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Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein–coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Paleozoic origin of all modern beetle suborders and a Triassic–Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.

Alcestoma queenslandicum sp. n. (Figs 2–3, 6, 9–10, 28) Diagnosis. This species resembles A. serropalpoides in general shape and coloration, but differs in the coarser, sparser pronotal punctation, more densely setose upper surfaces and more sharply serrate antennomeres 4–10; unfortunately the aedeagus of that species has not been described. A. queenslandicum differs from A. insulare, not only in general shape and colour (darker and more slender) but in the structure of the aedeagus, which has a narrowly rounded parameroid apex, lacking the apical row of spines which characterises the Seychelles species. Description. Total length: males, 7.80–13.10 mm (10.60 ± 1.70, n = 22), females, 7.00–22.00 (13.00 ± 3.60, n = 69); body 4.05–5.25 (4.13) times as long as combined elytral width. Colour of head black above; pronotum reddish-brown to dark brown or black, sometimes reddish along margins; elytra reddish-brown to dark brown or black, sometimes reddish along suture; undersurfaces, legs and antennae yellowish-brown to reddish-brown or dark brown. Upper surfaces densely, more or less evenly clothed with moderately long, inclined, fine hairs; undersurfaces with somewhat shorter hairs. Head 1.10 times as long as wide, abruptly constricted to form neck, which is 0.64 times as wide as head. Eye about 0.54 times as long as head width, slightly emarginate anteriorly, protruding and finely facetted, with short, fine interfacetal hairs; distance across eyes 10.4 times shortest distance between them both dorsally and ventrally. Vertex without transverse ridge. Temples 0.13 times as long as eye length. Antennal insertions concealed by frontal ridges which extend slightly into eye emargination. Clypeus 0.30 times as long as wide with parallel sides and truncate apex. Labrum 0.65 times as wide as clypeus, 0.61 times as long as wide, with sides slightly curved, apex subtruncate and surface densely setose. Antennae in male about 1.1 times as long as head width behind eyes; antennomere length ratio: 1.67: 1.00: 1.87: 1.93:1.80: 1.80: 1.67: 1.87: 1.67: 1.53: 2.47; antennomere length/width: 1.67, 1.07, 2.15, 1.81, 1.50, 1.59, 1.67, 1.93, 1.79, 1.77, 3.70. Antenna in female 1.73 times as long as head width behind eyes; antennomere length ratio: 1.59: 1.00: 1.59: 1.71: 1.82: 1.94: 1.88: 2.00: 1.94: 1.88: 3.77; antennomere length/width: 1.59, 1.13, 1.42, 1.00, 1.03, 1.18, 1.28, 1.31, 1.43, 1.52, 1.77. In both sexes antennomeres 1–3 slightly widened apically; 4–10 distinctly serrate; 11 widest at about middle, acute at apex and not weakly divided by transverse impression; mandible about 1.9 times as long as wide; outer edge sharply curved at apical third, apex bidentate but with smaller tooth subapical; incisor edge simple; mola absent; prostheca consisting of a patch of short setae. Galea 1.2 times as long as wide, widest near broadly rounded apex, densely setose; lacinia slightly shorter than and 0.2 times as wide as galea, subacute apically; first maxillary palpomere in male 1.5 times as long as wide and slightly shorter than second, which is 0.87 times as long as wide; third palpomere about twice as long as second, more or less globular and deeply excavate and fourth palpomere 1.5 times as long as third, 4.3 times as long as wide and narrowly rounded apically; palp organ relatively complex, with at least 8 primary lobes and numerous secondary lobes. Ratio of palpomere lengths in female 1.00: 3.60: 2.80: 5.60; apical palpomere 2.67 times as long as wide, widest near apex, which is obliquely truncate. Mentum 0.6 times as long as wide, widest at truncate apex; ligula longer than mentum, widest at apex, which is slightly emarginate; labials palps separated by slightly less than the basal width of one; ratio of palpomere lengths 1.00: 2.60: 3.80; apical palpomere 2.09 time as long as wide, widest subapically, with obliquely truncate apex. Submentum elongate, not clearly separated from gula; gular sutures slightly converging anteriorly. Cervical sclerites divided, anterior plate about twice as long as posterior one, each broadly curved at each end and distinctly concave in middle. Pronotum (Figs 9–10) 1.10–1.30 (1.20) times as long as wide, widest at posterior fifth; sides slightly curved, continuous with broadly curved anterior edge, so that anterior angles are absent; posterior angles subright with rounded tips; posterior edge weakly biemar-ginate; lateral margins slightly explanate and somewhat elevated, extending onto anterior edge; disc finely and densely punctate (Fig. 6), punctures usually separated by less than half a puncture diameter and bearing moderately long setae, usually decumbent but suberect in places; interspaces relatively smooth and shiny. Prosternum 1.33 times as long as mid length of procoxal cavity, slightly, evenly convex, with straight, complete notosternal sutures. Prosternal process very short, broad at base, narrowing to subacute apex. Procoxal cavities large and broadly open externally and internally, with small notch at base of notosternal suture; postcoxal (hypomeral) process very short and angulate; protrochantin reduced but externally visible, more or less trapezoidal; endopleuron short, broadly expanded at apex. Scutellum 0.8 times as long as wide, not elevated posteriorly, without sub-basal curved carina, with sides slightly converging to subtruncate apex, distinctly separated from scutum, which has median endocarina. Elytra 3.10–4.08 (3.62) times as long as wide and 3.11–4.08 (4.13) times as long as pronotum, with sides subparallel or slightly wider behind middle, outer angles more or less rounded and apices independently narrowly rounded but narrowly separated; humeri weakly developed; disc with four very weak, longitudinal costae; punctation very fine, moderately dense and confused, punctures usually separated by more than a puncture diameter and bearing a decumbent seta; interspaces moderately sculptured and somewhat shiny. Mesoventrite triangular, 0.95 times as long as wide, widest at posterior end and gradually narrowed to anterior end which is very narrow and truncate, separating the broad mesanepisternal coxal rests; anterior portion of mesoventrite with pair of longitudinal, converging ridges extending from the posterior end of each coxal rest to the apex and joined at about middle by a transverse, slightly curved ridge. Posterior edge of mesoventrite forming a short, apically acute intercoxal process. Mesocoxal cavities large, weakly impressed, almost completely contiguous, broadly closed laterally by the mesepimeron; trochantin visible, small, irregularly quadrangular. Metaventrite 1.44 times as long as wide, strongly convex, longest at midline, with discrimen extending anteriorly almost to the posterior edges of the mesocoxal cavities; posterior edges of metaventrite on either side of discrimen straight but strongly oblique, extending from posterior edge of metanepisternum on each side posteromesally to a small, cleft, mesal projection adjacent to base of discrimen. Metanepisernum 12 times as long as wide, widest anteriorly, sides narrowed to narrowly truncate apex. Metendosternite with very long stalk, paired, elongate-oval laminae about twice as long as wide, slender lateral arms extending dorsally and a pair of anterior tendons arising from the mesal edges of the laminae. Hind wing about 2.4 times long as wide; apical field 0.43 times as long as wing length, with three linear sclerites forming an irregular epsilon: anterior oblique moderately thick sclerite, more slender posterior oblique sclerite and median, slender, vein-like sclerite; another lightly pigmented triangular sclerite at the base of the others and crossing the r4 cross-vein. Radial cell about 0.10 times wing length, 3.7 times as long as wide, with curved, oblique base forming an inner posterobasal angle of about 120°; cross-vein r3 absent, r4 moderately long and straight; radio-medial loop forming a 45° angle with apex of MP 1+2 and basal portion of RP extending to about midwing. MP 1+2 strongly developed, with medial spur straight, extending to wing margin. MP 3+4 undivided and extending to wing margin, basally with short, oblique cross-vein and very short basal stub. CuP meeting AA 3 without forming wedge cell, giving rise to CuA 1+2, which is moderately long, oblique and forked to form short CuA 1 meeting MP 3+4 and a long CuA 2 extending to wing margin; AA 4 extending almost to wing margin. Anal lobe well-developed, without embayment, with only one vein (AP 3+4). Procoxa 3.1 times as long as wide, widest near base and slightly narrowed to apex, strongly projecting. Mesocoxa 1.8 times as long as wide, widest near base and narrowed apically, slightly projecting near midline. Metacoxa 3.12 times as long as wide, strongly oblique and slightly projecting near midline, with very short, curved coxal cowling, a long, sinuate line occupying almost entire width and a straight groove and internal ridge. Profemur 3.07 times as long as wide; protibia 9.2 times as long as wide; protarsi about 0.72 times as long as tibia, with segment lengths 3.43: 1.43: 1.14: 1.00: 2.57. Mesofemur 3.56 times as long as wide; mesotibia about 15.5 times as long as wide, slightly narrower at base than apex; mesotarsi about 0.84 times as long as tibia, with segment lengths 4.14: 2.00: 1.57: 1.00: 1.57. Metafemur, metatibia and metatarsi similar to those of midleg. Pretarsal claws moderately long and slender with relatively small, bisetose empodium. Abdomen with first ventrite about 0.7 times as long as second, ventrites 2–4 equal in length and 5 slightly shorter than 4 and broadly rounded; intercoxal process on ventrite 1 very short and angulate, continued internally as a median carina separating metacoxae; concealed sternite II well-developed, with median carina; abdominal tergite VII (pygidium) 1.66 times as long as wide, widest at base, with sides gradually converging to broadly rounded apex and lined with pair of sublateral struts extending almost to apex. Abdominal sternite VIII in male about as long as wide; sides subparallel basally, then converging to broadly rounded apex; tergite VIII 1.48 times as long as wide; sides subparallel at base and converging to narrowly rounded apex; disc with a narrow, longitudinal translucent area at middle. Segment IX in male 1.87 times as long as wide, widest at apical third, apex deeply emarginate to form two rounded lobes; pregenital ring broadly rounded anteriorly; posteriorly segment X not clearly separated from IX. Aedeagus normally oriented, with tegmen dorsal to penis; basale 0.80 times as long as apicale and 1.67 times as long as wide in dorsal view, with narrowly rounded base; widest at basal fourth with sides beyond this point subparallel; apex almost completely fused to apicale, except apicolaterally where there are small condyles; apicale 1.85 times as long as wide, sides subparallel basally, diverging beyond middle than converging subapically; apex deeply emarginate forming pair of broad, mesally curved, apically rounded, setose parameroids; base of apicale with pair of broad tegminal struts extending anteriorly and mesally to join at midline near base of penis. Penis about as long as tegmen, 15 times as long as wide, distinctly curved ventrally, its base with broad, median, oblique, ventral projection attached to tegminal struts. Ovipositor about 8 times as long as wide, with distinct proctigeral, paraproctal and coxital bacula; paraprocts 1.5 times as long as gonocoxites, which are 3.0 times as long as combined width; each gonocoxite subdivided into a basal lobe about twice as long as wide and lightly sclerotised with heavily sclerotised oblique baculum and apical lobe, which is 2.7 times as long as basal lobe, 10 times as long as wide, parallel-sided, densely setose and narrowly rounded at apex; apical lobes relaively widely separated and parallel; gonostylus 0.12 times as long as gonocoxite, 5 times as long as wide, slightly expanded apically and setose. Types: Holotype, ♁ “ 18.55S 149.09E QLD Mt. Spec. S3 880m 6 Dec. 1994 – 10 Jan. 1995 M. Cermak F I Trap JCU 5 m ” (ANIC #25-075148) Paratypes: QLD: Bald Mtn. Area, 3-4000 ft., via Emu Vale, 22–27.ii.1971, L. Hill (1♀, QMB); Bartle Frere, West Base (17°23’S, 145°46’E), 50m, 25.xi.1994 – 10.i.1995, flight intercept trap, Monteith & Hasenpusch (1♀, QMB); Bartle Frere, West Base (17°23’S, 145°46’E), 700m, 10.i.–31.iii.1995, flight intercept trap, Monteith & Hasenpusch (2♀♀, QMB; 1♀, GS, ANIC); Bartle Frere, West Base (17°23’S, 145°46’E), 700m, 7.iii.–5.v.1995, flight intercept traps, Monteith & Hasenpusch (4♀ ♀, QMB); Boggom, via Taroom (25°27’S, 150°08’E), 13.xi.1996, at light, G. B. Monteith (1♀, QMB); Charmillan Ck. Xing, Tully Falls Road, 950m, 8.xii.1989 – 5.i.1990, pitfall & intercept traps, Monteith, Thompson & Janetski (1♀, QMB); Davies Creek Rd., 20 km ESE Mareeba, 750m, 4–13.xii.1988, flight intercept trap, Monteith & Thompson (2♀♀, QMB); Hugh Nelson Range, 2.5 km S of Cra-ter N.P., 1100m, 5–14.xii.1988, flight intercept trap, Monteith & Thompson (3♀♀, QMB); Hughes Road, Topaz (17°26. S, 145°42’E), 650m, 6.xii.1993 – 25.ii.1994, RF intercept, Monteith, Cook, Janetzki (6♀♀, QMB); KarnakDevil’s Thumb, 8–12 km NW Mossman, Site 4, 300m, 26.xii.1989 – 15.i.1990, flt. Intercept, ANZSES Expedition (1♀, QMB); Kenny Road (17°28’S, 145°32’E), 850m, 8.xii.1990 – 5.ii.1991, flight intercept trap, Monteith & Seymour (2♀♀, QMB); Mossman Bluff Track, 5–10 km W Mossman, Site 4, 600m, 20.xii.1989 – 15.i.1990, flight intercept, Monteith, Thompson, ANZSES (1♀, QMB); Mossman Bluff Track, 5–10 km W Mossman, Site 5, 760m, 16–30.xii.1988. flight intercept, Monteith, Thompson, ANZSES (1♀, QMB); Mossman Bluff Track, 5–10 km W Mossman, Site 7, 1000m, 16–30.xii.1988, flight intercept, Monteith, Thompson, ANZSES (1♀, QMB); Mt. Edith (17°06’S, 145°37’E), GS2, 1050 m, 1.xii.1994 – 3.i.1995, Malaise trap, P. Zborowski (1♁, ANIC); Mt. Fisher (17°33’S, 145°32’E), BS2, 1150m, 4.ii–21.iii.1995, Malaise traps, P. Zborowski (1♁, ANIC); Mt. Huntley (28°08S, 152°26E), 1250m, 29–30.i.1993, G. B. Monteith (1♁, QMB); Mt. Lewis Rd., 22 km from Highway, Site 3, 1000m, 18.xii.1989 – 13.i.1990, flight intercept, Monteith, Thompson, ANZSES (1♀, QMB); Mt. Lewis Rd., 11 km from Highway, Site 1, 1000m, 18.xii.1989 – 13.i.1990, flight intercept, Monteith, Thompson, ANZSES (1♁, QMB); Mt. Massey Creek (17°37S, 145°34E), BS3, 1000m, 1.xii.1994 – 3.i.1995, flight intercept trap. JCU (West), P. Zborowski (1♁, ANIC); Mt. Misery Summit (15°52’S, 145.14’E), Site 2, 850m, 16.xii.1990 – 17.i.1991, Flight intercept, QLD Mus. & ANZSES (3♀♀, QMB,); Mt. Spec, S 3 (18°55’S, 146°09’E), 880m, 6.xii.1994 – 10.i.1995, flight intercept traps, M. Cermak (8♁♁, 7♀♀, ANIC); Mt. Spec, S 3 (18°55’S, 146°09’E), 880m, 10.i.–6.ii.1995, flight intercept traps, M. Cermak (1♀, ANIC); Mt. Spec, S3 (18°55’S, 146°09’E), 880m, 6.ii–9.iii.1995, flight intercept traps, M. Cermak (7♁♁, 12♀♀, ANIC); Mt. Spec, S2 (18°55’S, 146°10’E), 880m, 9.iii–6.iv.1995, flight intercept traps, M. Cermak (1♁, 1♀, ANIC); Paluma Dam Rd. Site 4, 750m, 17.xi–8.xii.1990, flight intercept trap, Monteith & Sey-mour (1♁, QMB); Paluma Dam Rd. Site 4, 750m, 8.xii.1990 – 5.ii.1991, flight intercept trap, Monteith & Seymour (1♀, QMB); Paluma Dam Rd. Site 5, 850m, 8.xii.1990 – 5.ii.1991, flight intercept trap, Monteith & Seymour (3♀♀, QMB); South Koombooloomba Dam, 1.5 km N Tully River Xing, 750m, 8xii.1989 – 5.i.1990, pitfall & intercept traps, Monteith, Thompson, Janetski (1♁, 12♀♀, QMB); Stone Creek (Hasenpusch) (17.28’ S, 146.01’ E), 100m, 1.x–1.xi.1995, intercept trap, J. Hasenpusch (1♀, QMB); Tower nr. The Crater NP (17°27’S, 145°29’E), 1230m, 10.i.1995 – 31.iii.1996, FIT intercept trap, Monteith & Hasenpusch (1♀, QMB); Tully, 11 km NNW, Upper Boulder Ck., 850m, 16–19.xi.1984, Cook, Monteith, Thompson (1♀, QMB); Westcott Rd., Topaz (17°24’S, 145°41’E), 680m, 6.xii.1993 – 25.ii.1994, intercept, Monteith, Cook, Janetzki (1♀, QMB); Windsor Tableland, site 1, 27.i.1988 – 10.i.1989, flight intercept, E. Schmidt & ANZSES (1♀, QMB). Distribution. The species is restricted to Queensland, with records from Mt. Huntley in Main Range National Park, close to the New South Wales border north at least to Mt. Misery near Cooktown. Biology. Nothing is known on the biology of this species. Adults are usually collected in flight intercept traps or Malaise traps., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on pages 214-217, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473

Atractocerus crassicornis Clark, 1931 (Figs 22–23) Atractocerus crassicornis Clark, 1931: 120. Type, Diagnosis. Comparison with other species in the genus is not possible due to lack of material, and differences from other Australian lymexylid genera may be found in the generic key. Specimens Examined. NT: Dorisvale HS, 29 mi. SW (14°48’S, 131°02’E), 9.viii.1968, M. Mendum (2♀♀, ANIC); Katherine, Manbulloo Station 10.vii.1929, T. G. Campbell (1♀, ANIC); Victoria River Roadhouse, 1.2 km NNW, Victoria River Highway (15°06’S, 131°18’E), 23.iii.1999, D. C. F. Rentz, P. Naskrecki (1♀, ANIC). Distribution. Known only from the Northern Territory. Biology. The type was collected at lights at night, but nothing else is known of the biology., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 225, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Clark, J. (1931) A new species of Lymexylonidae (Coleoptera) Proceedings of the Royal Society of Victoria, New Series, 43 (2), 120 - 122."]}

Melittomma Murray, 1867 (Wheeler, 1986, figs 225–267) Melittomma Murray, 1867: 314. Type species: Hylecoetus brasiliensis Laporte 1832, by original designation. Neohylecoetus Kurosawa, 1956: 80. Type species: N. javanus Kurosawa, 1956: 81 (= Hylecoetus javanicus Chevrolat, 1829; Neohylecoetus philippensis Kurosawa, 1956). Kurosawa, 1985 (synonymy). Diagnosis. According to Wheeler (1986), adults of the genus Melittomma may be distinguished from other members of the family by having large, often proximal eyes, palp organs in the male, entire or almost entire elyra, appendiculate paratergites on abdominal segment IX and small parameroids, while lacking setose pads on segment VIII in the male. Larvae may be distinguished by the presence of a heavily sclerotised abdominal apex terminating in a concavity lined by a deeply notched ridge. Adults of Australymexylon species differ in lacking palp organs and having setose pads on segment VIII in the male, while Alcestoma species lack the appendiculate paratergites on segment IX and have more elongate parameroids. Note. The genus Melittomma currently contains about 20 species worldwide, but it is not clear if all of these species share the diagnostic features mentioned above. Wheeler (1986) moved M. insulare Fairmaire (1893) and M. curtipenne Pic (1939) to the genus Protomelittomma Wheeler (= Alcestoma Fairmaire) and considered them to be synonymous, but some Melittomma types were not seen and some names were based on females only. This problem can only be addressed by those with access to the relevant types. The only true Melittomma occurring in Australia, based on male genital and pregenital features listed above, is M. pervagum, which is redescribed below., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on pages 218-219, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Murray, A. (1867) List of Coleoptera received from Old Calabar, on the west coast of Africa. Annals and Magazine of Natural History, Series 3, 22, 314 - 323. https: // doi. org / 10.1080 / 00222936708646336","Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210.","Kurosawa, Y. (1956) A new genus and the species of the family Lymexylonidae, with the additional notes on some Japanese species (Coleoptera). Bulletin of the National Science Museum of Tokyo, 39, 80 - 82.","Kurosawa, Y. (1985) Revisional notes on the family Lymexylonidae (Coleoptera) in eastern and southeastern Asia. Bulletin of the National Science Museum, Tokyo, (A), 11, 109 - 119.","Fairmaire, L. (1893) Communications. Annales de la Societe Entomologique de France Bulletin, 62, cccxxii-cccxxv.","Pic, M. (1939) Diagnoses de Coleopteres exotiques. L'Echange, Revue Linneenne, 55 (577), 31 - 32."]}

Leptonetron victoriense (Blackburn, 1891) comb. nov. (Figs 15–16, 19) Atractocerus victoriensis Blackburn, 1891: 306. Type, female, Victoria, Collection of C. French (MVM). Atractocerus tasmaniensis Lea, 1917: 143, syn. nov. Type, male, Tasmania: Triabunna (G. H. Hardy). Type I. 6740 (SAM). Fusicornis tasmaniensis (Lea, 1917), Paulus (2004). Redescription. Total length: 9.80–24.80 (15.37 ± 4.20, n = 16); body 12.60–17.32 (14.45) times as long as combined elytral width. Head 1.25 times as long as wide, more or less globular, not constricted posteriorly, without temples or neck; base weakly biemarginate dorsally, usually with median, longitudinal groove between eyes. Eye about 0.78 times as long as head width, distinctly emarginate anteriorly, protruding and finely facetted, with fine interfacetal hairs; distance across eyes dorsally is 4.88 times and ventrally 1.86 times distance between them.Antennal insertions concealed by frontal ridges which extend slightly into eye emargination. Frontoclypeal suture absent. Labrum about 0.6 times as long as wide, densely setose, with base slightly curved and sides strongly converging to narrowly rounded apex. Antennae 12-segmented in both sexes, slightly shorter than head width in male, with antennomere 2 widest, ratio of antennomere lengths 2.60: 2.20: 2.00: 1.20: 1.00: 1.00: 1.00: 1.20: 1.00: 1.20: 1.40: 2.00 and antennomere length/width ratios 1.44, 1.10, 1.11, 0.75, 0.83, 1.11, 2.00, 2.40, 2.00, 2.40, 2.80, 5.00; slightly longer than head width in female, with antennomere 3 widest, ratio of antennomere lengths 3.00: 1,83: 2.00: 1.50: 1.50: 1.33: 1.33: 1.25: 1.17: 1.00: 1.00: 1.33 and antennomere length/width ratios 2.00, 1.10, 0.75, 0.56, 0.60, 0.57, 0.67, 0.75, 0.78, 0.71, 0.75, 2.00; terminal antennomere in both sexes acute at apex. Mandible about 1.43 times as long as wide; outer edge densely clothed with long, fine setae; apex bidentate with ventral tooth somewhat smaller than dorsal one; mola and prostheca absent. Maxilla highly reduced, with short lacinia narrowly rounded at apex, longer galea broadly rounded at apex and both densely clothed with long setae; first maxillary palpomere 1.67 times as long as wide, second and third closely associated by oblique joint, the two together about 3 times as long as wide, fourth palpomere 4 times as long as wide and narrowly rounded apically; palp organ simple, with two sets of paired lobes. Labial palps approximate, apical palpomere 2.0 times as long as basal one, 2.3 times as long as wide, with narrowly rounded apex. Gular sutures parallel. Cervical sclerites very large, the combined lengths of the two lobes about equal to head width. Pronotum 1.16–1.52 (1.34) times as long as wide, widest anteriorly; sides gradually converging posteriorly, weakly sinuate, without lateral carinae; anterior edge broadly curved; anterior angles not produced, broadly rounded; posterior angles rounded; posterior edge weakly biemarginate laterally and truncate mesally; disc moderately convex, coarsely and densely punctate and clothed with fine, suberect setae; longitudinal groove (endocarina) extending from anterior fourth to posterior fifth. Prosternum about 1.5 times as long as mid length of a procoxal cavity, distinctly convex, with biconcave anterior edge; notosternal sutures straight and complete. Prosternal process short, broad and broadly truncate at apex. Procoxal cavities large and broadly open externally and internally, with notch at base of notosternal suture; postcoxal (hypomeral) process absent; protrochantin well-developed and externally visible, more or less trapezoidal; endopleuron short, broadly expanded at apex. Scutellar shield not clearly separated from remainder of exposed scutellum, slightly elongate and rounded at apex. Elytra 1.77–1.79 (1.78) times as long as greatest combined width and 1.40–1.85 (1.57) times as long as pronotum, separated for their entire lengths, but almost meeting at anterior fourth just behind scutellar shield; each elytron about 1.8 times as long as pronotum and 3.5 times as long as wide, widest at anterior fourth with outer edge sinuate beyond middle and inner edge concave beyond anterior fourth; outer elytral angles more or less rounded and apices independently broadly rounded and well separated; disc with punctation confused, dense, coarse anteriorly and fine posteriorly, each puncture bearing a short fine, inclined seta. Mesoventrite 0.68 times as long as wide, widest at anterior end and gradually narrowed posteriorly; anterior edge emarginate, without coxal rests, with weak median endocarina extending to posterior third and a small, deep pit just before short intercoxal process. Mesocoxal cavities virtually absent. Metaventrite 1.75 times as long as wide, strongly convex, without discrimen; metacoxal cavities strongly oblique and narrowly separated. Metendosternite with long, narrow stalk and paired, slender lateral arms forming an angle of about 30°. Hind wing about 4 times long as wide; apical field 0.26 times as long as wing length, with two short radial extensions; radial cell absent; radial and medial veins joined at apical fourth of wing by transverse connection straight anteriorly, curved posteriorly and probably derived from cross-vein r4 and the radiomedial loop; MP1+2 long, straight and continuing as medial spur to wing edge; medial field with three veins reaching wing edge and probably corresponding to CuA, CuP + AA 3 and AA 4. Anal lobe reduced, without embayment, with single vein (AP 3+4). Procoxa 3.00 times as long as wide, widest near base and very slightly narrowed to apex, strongly projecting; Mesocoxa 1.85 times as long as wide, widest near base and slightly narrowed apically, slightly projecting near midline. Metacoxa about 0.38 times as long as wide, strongly oblique, almost contiguous and projecting near midline, without coxal cowling, Fore legs relatively short: profemur slightly shorter than coxa, 2.4 times as long as wide; protibia about equal in length to femur, 5.3 times as long as wide, parallel-sided, not expanded at apex; protarsi combined about equal in length to tibia, with segment length ratio 3.43: 1.43: 1.14: 1.00: 2.57. Mid and hind legs very long and slender, each with femur about 3.42 times as long as wide and widest at middle; tibia 2.00 times as long as femur, about 16 times as long as wide and more or less parallel-sided; combined tarsal length about 2.00 times as long as tibia, with segment lengths 3.17: 2.22: 1.72: 1.17: 1.00. All tarsomeres simple, without ventral lobes, but densely lined with fine, erect hairs; pretarsal claws moderately long and slender, empodium with several setae. Abdomen with tergites and sternites more or less equal in length and degree of sclerotisation; first segment (sternite and tergite II) about 0.85 times as long as second, with intercoxal process absent; segments 2–6 subequal in length; ventrite 7 (sternite VIII) about 0.85 times as long as 6 and subacute at apex, tergite VIII (pygidium) slightly longer and narrowly rounded at apex. Segment IX in male about 3.0 times as long as wide, widest at posterior third, apex deeply emarginate to form two rounded lobes; pregenital ring anteriorly subacute; tergite X not separated from IX, the apex of which is narrowly rounded. Basale 1.4 times as long as wide, widest at apex, with broadly rounded base and truncate apex; parameroids 1.13 times as long as phallobase, contiguous at base, but well separated apically, each 1.7 times as long as basal width, 2.4 times as long as mid width and narrowly rounded at apex. Penis short, slender, strongly curved dorsally and more or less enclosed within phallobase. Specimens Examined. TAS: Collinsvale, 27.i.1989, N. W. Rodd (1♁, AMS); Triabunna, G. H. Hardy (SAM, Type I. 6740). VIC: Beaconsfield, 8.xii.1923, G. F. Hill (3♁♁#, ANIC); No specific locality (Type, MVM; 2 Cotypes SAM); Woori Yallock, 22.xii.1936, F. E. Wilson (3♁♁, 1♀, MVM); Wyperfield Nat. Park (35°41’S, 141°36’E), 28.xi.1997, J. & A. Skevington, S. Winterton, C. Lambkin (1♁, QMB); Yarragon, Moe River, “taken out of piles”, W. Kershaw (3♁♁, 1♀, MVM). NSW: Grafton, 12 km S, 10.i.1958, E. F. Riek (2♁♁, ANIC); Singleton, 48 km N, “Tuglo”, 31.i.1977, C. N. Smithers (1♀, AMS); Tamworth, 23 km SE, 25.xi.1982, D. S. Horning (1♁. ANIC). QLD: Brisbane, 16.x.1911, H. Hacker (1♁, QMB); Mt. Coot-tha (27°29’S, 152°57’E), 8.xi.1997, C. Lambkin (1♁, QMB); Mt. Coot-tha (27°29’S, 152°57’E), 170m, 26.x.1997, hilltoping, J. Skevington (1♁, QMB). WA: Warren River, nr. Pemberton (3♀♀, AMS). Distribution. Southern Australia from TAS to southern QLD and southern WA. Biology. Specimens from Yarragon River in Victoria were “taken out of piles” (apparently logs)., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on pages 226-227, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Blackburn, T. (1891) Further notes on Australian Coleoptera, with descriptions of new genera and species. Transactions of the Royal Society of South Australia, 14, 292 - 345.","Lea, A. M. (1917) Notes on some miscellaneous Coleoptera, with descriptions of new species-Part III. Transactions of the Royal Society of South Australia, 41, 121 - 322, pls. 12 - 15. https: // doi. org / 10.5962 / bhl. part. 15329","Paulus, H. F. (2004) Urtrea graeca nov. gen. et nov. spec., der erste Vertreter der tropischen Atractocerinae in Europa sowie eine Beschreibung von Homaloxylon aspoecki nov. spec. aus Yunnan (China) (Coleoptera, Cucujiformia, Lymexylidae, Atractocerinae nov. status). Denisia, 13, 277 - 290."]}

Raractocetus Kurosawa, 1985 Raractocetus Kurosawa, 1985: 111. Type species: Atractocerus emarginatus Laporte, 1836 (original designation). Diagnosis. Species of the genus Raractocetus differ from those of Arractocetus Kurasawa, Fuscicornis Philippi and Hymaloxylon Kurosawa and Leptonetron gen. nov. in having the eyes very large and contiguous (male) or subcontiguous (female) in frontal view. Raractocetus is distinguished from Atractocerus by the larger, shorter and more vertically oriented head, as wide as or wider than the anterior pronotal margin, more robust antennae and shorter elytra. A male from Barro Colorado Island, Panama fits the description of Atractocerus procerus Schenkling (1914) and has the head characters of Raractocetus, but A. procerus cannot be transferred to Raractocetus without examination of the type. If this specimen does belong to A. procerus, then the Neotropical species differs from both R. emarginatus and R. kreusleri in having a median groove behind the eyes and a median groove extending the entire length of the pronotum., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 227, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Kurosawa, Y. (1985) Revisional notes on the family Lymexylonidae (Coleoptera) in eastern and southeastern Asia. Bulletin of the National Science Museum, Tokyo, (A), 11, 109 - 119.","Schenkling, S. (1914) Beitrage zur Kenntnis der Lymexyloniden (Col.) I. Entomologische Mitteilungen, 3, 217 - 221. https: // doi. org / 10.1002 / mmnd. 191419140303"]}

Australymexylon fuscipennis (Lea) (Fig. 20) Hylecoetus fuscipennis Lea, 1912: 466. Type, ♀, SAM. Examined by Wheeler (1986). Diagnosis. Australymexylon fuscipennis differs from A. australe in its lighter colour, especially at the elytral bases and males with strongly serrate antenna, smaller setose pads on segment VIII and much smaller penis. Specimens examined. QLD: Bald Mtn.Area, via Emu Vale, 3-4000’, 22–27.i.1971, G. B. Monteith, etc. (2♁♁, 8♀♀, QMB); Bald Mtn. area, via Emuvale, 3-4000 ft., 22–27.i.1972, L. Hill (1♀, ANIC); Bald Mtn. Area, via Emu Vale, 3-4000;, 26–30.i.1973, T. Spencer (1♀, QMB); Bald Mtn. Area, via Emu Vale, 3500–4000’, 26–30.i.1973, I. D. Naumann (1, QMB); Brisbane, x–xi.1957, 1961, W. Haseler (4♀♀, QMB); Cooloola, Freshwater Rd., rainforest, 9.xi.1974, G. Mon- teith (1♀, QMB); Glen Alpin, 4–8.x.1964, P. Kerridge (1♀, Q! MB); Inglewood, 1.ii.1954, R. J. Turner (1♀, QMB); Middle ridge, 27.xi.1962, A. Macqueen (1♀, QMB); Mt. Moffat (24.57’S, 148.02’E), Mahogany Forest, 1000m, 26.ix–26.xi.1995, intercept, G. B. Monteith (1♀, QMB); National Park, xii.1919, H. Hacker (1♁, 2♀♀, QMB); Stan-thorpe Dr., xi.1919, H. Jarvis (1♀, QDAF); Toowoomba, R. Illidge (1♀, QMB). NSW: Arrawarra, xii.1961, C. W. Frazier (3♀♀, ANIC); Barrington House, via Salisbury, 26–28.xii.1965, B. Cantrell (2♁♁, 1♀, QMB); Barrington Tops N. P., Cobark Forest Area, 16–17.i.1996, M. J. Fletcher (1♀, ASCU); Barrington Tops N. P., Upper Williams River, 550m, 26.i.1987, R. De Keyzer (2♀♀, AMS); Bawley Point (30°30’S, 150°24’E), 30.xii.1997, D. C. F. Rentz (2♀♀, ANIC); Beaucroft, 10.i.1968, O. M. Williams (1♀, ASCU); Blue Mts. (1♀, ANIC); Bombala, Bondi State For., iii.1984, Ipsenol tra, Pinus radiata, D. M. Campbell (1♁, ASCU); Buccleuch St. For., Black Andrew, 26.i, 28. ii.2007, I. Hides (1♁, 1♀, ASCU); Chichester State For., via Dungog, 25.xii.1965, T. Weir, B. Cantrell (2♀♀, QMB); Congo, xii.1979 – i.1980, M. S. Upton (3♁♁, 2♀♀, ANIC); Copeland Tops Forest Rd., 19 km from Gloucester, 4.i.1982, E. B. Britton (1♀, ANIC); Cutler’s Pass, near, 1250 ft., i.1943, A. Musgrave (1♀, AMS); Cumberland Nat. For., 10.i.1988, light trap, D. S. Kent (1♀, ASCU); Dingo Tops Forest Park, NW of Wingham, 8.i.1984, G. Williams (1♀, ANIC); Dorrigo, W. Heron (1♀, ANIC); Durras, i.1939, G. F. Hill (1♀, ANIC); Greenwich, xii.1970, P. Zborowski (1♀, ANIC); Gundaroo Road property “Calosoma”, 4.iii.1977, B. P. Moore (1♀, ANIC); Jerilderie, 1.ii.1970, H. Burton (1♀, QMB); Jindabyne (36°25’S, 148°.37’E), 920m, 7.ii.1993, UV blacklight, A. Newton, M. Thayer (1♀, ANIC); Lansdowne, 3 km N via Taree, 3.xii.1988, wet forest complex, UV light, G. Williams (1♀, AMS); Lansdowne, 3 km N via Taree, 4.i.1989, UV light at rainforest margin, G. Williams (1♀, AMS, K517751); McCarr’s Creek, Ku-ring-gai Chase, 31.xii.1970, M lamp, D. K. McAlpine (1♁, 1♀, AMS); Middle Brother St. For, Bird Tree, 23.xi.2001, MV light, D. Britton, B. Walsh (1♀, AMS); Mt. Kaputar Plateau, Mt. Kaputar N. P., NE of Narrabri, 11.i.1989, on Acacia foliage, G. Williams (1♀, AMS); Narara, 24.xii.1949, I. Mosse-Robinson (1♀, ASCU); Narara, 2.i.1950, I. Mosse-Robinson (1♀, ASCU); Narrabri Ag. Res. Sta., 24.ii.1966, W. E. Wright (1♀, ASCU); Nepean River, xii.1922, A. Musgrave, T. Campbell (1♀, AMS); Queanbeyan, 2.7 km NE, 670m, 30.i.1980, I. F. B. Common (1♀, ANIC); Richmond, 3.iii.1993, C. Carr (1♀, ASCU); Royal Nat. Park, 28.xii.1970, MV lamp, D. K. McAlpine (1♀, AMS, K517727); Rydalmere, xii.1976, light trap, G. R. Brown (1♀, ASCU); Starr’s Creek, Lansdowne, 23.xii.1990, S. G. Watkins (2♀♀, ANIC); Tooloom Plateau, via Urbenville, 16–17.xii.1972, I. Naumann (1♀, QMB); “ Tuglo ”, 48 km N of Singleton, 1.i, 16.xii.1977, 20.xii.1980 C. N. Smithers (4♀♀, AMS, K517745–48); Upper Colo, 1.i.1979, at UV light, D. P. Carne (1♀, ANIC); Upper Horseshoe Creek, near Kyogle, 24.xi.1986, at black light, D. J. Scambler (1♀, AMS); Wauchope, 72 km W on Oxley Hwy. (31°27’S, 152°44’E), 4.i.1970, at light, Britton, Holloway, Misko (1♁, 2♀♀, ANIC); Westleigh, Sydney, 15.xii.1986, at light, G. A. Webb (1♀, ASCU); West Pennant Hills, 3.i.1984, light trap, E. E. Taylor (1♀, ASCU); West Pimble, near Sydney, 1984, D. J. Scambler (2♀♀, AMS); West Pimble, near Sydney, 22.xii.1986, at black light, D. J. Scambler (1♀, AMS); West Pimble, near Sydney, 10.xii.1987, at black light, D. J. Scambler (1♀, AMS); Wedderburn, 3 km E (34°08’S, 150°49’E), 16,21. xii.2005, at MV light, D. Britton (1♁, 6♀♀, AMS); Wedderburn, 3 km E (34°08’S, 150°49’E), 1.i.2011, at MV lamp, D. R. Britton, K. Hill (1♁, 2♀♀, AMS,); Yarras, Yarras Timber Mill, ii & xi.1964, M. V. light trap, M. Thompson (4♀♀, ASCU). VIC: Cann River, 20.i.1967, G. Monteith (4♀♀, QMB); Hall’s Gap, 31 km S, Grampians Nat. Park (37.25’S, 142.29’E), 250m, 6.ii.1997, D. C. F. Rentz, E. S. Ross (1♀, ANIC); Valencia Creek, 10 mi. N via Maffra, River Crossing, 1.i.1966, T. Weir (2♀♀, QMB). ACT: Black Mtn., 18.ii.1951, P. B. Carne (1♀, ANIC); Lyneham, Canberra, 1.1964, at light, B. P. Moore (1♁, ANIC). SA: Coorong, 25.xii.1928, A. H. Elston (1♀, AMS); Inglewood, nr. Adelaide, 24.xi.1984, G. R. Brown (1♀, ASCU); Mt. Lofty Rgs., A. H. Elston ((1♀, AMS)., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 217, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Lea, A. M. (1912) Descriptions of new species of Australian Coleoptera. Proceedings of the Linnean Society of New South Wales, 36 (3), 426 - 478, pl. 17. [1911]","Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210."]}

Atractocerinae Laporte, 1840 The Atractocerinae are unique among the Coleoptera in many respects, but notably in the extreme reduction of the elytra combined with the lack of transverse folds in the hind wings, which are almost completely exposed, lying over the abdomen. Based on the wing character alone, this subfamily was treated as a beetle suborder, Aplicalae, by King (1955); but Selander (1959) demonstrated that this wing type could be derived from that found in the lymexylid genus Melittomma. Likewise, Baehr (1976) in his detailed descriptive paper on the prothoracic morphology of “ Atractocerus sp.” commented on several structures (e.g., spinasternum, dorsal cervical sclerites, karepisternum, which “have been lost” in all other Coleoptera. Although this group appears to be derived from an ancestor well within the Polyphaga, the family Lymexylidae being part of or sister to the large superfamily Tenebrionoidea, the adult has certainly undergone major changes in body structure. The ideas on relationships of atractocerines to other lymexylids have not really changed from those of Wheeler (1986), with the Palaearctic genus Lymexylon Fabricius as the sister group. It may be time, however, for a worldwide analysis to be performed based on adult and larval morphology and molecular data. This, however, is beyond the scope of the present study. As noted in the Introduction, the subfamily now includes six extant genera: Arractocetus Kurosawa (1985), with ten species from Japan, Taiwan, Southeast Asia, the Philippines, India, Indonesia, New Guinea and the Moluccas; Atractocerus de Beauvois (1801), with eight species from Central and South America, the West Indies, tropical Africa, Madagascar, India, Sri Lanka, Sumatra and southern China; Fusicornis Philippi (1866), with a single species from Chile; Hymaloxylon Kurosawa (1985), with two species from India, Nepal and western China; Raractocetus Kurasawa (1985), with two species from India, the Philippines, Taiwan, Southeast Asia, Indonesia and Australia (introduced with Eucalyptus into New Zealand and Chile); and Urtea Paulus (2004) with one species from northern Greece. In addition, the following fossil Atractocerinae have been reported: Cretoatractocerus grimaldii WolfSchwenninger (2011), the earliest member of the group from the Lower Cretaceous (Upper Aptian) Crato Formation of Brazil, plus Vetatractocerus burmiticus Yamamoto,(2019), Raractocetus extinctus Yamamoto (2019) and R. fossilis Yamamoto (2019) from the mid-Cretaceous Burmese amber (earliest Cenomanian), R. balticus Yamamoto (2019) from mid-Eocene Baltic amber and Atractocerus sp. from Miocene Dominican amber (Yamamoto, 2019)., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 223, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Laporte, F. L. N. C. de (1840) Histoire naturelle des insectes Coleopteres; avec une introduction renfermant l'anatomie et la physiologie des animaux articules, par M. Brulle. Tome deuxieme. Histoire naturelle des animaux articules, annelides, crustaces, arachnides, myriapodes et insectes. Tome Troisieme. P. Duymenil, Paris, 563 pp., 38 pls.","King, E. W. (1955) The phylogenetic position of Atractocerus Palis. The Coleopterists Bulletin, 5, 65 - 74.","Selander, R. B. (1959) Additional remarks on wing structure in Atractocerus. The Coleopterists Bulletin, 13, 1 - 5.","Baehr, M. (1976) Das Prothorakalskelett von Atractocerus (Lymexylonidae) und seine Bedeutung fur die Phylogenie der Cole- opteren, besonders der Polyphagen (Insecta: Coleoptera) Zoomorphologie, 85, 39 - 58. https: // doi. org / 10.1007 / BF 00996064","Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210.","Kurosawa, Y. (1985) Revisional notes on the family Lymexylonidae (Coleoptera) in eastern and southeastern Asia. Bulletin of the National Science Museum, Tokyo, (A), 11, 109 - 119.","Philippi, R. A. (1866) Einige Insekten von Chile. Stettiner Entomologische Zeitung, 27, 109 - 117.","Paulus, H. F. (2004) Urtrea graeca nov. gen. et nov. spec., der erste Vertreter der tropischen Atractocerinae in Europa sowie eine Beschreibung von Homaloxylon aspoecki nov. spec. aus Yunnan (China) (Coleoptera, Cucujiformia, Lymexylidae, Atractocerinae nov. status). Denisia, 13, 277 - 290.","Yamamoto, S. (2019) Fossil evidence of elytral reduction in ship-timber beetles. Nature, Scientific Reports, 9 (4938), 1 - 10. https: // doi. org / 10.1038 / s 41598 - 019 - 41310 - 1"]}

Australymexylon australe (Erichson) (Fig. 21) Lymexulon australis Erichson, 1842: 147. Type, ♀, Museum für Naturkunde, Berlin. Examined by Wheeler (1986). Hylecoetus linearis Lea, 1894: 803. Type, SAM. Examined by Wheeler (1986). Lymexylon adelaidae Blackburn, 1898: 34. Type, SAM. Examined by Wheeler (1986). Diagnosis. Australymexylon australe differs from A. fuscipennis in its much darker colour, weakly serrate male antenna, males with larger setose pads on segment VIII and penis about as long as basale and apicale combined. Specimens examined. TAS: Bronte Park, 12 km NNE (42°02’S, 146°33’E). 20.i.1983, J. Cardale (3♁♁, ANIC); Liffey Valley, 11.ii.1987, S. Fearn (1, ANIC); Mt. Field Nat. Park (42°41’S, 146°43’E), 160–240m, 30.i–4.ii.1980, on bark surface, J. Lawrence, T. Weir (1♀, ANIC); Mt. Nelson, Hobart, 100m, 6.ii.1982, at light, G. Bornemissza (1♀, ANIC); St. Patrick’s R., 5.ii.1914, A. H. Elston (1♁, 1♀, AMS); VIC: Cape Conran Coastal Park, 24 km SE Orbost (37°47’S, 148°44’E), 12.i–13.ii.2005, C. Lambkin. N. Starick (1♁?, ANIC); Halls Gap, 3 km S, Grampians Nat. Park (37°25’S, 142°29’E), 250m, 6.ii./1997, D. C. F. Rentz, D., E. S. Ross (1♀, ANIC); Little Boys Camp area, 25.i.1940 (1♀, ANIC); Lorne, 25.i.1959, B. P. Moore (1?, ANIC); North Cascade Camp, 25.i.1940, Eucalyptus regnans (1♀ GS, ANIC); NSW: Blacktown, Sydney, 17.xi.1980, R. De Keyzer (1♀, QMB); Gloucester Tops, 17.i.1979, C. N. Smithers (1♀. AMS); Lord Howe Island, Cowan Track, 400m, before end on right, 33°39’14”S, 151°10’’19”E), 3.xii.2000, Flight int. trap, SLASS1.int27, S. A. Lassau (1♁, AMS)., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 217, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210."]}

Australymexylon Wheeler, 1986 (Wheeler, 1986, figs 188–215) Australymexylon Wheeler, 1986: 160. Type species: Lymexylon australis Erichson, 1842, by original designation. Diagnosis. Australymexylon species differ from other Melittommatinae in lacking a maxillary palp organ in both sexes, base of pronotum with pair of small deep impressions, elytra lying flat over abdomen (with incomplete interlocking mechanism), scutellum not separated from scutum, and males with paired setose pads on sternite VIII., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 217, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210."]}

Atractocerus Palisot de Beauvois, 1801 Atractocerus Palisot de Beauvois, 1801: 3. Type species: Atractocerus necydaloides Palisot de Beauvois (monotypy) (= Necydalis brevicornis Linnaeus, 1766). Diagnosis. Species of Atractocerus in the strict sense are characterised by the large, elongate eyes, elongate, horizontal head, which is narrower than the pronotum, and very short elytra. Distribution. At least ten species of Atractocerus have been described from California to Brazil, Africa, Madagascar, India, Sri Lanka, Southeast Asia, Indonesia and northwestern Australia, but it is not certain that all of these species are congeneric. According to Kurasawa (1985), A. brevicornis (Linnaeus), A. reversus Walker and A. brasiliensis Lepeletier & Audinet-Serville belong to this genus, and A. crassicornis Clark should be added to this list., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 225, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473

Melittommatinae, Wheeler, 1986 The subfamily Melittommatinae was proposed by Wheeler (1986) for those members of the family Lymexylidae in which the elytra completely conceal the abdomen or expose at most two abdominal terga, and the larvae have a heavily sclerotised, truncate tergite IX. The largest genus in this group is Melittomma, with about 20 species primarily in tropical and subtropical regions of both Western and Eastern Hemispheres. Other currently recognised genera include Melittommopsis Lane (1955), with at least four species from South America, the monotypic Protomelittomma Wheeler (1986) from the Seychelles and Madagascar and the Australian Australymexylon Wheeler (1986). The remaining fossil species, Ponomarenkylon alexandri Kirejtshuk (2008), an apparent Melittommatine from Baltic amber differ from other lymexylids in having: 1) very large eyes, (2) unusually short prothorax 3) filiform antennae and 4) the apparent connation of the first three abdominal ventrites (straight margins separating the first three ventrites and curved margins separating 3 from 4 and 4 from 5). Only a re-examination of the holotype will support or refute the author’s hypothesis. More than 20 years ago, the genus, Alcestoma Fairmaire (1895) was transferred to Lymexylidae from Melandryidae in a supplement to a paper dealing with a portion of the beetle fauna of the Prioksko-Terrasny Biosphere Reserve in the Moscow Region, Russia (Nikitsky et al. 1998). Fortunately, Dr. Nikitsky photographed the types in his possession and sent me a habitus figure plus several detailed images. Based on these images, there is little doubt that this species is congeneric with Melittomma insulare Fairmaire (1893) from the Seychelles, which was made the type species of Protomelittomma Wheeler (1986)., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 213, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Wheeler, Q. D. (1986) Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bulletin of the American Museum of Natural History, 183, 113 - 210.","Lane, F. (1955) Novos generos e especies de Coleoptera Lymexylonidae e notas sobre Melittomma Murray, 1867. Papeis Avulsos do Departamento de Zoologia, Sao Paulo, 12, 141 - 164.","Kirejtshuk, A. G. (2008) A new genus and species of timber beetle (Coleoptera: Lymexylidae) from Baltic amber. Paleontological Journal, 42 (1), 63 - 65.","Fairmaire, L. (1895) Materiaux pour la faune coleopterique de la region Malgache. Annales de la Societe Entomologique de Belgique, 29, 336 - 398.","Nikitsky, N. B., Semenov, V. B. & Dolgin, M. M. (1998) The beetles of the Prioksko-Terrasny Biosphere Reserve Xylobionts, Mycetobionts, and Scarabaeidae (with a review of the Moscow Region fauna of these groups). Supplement 1 (with remarks on nomenclature and systematics of some Melandryidae of the world fauna) Sbornik Trudov Zoologicheskogo Muzeya M. G. U. (Archives of the Zoological Museum of Moscow State University), 36 (Supplement 1), 1 - 63. [in Russian with English summary]","Fairmaire, L. (1893) Communications. Annales de la Societe Entomologique de France Bulletin, 62, cccxxii-cccxxv."]}

Arractocetus Kurosawa, 1985 Arractocetus Kurosawa, 1985: 112. Type species: Atractocerus nipponicus Nakane, 1985 (original designation). Diagnosis. Species of Arractocetus are distinguished from other atractocerines by the head, which is declined, flattened above and abruptly declined posteriorly, the head and pronotum densely clothed with hairs, the pronotum subquadrate with a distinct median groove, and the hind wings often darkened or metallic. Eyes in this genus are very finely facetted, with individual facets flattened rather than convex. Distrubution. Arractocetus is primarily an Oriental genus, distributed from India to Southeast Asia, Indonesia, New Guinea, northern Australia, the Philippines, Taiwan and southern Japan. Biology. As noted by Kurosawa (1985), the eye type in this genus is indicative of diurnal habits and the colour patterns appear to mimic those of various Hymenoptera., Published as part of Lawrence, John F., 2020, The Australian Lymexylidae (Coleoptera: Tenerionoidea) with one new genus and two genera new to Australia, pp. 211-238 in Zootaxa 4895 (2) on page 223, DOI: 10.11646/zootaxa.4895.2.3, http://zenodo.org/record/4322473, {"references":["Kurosawa, Y. (1985) Revisional notes on the family Lymexylonidae (Coleoptera) in eastern and southeastern Asia. Bulletin of the National Science Museum, Tokyo, (A), 11, 109 - 119."]}