1963; Linsley 1959). The eggs generally take 2–4 weeks to hatch (Duffy 1953, 1963). As they emerge from the egg, the larvae tunnel into the bark of the host plant and over time make extensive oval-shaped galleries throughout the infested timber; the galleries become loosely packed with granular frass and coarsely shredded pieces of wood (Craighead 1923; Froggatt 1923; Hay 1968; Solomon 1977; Hawkeswood 1992b). When the prionine larva is ready to pupate, it makes a pupal chamber at the end of a feeding gallery, lined with the coarsely shredded wood and large enough to contain the more sizeable pupa (Craighead 1923; Duffy 1953). Prior to pupation, larvae stop feeding, become very inactive, contract in body length and occasionally oscillate, consolidating the lining of the pupal chamber (Duffy 1953). Pupal duration is generally 3–4 weeks and newly emerged (teneral) adults generally take one to several weeks to become fully sclerotised (Duffy 1953, 1963). Adults prionines usually do not feed and generally live for only several weeks to a month (Duffy 1953; Švácha & Lawrence 2014).

Diagnosis of subfamily Prioninae

Prioninae adults are easily distinguished from the other cerambycid subfamilies represented in the Australo-Pacific region by the prothorax with variously developed lateral carina, the strongly transverse procoxae with large exposed trochantins and the protibia often expanded and serrate along the external edge. Prioninae larvae differ from the legless larvae of Lamiinae by having well-developed legs and the head transverse with the occipital foramen divided, and from the more closely similar larvae of Cerambycinae by having mandibles with the incisor areas grooved, the maxillary mala covered by distinct dense setae and the antenna often 2-segmented.

Adult

Detailed morphology of adult and larval Cerambycidae is covered by Švácha and Lawrence (2014) and in Volume 1 of this series (Ślipiński & Escalona 2013). The nomenclature of the main body parts of adults is explained on the schematic Figures 9–11.

Head

prognathous to moderately inclined. Frontoclypeus usually transverse; median longitudinal groove (Fig. 10C) and associated internal endocarina usually present (absent in Parandrini) but often incomplete posteriorly. Postocular constriction absent. Frontoclypeal suture (Fig. 11E) usually impressed, arcuate or angulate, rarely not externally visible in median section (some Parandrini). Labrum transverse, setose, usually free but occasionally fused (Fig. 11B) with anteclypeus (Osphryonini, Parandrini). Mandibles (Fig. 10D,E) variable, often longer and stronger in males apically pointed, but sometimes (Catypnini, Rhipidocerini and most of Parandrini) bearing ventral subapical tooth (Fig. 11F). Maxilla (Fig. 10F) with distinct, setose galea and strongly reduced lacinia. Labial (Fig. 10G) and maxillary palps elongate, terminal palpomeres fusiform to somewhat expanded apically.

Eyes

variable in size, usually large, shallowly to deeply emarginate near antennal insertions, finely to very coarsely facetted; moderately to very narrowly separated dorsally. Antennal insertions usually close to mandibular condyles with foramen bearing thin rim, usually directed laterally and articulation point situated on the rim.

Antenna

11-segmented, rarely 12-segmented, strongly sexually dimorphic, with males having longer and more strongly modified antennomeres; antenna is usually moniliform or filiform but can be serrate, flabellate or biflabellate. Scape always distinctly longer than pedicel; pedicel short, usually transverse, always shorter than antennomere 3.

Prothorax

transverse with lateral carina usually complete (Fig. 11A,D,E) and often dentate or spinose; in Rhipidocerini lateral carina is reduced and often present only as a median spine. Prosternal process broad, distinctly separating procoxae, almost always extending to mesoventrite. Procoxal cavities strongly transverse with exposed protrochantin, open internally and externally; postcoxal hypomeral projections variable.

Pterothorax.

Scutellum always visible (Fig. 9A); mesoscutum without stridulatory file. Mesocoxal cavities closed or rarely open to mesepimeron (Macrotomini). Mesocoxae flat; mesotrochantin always visible. Elytra usually completely covering abdomen, rarely narrowing apically and dehiscent. Wings present (reduced in female of Psalidosphryon and in male of Storeyandra), always with radial cell, usually with wedge cell (absent in Parandrini) and with 3 or 4 free veins in median field (Fig. 9C). Legs usually strong and in Macrotomini often bearing rows of spines along ventral and outer surfaces. Tibial spurs well developed and subequal (distinctly uneven in Parandrini), usually paired, sometimes reduced to single spur. Tarsi with tarsomere 3 usually forming large, divided lobe.

Abdomen

with ventrites 1 to 4 subequal and freely articulated; sclerotised and at most shortly setose; ventrite 5 apically arcuate and usually setose apically. Aedeagus with long anterior struts; endophallus mostly with pair of basal sclerites and patches of microspinules in middle and apical sections, rarely with flagellum; parameres well developed and separate.

Ovipositor

usually long and flexible with short lateral styli but ovipositor distinctly sclerotised with laterodorsal styli in Parandrini.

Larva

Body subcylindrical or slightly depressed with dorsal and ventral ampullae on abdominal segments (Fig. 13E,F).

Head

(Fig. 12) prognathous, usually slightly transverse, and deeply retracted into prothorax; posterior edge of head capsule usually notched posteriorly. Epicranial stem (coronal suture) and median endocarina present. Stemmata variable, 6 to none. Frontoclypeal suture present. Clypeus trapezoidal, filling the area between dorsal mandibular articulations. Labrum (Fig. 12C) free, usually transverse but cordate and very long in Parandrini. Antennae (Fig. 313C) moderately long, 2- or 3-segmented with terminal antennomere usually well developed, sensorium flat or protuberant. Mandibles short with well-developed pseudomola, unidentate or bidentate apically. Ventral mouthparts slightly retracted (Fig. 13A); maxillary articulating area well developed, usually divided; maxillary mala (Fig. 13D) cylindrical to expanded with short and sparse setae; maxillary palp 3-segmented; labial palp (Fig. 13B) 2-segmented. Hypostomal rods strongly convergent posteriorly. Tentorial bridge well developed, dividing occipital foramen into two parts (Fig. 12B).

Legs

short (Fig. 13F), 3-segmented.

Abdomen

with tergum IX simple.

Figure 9. Adult morphology

Adorsal view

Blateral view

Chind wing

[Anne Hastings from Ślipiński & Escalona 2013, modified]

Figure 10. Adult morphology

Aventral view

Bhead, ventral

Chead, dorsal

Dmandible, ventral

Emandible, dorsal

Fmaxilla, ventral

Glabium, ventral

[Anne Hastings from Ślipiński & Escalona 2013]

Figure 11. Adult morphology

A,B Storeyandra frenchi (Parandrini), head and prothorax:

Alateral

Bdorsal

C–E Cryptipus frenchi (Macrotomini), head and prothorax:

Clateral

Dventral

Edorsal

FRhipidocerus australasiae (Rhipidocerini), head anterior view

[from Ślipiński & Escalona 2013]

Figure 12. Larval morphology, head: Cnemoplites sp. (Prioninae)

Adorsal view

Bventral view

Cfrontal view

Dantero-lateral view

[from Ślipiński & Escalona 2013]

Figure 13. Larval morphology: Cnemoplites sp.

Amouthparts, ventral

Blabium and maxilla, ventral

Cantenna

Dmaxilla, ventral

Edorsal habitus

Flateral habitus

[from Ślipiński & Escalona 2013]

Higher classification of Prioninae

Current generic and tribal classifications of Prioninae have been inherited from comprehensive works by T. Lacordaire (1868) and A. Lameere (1912–1919) and have been perpetuated for over a hundred years with various adjustments, reflecting ongoing research, mostly on regional scales. Parandrinae, treated as a tribe in Prioninae by Lacordaire (1868) and Lameere (1919) were treated more recently by some authors as a separate subfamily (Linsley 1962; Villiers 1978; Napp 1994; Monné 2006). However, the status of Parandrini as a tribe within Prioninae, considered by Švácha and Lawrence (2014), has been further stabilised by molecular evidence (Nie et al. 2021; Jin et al. 2022b, 2023) and thus is treated as such in this book.

The current tribal classification of Prioninae (Bousquet et al. 2009) is unsatisfactory, mostly artificial, and requires thorough revision (Švácha & Lawrence 2014). The molecular studies by Jin et al. (2020a), and especially by Jin et al. (2023), although heavily focused on Australian representatives, are the only larger molecular studies of world Prioninae that provide some evidence of the relationships of the major branches of this group (Fig. 14). They show that many traditionally recognised groups like Anacolini, Eurypodini, Aegosomatini or Calipogonini are paraphyletic, and that several Australian genera have been misplaced and required establishment of new tribes. Jin et al. (2023) erected two new tribes (Rhipidocerini and Osphryonini) and resurrected tribes Catypnini and Sceleocanthini for misplaced Australo-Pacific genera. They also found that the New Caledonian endemic Acideres ricaudii Guerin-Meneville, classified in Aegosomatini, does not belong to Prioninae but to Cerambycinae as already supposed by Vives et al. (2008).

The simplified cladogram (Fig. 14) summarises the results of Jin et al. (2023), showing that the Australo-Pacific Prioninae comprise several clades with a highly diverse Macrotomini the largest and most diverse group. Representation of world Prioninae was limited in the study by Jin et al. (2023), and many critical genera and species have not been examined. However, based on the tentative results in Jin et al. (2023) the tribes Osphryonini and Catypnini appear to be confined to the Australo-Pacific Region, Sceleocanthini is an Australian endemic, and Rhipidocerini, having closely related representatives in Australia, New Zealand and Chile, can be regarded as the only Gondwanan group identified so far.

Figure 14. Phylogenetic relationships of major clades within subfamily Prioninae (based on Jin et al. 2023).

Legend

Classification of Australo-Pacific Prioninae

Subfamily: Prioninae Latreille, 1802

Tribe: Macrotomini Thomson, 1861

Subtribe: Remphanina Pascoe, 1869

Genus: Agrianome Thomson, 1864

Species: A. fairmairei (Montrouzier, 1861)

A. howei (Olliff, 1889)

A. loriae Gestro, 1893

A. spinicollis (Macleay, 1826)

Genus: Analophus Waterhouse, 1877

Species: A. niger Gahan 1894

A. parallelus Waterhouse, 1877

A. septentrionalis Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

Genus: Archetypus Thomson 1861

Species: A. acutus Komiya & Drumont, 2022

A. longicornis Komiya & Drumont, 2022

A. marginatus Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

A. parandroides Thomson, 1861

A. splendens sp. nov.

Genus: Brephilydia Pascoe 1861

Species: B. fearni Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

B. jejuna (Pascoe, 1864)

Genus: Cnemoplites Newman 1842

Species: C. argodi Lameere, 1903

C. australis (Erichson, 1842)

C. binnaburensis sp. nov.

C. cnemoplitoides (Thomson, 1877)

C. edulis Newman, 1842

Genus: Cnemoplitodes gen. nov.

Species: C. cephalotes (Pascoe, 1864) comb. nov.

Genus: Cryptipus Jin, de Keyzer & Ślipiński, 2020a

Species: C. frenchi (Blackburn, 1892)

Genus: Cryptobelus Thomson, 1877

Species: C. gestroi Thomson, 1878

C. papuanus sp. nov.

Genus: Drumontia gen. nov.

Species: D. edwardsii (Montrouzier, 1861) comb. nov.

Genus: Eurynassa Thomson, 1864

Species: E. australis (Boisduval, 1935)

E. servillei Thomson, 1864

E. stigmosa Newman, 1840

E. tuberculicollis Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

Genus: Geoffmonteithia Jin, de Keyzer & Ślipiński, 2020a

Species: G. aurivillii (Lameere, 1903)

Genus: Gnathonyx Gahan, 1864

Species: G. amplitarsalis Komiya & Nylander, 2005

G. heteromandiblaris Komiya & Nylander, 2005

G. inermis Komiya & Nylander, 2005

G. longiscapis Komiya & Nylander, 2005 stat. nov.

G. orientalis Komiya & Nylander, 2005

G. piceipennis Gahan, 1894

Genus: Hagrides Jin, de Keyzer & Ślipiński, 2020a

Species: H. blackburni (Lameere, 1903)

H. gahani (Lameere, 1912) comb. nov.

H. gracilis sp. nov.

H. incertus sp. nov.

H. mandibularis Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

H. princeps (Gahan, 1893)

Genus: Hermerius Newman, 1844

Species: H. fairmairei (Lameere, 1912)

H. howei (Thomson, 1864)

H. impar Newman, 1844

H. occidentalis Jin, de Keyzer, Hutchinson, Pang & Ślipiński, 2020b

H. prionides (Thomson, 1864)

Genus: Komiyaphus gen. nov.

Species: K. vicksoni (Nylander & Komiya, 2005) comb. nov.

Genus: Pseudoplites Lameere, 1916

Species: P. hamali (Lameere, 1903)

P. rugosus sp. nov.

Genus: Teispes Thomson, 1864

Species: T. insularis (Hope, 1842)

Genus: Utra Jordan, 1895

Species: U. nitida Jordan, 1895

Genus: Xaurus Pascoe 1867

Species: X. bennigseni Lameere, 1912

X. depsarius Pascoe, 1867

X. papuus Lansberge, 1884

Subtribe: Macrotomina Thomson, 1961

Genus: Anomophysis Quentin & Villiers, 1981

Species: A. australis Quentin & Villiers, 1981

A. coxalis (Gahan, 1900)

Subtribe: Mallodonina Thomson, 1861

Genus: Aurilethrius nom. nov.

Species: A. cariosicollis (Fairmaire, 1877) comb. nov.

A. macrothorax (Montrouzier, 1861) comb. nov.

Genus: Samolethrius Vitali, 2008

Species: S. fairmairei nom. nov.

S. subnitidus (Aurivillius, 1928)

Genus: Olethrius Thomson, 1861

Species: O. admiralis Gressitt, 1959

O. koronivia sp. nov.

O. salomonum Vitali, 2008 stat. nov.

O. scabripennis Thomson, 1865

O. tyrannus (Thomson, 1861).

Subtribe: Xixuthrina Lameere, 1912

Genus: Hastertia Lameere, 1912

Species: H. bougainvillei Lameere, 1912

Genus: Omotagus Pascoe, 1867

Species: O. lacordairii Pascoe, 1867

Genus: Xixuthrus Thomson, 1864

Subgenus: Xixuthrus Thomson, 1864

Species: X. arfakianus Lansberge, 1884

X. axis Thomson, 1877c

X. costatus (Montrouzier, 1855)

X. drumonti Zubov & Titarenko, 2020

X. fominykhi Titarenko & Zubov, 2018

X. ganglbaueri Lameere, 1912

X. gressitti Marazzi, Marazzi & Komiya, 2006

X. lansbergei Lameere, 1912

X. microcerus (White, 1853)

X. nycticorax Thomson, 1877b

X. parallelus (Gressitt, 1959) comb. nov.

X. solomonensis Marazzi & Marazzi, 2006

X. stumpei Titarenko & Zubov, 2018

Subgenus: Daemonarthra Lameere, 1903

Species: X. (D.) granulipennis Komiya, 2000

X. (D.) helleri (Lameere, 1903)

X. (D.) lameerei Marazzi, Marazzi & Komiya, 2006

X. (D.) thomsoni Marazzi, Marazzi & Komiya, 2006

Subgenus: Megathurus subgen. nov.

Species: X. (M.) heros (Gräffe, 1868)

X. (M.) terribilis Thomson, 1877

Tribe: Osphryonini Jin, de Keyzer, Ashman, Zwick & Ślipiński, 2023

Genus: Osphryon Pascoe, 1869

Species: O. adustus Pascoe, 1869

O. bispinosus Nylander, 1998

O. delahayei Voitsekhovskii, 2020

O. elina Voitsekhovskii, 2020

O. excilis Komiya, 2020

O. forbesi Gahan, 1894

O. granuliger Aurivillius, 1926

O. hirticollis Gahan, 1894

O. pallidipennis Gressitt, 1951

O. satori (Zhang & Barclay, 2021) comb. nov.

O. subitanus Gressitt, 1959

O. sudestus Gressitt, 1959

O. tridentatus Gressitt, 1959

O. wauensis Nylander, 1998

O. woodlarkensis Gressitt, 1959

Genus: Psalidocoptus White, 1856

Species: P. scaber White, 1856

Genus: Psalidosphryon Komiya, 2001

Species: P. andreevi Delahaye, Komiya, Drumont & Shapovalov, 2021

P. spiniscapus (Schwarzer, 1924)

Tribe: Rhipidocerini Jin, de Keyzer, Ashman, Zwick & Ślipiński, 2023

Genus: Eboraphyllus McKeown, 1945

Species: E. middletoni McKeown, 1945

Genus: Enneaphyllus Waterhouse, 1877

Species: E. aeneipennis Waterhouse, 1877

E. rossi Blackburn, 1890

Genus: Prionoplus White, 1843

Species: P. reticularis White, 1843

Genus: Rhipidocerus Westwood, 1842

Species: R. australasiae Westwood, 1842

R. weiri Jin, de Keyzer & Ślipiński, 2022

Tribe: Scleocanthini Lacordaire, 1868

Genus: Sceleocantha Newman, 1840

Species: S. carteri McKeown,1938

S. cuneata McKeown, 1938

S. garnseyi McKeown, 1938

S. gigas (Carter, 1913)

S. glabricollis Newman, 1840

S. gracilis sp. nov.

S. pilosicollis (Hope, 1834)

Tribe: Tereticini Lameere, 1913

Genus: Aesa Lameere, 1912

Subgenus: Aesomima subgen. nov.

Species: A. (A.) dimidiata (White, 1853)

A. (A.) glabra Komiya & Drumont, 2013

Subgenus: Aesa s. str.

Species: A. (A.) carpentariae (Blackburn, 1894) comb. nov.

A. (A.) iriana Komiya & Drumont, 2013

A. (A.) lemannae sp. nov.

A. (A.) magnetica sp. nov.

A. (A.) rentzi sp. nov.

Genus: Elaptus Pascoe, 1867

Species: E. brevicornis Pascoe, 1875

E. prionoides (Pascoe, 1875)

E. simulator Pascoe, 1867

Genus: Howea Olliff, 1889

Species: H. angulata Olliff, 1889

H. doensis (Delahaye, Drumont & Salesne, 2022) comb. nov.

H. kudrnai (Drumont & Vives, 2007) comb. nov.

H. nearnsi (Komiya & Drumont, 2013) comb. nov.

Genus: Paulhutchinsonia Jin, de Keyzer & Ślipiński, 2020a

Species: P. gamora sp. nov.

P. nebula sp. nov.

P. pilosicollis (Wilson, 1923)

Tribe: Catypnini Lacordaire, 1868

Genus: Bifidoprionus Komiya & de Keyzer, 2011

Species: B. rufus Komiya & de Keyzer, 2011

Genus: Cacodacnus Thomson, 1861

Species: C. hebridanus (Thomson, 1864) complex

C. occidentalis sp. nov.

C. planicollis (Blackburn, 1895)

Genus: Catypnes Pascoe, 1864

Species: C. macleayi (Pascoe, 1864)

C. marazziorum Drumont, Komiya & Weigel, 2021

C. salesnei (Delahaye, Drumont & Komiya, 2016)

Genus: Catypnodes gen. nov.

Species: C. pascoei (Lameere, 1904) comb. nov.

Genus: Methylethelius Zubov & Titarenko, 2019 stat. nov.

Species: M. dentifrons (Aurivillius, 1926) comb. nov.

M. kozlovantoni (Zubov & Titarenko, 2019) comb. nov.

Genus: Papunya Jin, de Keyzer & Ślipiński, 2020a

Species: P. picta Jin, de Keyzer & Ślipiński, 2020a

Genus: Phaolus Pascoe, 1863

Species: P. metallicus (Newman, 1838)

Genus: Toxeutes Newman, 1840

Species: T. arcuatus (Fabricius, 1787)

Tribe: Parandrini Blanchard, 1845

Genus: Caledonandra Santos-Silva, Heffern & Matsuda, 2010

Species: C. austrocaledonica (Montrouzier, 1861)

C. passandroides (Thomson, 1867)

Genus: Komiyandra Santos-Silva, Heffern & Matsuda, 2010

Species: K. araucariae (Gressitt, 1959) comb. nov.

K. barclayi (Santos-Silva, 2011) comb. nov.

K. birai (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. bougainvillensis (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. drumonti Santos-Silva, Heffern & Matsuda, 2010

K. gressitti (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

Komiyandra irianjayana Santos-Silva, Heffern & Matsuda, 2010

K. menieri Santos-Silva, Heffern & Matsuda, 2010

K. norfolkensis (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. oberthueri (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. queenslandensis (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. rothschildi (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

K. solomonensis (Arigony, 1983) comb. nov.

K. striatifrons (Fairmaire, 1881) comb. nov.

K. weigeli (Santos-Silva, Heffern & Matsuda, 2010) comb. nov.

Genus: Malukandra Santos-Silva, Heffern & Matsuda, 2010

Species: M. hornabrooki Santos-Silva, Heffern & Matsuda, 2010

M. jayawijayana Santos-Silva, Heffern & Matsuda, 2010

Genus: Storeyandra Santos-Silva, Heffern & Matsuda, 2010

Species: S. frenchi (Blackburn, 1895)

Key to tribes of Prioninae in the Australo-Pacific Region

1Antenna short, usually not reaching beyond posterior margin of prothorax, moniliform with well-defined ventral sensory areas on antennomeres 3–10. Protibial spurs strongly uneven with outer one much longer than inner one, stouter and hook-like; empodium with single seta. Lateral carina of pronotum smooth and complete ( Fig. 11A,B ) . Lobe of tarsomere 3 narrow, only slightly broader than tarsomere 4, and often not divided . . . . . . . . . Parandrini

-Antenna almost always extending beyond posterior margin of prothorax, but variable. Protibial spurs almost straight, subequal or single; empodium with 2 setae. Lateral carina of pronotum almost always uneven, partly missing, serrate, crenulate or strongly dentate; lobe of tarsomere 3 much broader than tarsomere 4 and divided . . . . . . . . . 2

2(1) Mandible always with ventral preapical tooth ( Fig. 11F ) . . . . . . . . . 3

-Mandible without ventral preapical tooth ( Fig. 11D ) . . . . . . . . . 4

3(2) Metanepisternum strongly narrowing posteriorly over more than half its length ( Fig. 106D ) . Male antenna always flabellate, bi-flabellate with flabella broad and not distinctly setose. Prothorax with lateral carina strongly reduced, visible between lateral spine and posterior edge; lightly sclerotised green or yellowish-brown beetles . . . . . . . . . Rhipidocerini

-Metanepisternum only slightly constricted at posterior edge or narrowing from last third of its length. Male antenna almost always filiform to weakly serrate; if antenna flabellate then flabella densely setose and lateral carina of prothorax complete at least in anterior half; mostly brown and well-sclerotised beetles but with notable exceptions ( Figs 143 , 144 ) . . . . . . . . . Catypnini

4(2) Lateral carina of prothorax anteriorly explanate, expanding to sharp medial projection, then reduced posteriorly. Protibia flat, distinctly expanded and variously dentate along outer margin; remaining tibiae usually externally dentate. Protibial spurs slightly uneven, separated by median projection . . . . . . . . . Sceleocanthini

-Lateral carina of prothorax not as above, weakly crenulate, dentate to multidentate, or with multiple projections. Protibia usually narrow and spinose or smooth along outer margin; protibial spurs usually subequal and without projection between them. . . . . . . . . . 5

5(4) Prothorax with lateral carina having 2 to 4 long, sharp or rounded projections; elytral apex often bispinose, with outer spine sometimes reduced, rarely absent; labrum heavily sclerotised, usually firmly fused with clypeus; at least 4–5 terminal antennomeres with strong dorsal longitudinal ridges . . . . . . . . . Osphryonini

-Prothorax with lateral carina smooth, uneven, crenulate or dentate; elytral apex rounded or with only single spine at suture; labrum membranous or lightly sclerotised and free; antennomeres usually without longitudinal ridges or fine ridges on lateral surfaces only . . . . . . . . . 6

6(5) Eyes distinctly emarginate around antennal insertions. Small to medium brown beetles; antenna in male usually flattened and serrate ( Fig. 117A ) . Elytra rounded apically . . . . . . . . . Tereticini

-Eyes shallowly concave or not emarginate around antennal insertions. Medium and large brown or black beetles; antenna in male usually filiform. Elytra often with inner apical angle spinose . . . . . . . . . Macrotomini

Review of Australo-Pacific Prioninae

Tribe Macrotomini Thomson

Macrotomitae Thomson, 1861: 312. Type genus: Macrotoma Audinet-Serville, 1832.

Archetypi Lameere, 1912: 180. Type genus: Archetypus Thomson, 1861.

References: Jin et al. 2020a, 2020b, 2023 (in press).

Diagnosis

The Australo-Pacific taxa of Macrotomini are recognised by the eyes at most very shallowly emarginate, the mandible with a single apical tooth, the antenna usually filiform, the prothorax with the lateral carina variously dentate, and the elytral apices usually with a single sutural spine. Mescocoxal cavities broadly open to mesanepisternum and mesepimeron.

Description

Moderately large to very large (15–140 mm), brown or black, mostly glabrous beetles.

Head

oblique or prognathous with median groove and endocarina. Frontoclypeal suture usually distinct; clypeus and labrum separate; labrum densely setose. Antennal insertions lateral, separated from mandibular articulations, placed on tubercles. Mandibles in males often larger than in females, without ventral subapical tooth.

Antennae

variable, usually filiform; scape variable; antennomere 3 usually longer than scape, sometimes very long, extending to posterior margin of prothorax; antennomeres 3–11 with lateral or ventral sensory areas.

Prothorax

with variously developed, serrate to strongly spinose lateral carina. Procoxal cavities open externally. Prosternal process complete, extending to mesoventrite.

Pterothorax

. Mesocoxal cavities broadly open to mesanepisternum and mesepimeron; mesotrochantin large, exposed; metanepisternum broad, not strongly narrowing posteriorly; mesepimeron very narrow.

Legs

strong. Femora and tibiae often with various spines. Protibia with acute outer apical spine and 2 subequal spurs. Tarsi with tarsomere 3 usually broad and divided; empodium with 2 setae.

Remarks

The generic classification and the constitution of Macrotomini has a long and confused history. James Thomson (1861) erected the tribes Macrotomitae and Mallodonitae that have been adopted as Macrotomini and Mallodonini (or Mallodontini, Bouchard et al. 2011) by subsequent researchers. Further, Thomson (1864) proposed a rearranged classification, recognising tribe Macrotomini with subtribes Macrotomorphes and Mallodonmorphes. Mallodonini as defined by Thomson (1864) was distinguished from Macrotomini by having wider bodies (especially in males); shorter antennae, just surpassing the middle of the elytra and a stouter and longer antennal scape; the male pronotum with smooth facets, female pronotum scabrous or punctate; the prosternal process slightly projecting; the mesoventral process laminiform; the legs unarmed with tarsomere 5 as long as 1–4 combined (Thomson 1864; Santos-Silva & Galileo 2010).

Lacordaire (1868) retained the tribal levels for Macrotomides and Mallodontides of Thomson (1864) but proposed a separate tribe Remphanides for some genera he removed from Macrotomides that exhibit sexually dimorphic punctures on the male pronotum and sometimes on the abdomen. Pascoe (1869) catalogued cerambycid species from the Malay Archipelago and applied the Thomson-Lacordaire system pointing out that several distinguishing characters used in that system were questionable. Following his revision of the entire subfamily Prioninae of the world in several papers published between 1902 and 1912 and summarised his work in the series Genera Insectorum (Lameere 1919), Auguste Lameere in his final publication combined the three tribes recognised by Lacordaire, and subdivided them into seven subgroups, most of which are still recognised as subtribes by current authors. However, the debate concerning the distinctiveness of Macrotomini and Mallodonini has not subsided, and Quentin and Villiers (1975) recognised both as valid, being distinguishable by the shape and punctures of the pronotum. Following their definition of the tribe, Quentin and Villiers (1981) revised the tribe Macrotomini of the Old World (except for the Ethiopian region). The separation of Macrotomini and Mallodonini was adopted subsequently by Drumont and Komiya (2010), Bouchard et al. (2011) and Monné et al. (2016), building the current classification of family Cerambycidae. Vitali (2008) re-established the tribe Remphanini (as Rhaphipodini) mainly for the genera formerly included in Macrotomini (Rhaphipodi) by Lameere (1919), however the characters used to define this tribe (spiny legs, lighter colour, reduced elytral puncturing) are not useful for distinguishing higher taxa within the Prioninae.

Jin et al. (2020a and 2023) published a comprehensive molecular phylogeny of the Australo-Pacific Prioninae and delimited the subtribes and genera of the Macrotomini (Fig. 15). In addition to the strong molecular support for the Macrotomini clade, Jin et al. (2020a) found that mesocoxal cavity open to both mesanepisternum and mesepimeron and eye not emarginate near antennal insertions provide two unique morphological character states which can be used to separate Macrotomini from all remaining Prioninae in the region. We have confirmed these diagnostic characters in many exotic Macrotomini taxa, including Mallodon Lacordaire and Remphan Waterhouse, supporting synonymy of these tribes, as suggested partially by many authors (e.g. Monné & Hovore 2006; Silva-Santos & Martins 2006; Santos-Silva & Galileo 2010).

The current division of Macrotomini into subtribes Jin et al. (2023), Bousquet et al. (2009), Bouchard et al. (2011) requires further research based on larger world-wide sampling. It is currently based on restricted molecular evidence alone (Fig. 15) and no stable morphological characters are available to support it. Consequently, the genera in this book are arranged alphabetically.

The largest clade, Remphanina has not been thoroughly investigated as only two rather distinctly related genera from the outside of Australo-Pacific region have been included, and that clade Remphan + Rhaphipodus forms a sister group to the remaining highly diverse Australo-Pacific taxa. It is conceivable that with inclusion of more taxa, the Remphanina could be limited to the underrepresented taxa while the large sister clade will be regarded as Cnemoplitina. Based on Jin et al. (2023) there is no support for recognition of subtribe Archetypina as suggested by Bousquet et al. (2009) because the type genus, Archetypus is deeply embedded within the Rhemphanina.

Mallodonina and Xixuthrina form sister clades with strong support, however the sequence obtained from a specimen from the Solomon Islands, similar to Xixuthrus lansbergei Lameere has been recovered outside of the clade containing remaining Xixuthrina + Mallodonina which requires further research.

Anomophysis, in agreement with Quentin and Villiers (1975), is the sole representative of true Macrotomina in the Australo-Pacific Region. Macrotomina has been recovered as a sister taxon to the mostly Afrotropical Cantharocnemis Audinet-Serville suggesting that the tribe Cantharocnemini should be synonymised with Macrotomini or included there as a subtribe.

During our research we established that the genus Apsectogaster Thomson, 1877 (type species: A. flavipillis Thomson; Fig. 159I), sometimes (Lameere 1903) considered as a subgenus of Cnemoplites, is from South Africa not from Australia, and is a junior synonym of Erioderus Thomson, 1861 (new synonym). Consequently, Apsectrogaster flavipillis Thomson, 1877, which is likely a senior synonym of Erioderus candezei (Lameere, 1903) has been removed from the Australian fauna.

In the key and descriptions the abbreviation SDP refers to a dense punctation present in males of many species on the prothorax, scutellum and the ventral side of the body.

Figure 15. Phylogenetic relationships of major clades within tribe Macrotomini (based on Jin et al. 2023)

Legend

Key to genera of Australo-Pacific Macrotomini

1Elytra with dense setose pile between shiny and glabrous intervals ( Fig. 96C ) . . . . . . . . . Xixuthrus

-Elytra glabrous or with setae of various densities originating from distinct setiferous punctures . . . . . . . . . 2

2(1) Mandible with dorsal subapical tooth very close to apex of mandible so mandible appears bifid apically ( Fig. 79C ) ; pronotum and elytra without traces of distinct punctures or setae . . . . . . . . . Utra

-Mandible without additional tooth or with additional tooth positioned near middle of mandible . . . . . . . . . 3

3(2) Abdominal ventrites 1–4 with deep setose pockets ( Fig. 57E ) . . . . . . . . . Gnathonyx

-Abdominal ventrites glabrous, sparsely pubescent or with transverse patches of setae . . . . . . . . . 4

4(3) Abdominal ventrites 1–4 with transverse patches of yellow setae ( Fig. 41C ) . . . . . . . . . 5

-Abdominal ventrites sparsely setose or glabrous, without patches of dense setae . . . . . . . . . 8

5(4) Antennal scape extending to or beyond anterior pronotal margin ( Fig. 41B ) ; antennomere 3 distinctly longer than scape and extending to posterior margin of pronotum; pronotum glabrous or with few sparse setae laterally; protibia always with at least few spines along outer margin ( Fig. 41E ) . . . . . . . . . Cnemoplites [part: C. edulis , C. argodi , C. binnaburensis ]

-If antennal scape extending to close to anterior pronotal margin, then pronotal disc is regularly setose and/or protibia lacks spines along outer margin . . . . . . . . . 6

6(5) Pronotum with distinct, often dense yellowish setae ( Figs 67D ) ; antennal scape relatively long, almost extending to anterior margin of pronotum ( Fig. 67A ) ; protibia not spinose along outer margin ( Fig. 68A ) . . .