Australian Beetles Volume 2: Archostemata, Myxophaga, Adephaga, Polyphaga (part)

By Adam Slipinski and John F. Lawrence

This three-volume series represents a comprehensive treatment of the beetles of Australia, a relatively under-studied fauna that includes many unusual and unique lineages found nowhere else on Earth.

Volume 2 contains 36 chapters, providing critical information and identification keys to the genera of the Australian beetle families included in suborders Archostemata, Myxophaga, Adephaga and several groups of Polyphaga (Scirtoidea, Hydrophiloidea, Scarabaeoidea, Buprestoidea and Tenebrionidae). Each chapter is richly illustrated in black and white drawings and photographs. The book also includes colour habitus figures for about 1000 Australian beetle genera and subgenera belonging to the families treated in this volume.

This volume is a truly international collaborative effort, as the chapters have been written by 23 contributors from Australia, China, Czech Republic, Germany, Italy, Poland and USA.

Certificate of Commendation, The Royal Zoological Society of NSW 2020 Whitley Awards: Taxonomy

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Nov 1, 2019
• ISBN: 9781486311408

Book preview

1. ARCHOSTEMATA KOLBE, 1908

John F. Lawrence and Hermes E. Escalona

The genera Cupes Latreille and Omma Newman were included in the suborder Adephaga by Ganglbauer (1903), Kolbe (1901, 1903) and Lameere (1903), based primarily on wing venation, while Kolbe (1908) excluded cupedids from Adephaga and placed them in a separate group, Archostemata, within what is now known as Polyphaga. Forbes (1926) considered Archostemata to be one of the three suborders of beetles, containing Cupedidae Laporte (including Ommatidae Sharp & Muir) and Micromalthidae Barber, based on wing folding, while by Böving & Craighead (1931) and Peyerimhoff (1933) supported this based on larval evidence. This classification was used in Crowson’s (1955) classic work on the order. In a later paper, Crowson (1962) made the first attempt to integrate the work of paleontologists, such as A. Handlirsch, R. J. Tillyard, B. B. Rohdendorf and A. B. Martynov, on cupedid-like fossils with knowledge of the extant archostematan fauna known at that time. Ponomarenko (1964, 1966, 1968, 1969) published the first major work dealing with Archostemata, which was defined in a broad sense to include virtually all known Permian and Mesozoic fossils sufficiently complete to be described (11 families, 74 genera and 186 species) but lacking the distinctive features of either Polyphaga or Adephaga. The Lower Permian Tshekardocoleidae and several Upper Permian groups were placed in Protocoleoptera and Archecoleoptera, respectively, by Crowson (1975), but were considered to form a coleopterid stem group by Kukalová-Peck (1991). This was followed by Beutel (1997) and Beutel & Friedrich (2008), who added the Triassic Triadocupedidae to this paraphyletic stem group. Within the remaining Archostemata, relationships are still not clearly understood, particularly among the taxa with non-clathrate elytra (without window-punctures) such as Jurodidae, Ademosynidae, Schizophoridae and Catiniidae, and their possible affinities with the other three suborders, Adephaga, Myxophaga and Polyphaga. Numerous publications have appeared in the past 25 years on fossil Archostemata primarily from localities in Kazakhstan, Russia, Mongolia and northern China but also on those in the UK, USA, Spain, Australia and Argentina; included are Ashman et al. (2015); Jarzembowski & Wang (2016); Jarzembowski et al. (2012, 2013a, 2013b, 2015); Kirejtshuk (2005); Kirejtshuk et al. (2010a, 2010b, 2016); Lubkin (2003); Martin (2010); Martins-Neto et al. (2006); Ponomarenko (1994, 1997, 2000, 2004, 2006); Ponomarenko & Martinez-Delclòs (2000); Ponomarenko & Ren (2010); Ponomarenko et al. (2014); Ren & Tan (2006); Ren et al. (2006); Soriano & Delclòs (2006); Tan & Ren (2006a, 2006b, 2006c, 2007, 2009); Tan et al. (2007a); Tan et al. (2005); Tan et al. (2006a, 2006b, 2007b, 2007c); Tan et al. (2006c, 2013, 2012) and Yan et al. (2014).

In the study of Archostemata by Kirejtshuk et al. (2016), virtually all archostematans with clathrate or reticulate elytra are considered to belong to the single family Cupedidae, extending back to the Triassic, but without an adequate phylogenetic analysis and additional support from other members of the paleontological community, we continue to recognise the families Ommatidae and Cupedidae. Recent discoveries of Cupedidae in the Cretaceous amber of Myanmar may be found in the papers by Jarzembowski et al. (2017a, 2017b, 2017c).

Some important characters defining this suborder may be found in extant members of the Ommatidae and Cupedidae, but not in highly derived or little known groups like Micromalthidae (Beutel & Hörnschemeyer 2002a), Crowsoniellidae (Ge et al. 2011) or the doubtfully included Jurodidae (Yan et al. 2014). These include: (1) lack of cervical sclerites (also in Adephaga and Myxophaga); (2) external propleuron extending well in front of procoxal cavities (also in Adephaga, reduced in Myxophaga); (3) free, external protrochantin; (4) membranous joint between mesothorax and metathorax (also in some Polyphaga); (5) mesocoxal cavities broadly closed laterally by metanepisternum (narrowly so in a few Polyphaga); (6) well developed and exposed metatrochantins; (7) hind wing without a radial cell formed by meeting of RA1+2 and RA3+4 (also in Adephaga and Myxophaga); (8) hind wing with cross-veins between RP and MP1+2 forming an oblongum cell (also in Adephaga and Myxophaga), (9) hind wing with major fold crossing MP1+2 forming a sharp hinge (also in Adephaga and Myxophaga); (10) larva with 6-segmented legs and paired pretarsal claws (also in Adephaga); (11) larva with ligular sclerome.

There are relatively few extant species of Archostemata: the family Ommatidae contains four Australian species of Omma Newman and two species of Tetraphalerus Waterhouse, occurring primarily in Argentina and Bolivia, and the Cupedidae includes nine genera and 31 species occurring in North America, South America, the eastern Palaearctric and Oriental regions, the East Indies, New Guinea, Australia, New Caledonia, East Africa, South Africa and Madagascar (Neboiss 1984; Lawrence 1999; Hörnschemeyer 2009, 2016; Hörnschemeyer & Beutel 2016; Hörnschemeyer & Yavorskaya 2016). The group has become extinct in Europe, but several European species are known from the early Tertiary (Kirejtshuk 2005; Kirejtshuk et al. 2010a). Several morphological papers have been produced on both larvae (Beutel & Hörnschemeyer 2002a, 2002b; Hörnschemeyer et al. 2002; Grebennikov 2004; Yavorskaya et al. 2015) and adults (Baehr 1975; Hörnschemeyer et al. 2006; Beutel et al. 2008; Friedrich et al. 2009; Hünefeld et al. 2011). General summaries of extant Archostemata may be found in Hörnschemeyer (2016), Hörnschemeyer & Beutel (2016) and Hörnschemeyer & Yavorskaya (2016).

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Tan J, Ren D (2006b) New fossil Priacmini (Insecta: Coleoptera: Archostemata: Cupedidae) from the Jehol Biota of China. Journal of Natural History 40(47–48), 2653–2661. doi:10.1080/00222930601121445

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Tan J, Ren D, Shih C (2006b) New cupedids from the middle Jurassic of Inner Mongolia, China (Coleoptera: Archostemata). Annales Zoologici 56(1), 1–6.

Tan J, Ren D, Shih C, Ge S (2006c) New fossil beetles of the family Ommatidae (Coleoptera: Archostemata) from the Jehol Biota of China. Acta Geologica Sinica 80(4), 474–485.

Tan J, Huang D, Ren D (2007a) First record of fossil Mesocupes from China (Coleoptera: Archostemata: Cupedidae). Acta Geologica Sinica 81(5), 688–696. doi:10.1111/j.1755-6724.2007.tb00993.x

Tan J, Ren D, Shih C (2007b) New beetles (Insecta: Coleoptera: Archostemata) from the Late Mesozoic of North China. Annales Zoologici 57(2), 231–247.

Tan J, Ren D, Shih C (2007c) New ommatids of Ommatinae (Coleoptera: Archostemata: Ommatidae) from the Yixian Formation of western Liaoning, China. Progress in Natural Science 17(7), 803–811. doi:10.1080/10002007088537476

Tan J, Wang Y, Ren D, Yang X (2012) New fossil species of ommatids (Coleoptera: Archostemata) from the middle Mesozoic of China illuminating the phylogeny of Ommatidae. BMC Evolutionary Biology 12(113), 1–19.

Tan J, Ren D, Shih DC, Yang X (2013) New schizophorid fossils from China and possible evolutionary scenarios for Jurassic archostematan beetles. Journal of Systematic Palaeontology 11(1), 47–62. doi:10.1080/14772019.2011.637515

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2. OMMATIDAE SHARP AND MUIR, 1912

John F. Lawrence and Hermes E. Escalona

Fig. 2.1. Omma mastersi Macleay.

Introduction. Crowson (1962) recognised the cupedid subfamily Ommatinae (as Ommadinae of Sharp & Muir), including the genera Omma Newman and Tetraphalerus Waterhouse, each in its own tribe, and described the first fossil Omma from the Lower Jurassic of England. These were distinguished from the Cupedinae by short antennae, tridentate, vertically oriented mandibular apices, contiguous procoxae, simple fourth tarsomere, and non-overlapping abdominal ventrites. Ponomarenko (1964, 1966, 1968, 1969) considered these two genera, along with five other extinct taxa, to belong to the cupedid subfamily Ommatinae, and described eight more Tetraphalerus and five more Omma from the Lower Jurassic to the Lower Cretaceous of Russia, Kazakhstan and Kyrgyzstan. Since that time numerous other fossil taxa have been added to the Ommatinae sensu Ponomarenko from various deposits in Russia, Mongolia and China, as well as the UK, Spain and Australia. Lawrence (1999) suggested that several archostematan taxa be elevated to a higher rank in line with the current treatment of extant taxa (Triadocupedidae; Cupedidae with Mesocupedinae, Priacminae and Cupedinae; and Ommatidae with Lithocupedinae, Notocupedinae, Ommatinae and Tetraphalerinae). At least some palaeontologists (Kirejtshuk et al. 2010a, 2010b, 2016), however, consider the Ommatidae to be a subfamily of a broadly defined Cupedidae, which extends back to the Triassic. According to Tan et al. (2012) the group includes 13 genera and more than 100 species, but the relationships among extant and extinct species remain obscure. These workers presented a cladogram based on 27 characters in which the genera Notocupoides and Eurydictyon form a clade (unfortunately given an invalid name Pronotocupedini) sister to the remaining taxa, Notocupedini sister to the remaining four clades and Lithocupedini sister to (Brochocoleini (Ommatini and Tetraphalerini)). In the past 20 years, ommatids have been described from various sites in the UK (Jarzembowski et al. 2013a, 2013b; Ponomarenko 2006), Spain (Soriano & Delclòs 2006), China (Cai & Huang 2016; Jarzembowski et al. 2013a, 2013b; Ren et al. 2006; Tan et al. 2005, 2006, 2007, 2012), Mongolia (Ponomarenko 1994, 1997), Transbaikalia, Russia (Ponomarenko 2000), Myanmar (Jarzembowski & Wang 2016; Jarzembowski et al. 2017), and Australia (Ashman et al. 2015; Martin 2010).

Extant members of the family Ommatidae occur only in Australia (Omma) and in parts of southern South America (Tetraphalerus). The two groups differ in many respects and are placed in separate subfamilies. The South American Tetraphalerus Waterhouse is known from the relatively arid parts of northern Argentina and Bolivia but also occurs in southern Brazil. Species of Tetraphalerus differ from those of Omma in having a narrowly elongate, strongly flattened body almost entirely covered with apically flattened tubercles, each bearing a seta, head with sharply angulate temples and ventral antennal grooves, antennomeres 2–11 glabrous, prothorax with distinct lateral carinae, propleuron completely separated from sternum and extending almost to anterior edge of prothorax, and aedeagus with distinct phallobase and free parameres (Monrós & Monrós 1952; Crowson 1962; Vulcano & Pereira 1975; Beutel et al. 2007; Hünefeld et al. 2011). There are four species of Omma Newman in Australia. Omma stanleyi Newman occurs along the eastern coast but does not extend into the wet tropics, O. mastersi Macleay occurs mainly in the central and southern part of the continent, and O. rutherfordi Lawrence and O. sagitta Neboiss are southern and south-western in distribution (Hörnschemeyer & Beutel 2016).

Biology. An adult of Omma stanleyi was collected under bark of a Eucalyptus log in southern QLD, and it is likely that larvae of this species feeds in dead wood. An adult of O. mastersi was collected in southern QLD actively running on the trunk of a dead tree in full sun and its colour pattern was very similar to a clerid beetle (Stigmatium gilberti White) and a mutilid wasp (Ephutomorpha sp.). This is suggestive of a possible mimicry ring. An interesting parallel has been found in some fossil Zygadenia Handlirsch (Notocupedinae) from the Barremian (Early Cretaceous) of England; these beetles have elytral patterns similar to those in Omma mastersi and relatives and are likely to have been day-active (Jarzembowski et al. 2015). Males of O. rutherfordi were collected in flight-intercept traps at two sites in SA with deep sand and mixed mallee and the grass Triodia, while a female was collected in a swimming pool filter north of the Stirling Range in southern WA. The single known larva, which could belong either to O. sagitta or O. rutherfordi, was found in soil at Red Hill in southern WA and was likely feeding on underground roots or wood. It has been described by Lawrence (1999) and Grebennikov (2004).

Characteristics. Adults. 6–26 mm in length, and 2.4–3 times as long as wide, slightly flattened dorsally and distinctly so ventrally, with sides of pronotum and elytra slightly, independently rounded. Colour uniformly dark brown to black; upper surfaces tuberculate, (tubercles spine-like in Omma rutherfordi), and clothed with short, slightly thickened, decumbent setae in O. stanleyi, with flattened, ribbed scales which differ in colour and form a distinct pattern in other species of Omma (interspersed with stout, erect spines in O. rutherfordi). Head prognathous, always constricted posteriorly to form a neck, but without distinct temples in O. rutherfordi. Eyes entire and very finely facetted, without interfacetal setae. Antennal insertions lateral, barely concealed by frontal ridges; subantennal groove absent. Frontoclypeal suture absent. Labrum fused to clypeus. Antennae usually short, rarely exceeding posterior edge of pronotum, 11-segmented and filiform. Mandible tridentate with vertically aligned teeth, without mola or prostheca; its dorsal surface near base with setose cavity. Maxilla highly reduced and partly concealed by mentum, galea and lacinia slender, hyaline and setose, the latter without uncus. Apical maxillary palpomere expanded and securiform with small cavity bearing sensilla near outer edge of upper surface. Mentum about as long as wide, concealing small, rounded ligula and palpal insertions; apical labial palpomere expanded and securiform, with a cavity similar to that on the apical maxillary palpomere. Gular sutures well separated; gula longer than wide. Corpotentorium weakly developed or absent. Cervical sclerites absent. Prothorax usually distinctly narrower than distance across elytral humeri (less so in male O. rutherfordi), widest anteriorly with sides slightly sinuate; lateral pronotal carinae usually absent (irregular in O. rutherfordi). Prosternum in front of coxae moderately long, partly or completely fused to pleuron (pleurosternal suture incomplete or absent), without paired crural impressions. Prosternal process incomplete, narrowed apically and acute at apex. Notopleural suture complete to anterior edge, never joined by pleurosternal suture. Procoxae subglobular, not strongly projecting, without concealed lateral extensions, without or with reduced coxal plates; trochantin large, well sclerotised, broadly exposed and abutting sternum and pleuron. Procoxal cavities slightly transverse, contiguous, broadly open externally and internally. Scutellar shield flat, not elevated above adjacent sections of elytra. Elytra complete with independently rounded to acute apices and ten rows of circular or transversely oval window punctures; epipleura moderately broad and complete. Mesoventrite at middle with acute anterior projection separating paired procoxal rests extending onto mesanepisterna, mesoventral cavity absent; discrimen moderately long and transverse (mesokatepisternal) suture incomplete (not always visible externally). Mesoventral process divided into two short, acute processes. Mesocoxae more or less globular, not or slightly projecting, without or with short plates and broadly exposed trochantins. Mesocoxal cavities slightly transverse, contiguous, partly closed laterally by mesepimera and metanepisterna; meso- and metathoracic joint within coxal cavities membranous. Metaventrite strongly narrowed anteriorly; discrimen present; transverse (metakatepisternal) suture well developed. Metanepisternum completely exposed, widest near anterior end; exposed portion of metepimeron narrowly elongate and widest at posterior end; metatrochantins exposed. Metacoxae strongly transverse with subconical mesal projections, contiguous, extending laterally to meet elytra, with weak coxal plates. Metendosternite with a moderately long, slender stalk, short, broad ventrolateral processes, moderately long and distally slender lateral arms, moderately long, apically emarginate anterior process with tendons moderately widely separated on either side of emargination. Hind wing almost always well developed and moderately broad (reduced in males of O. rutherfordi), with very short bending zone near apex of radial bar (anterior wing strut); pterostigma absent; two cross-veins join RA1+2 and RA3+4 to form brachial cell; RP complete almost to wing base, joined to RA3+4 by four cross-veins; RP and MP joined by two cross-veins forming the oblongum cell. MP1+2 with very short bending zone or hinge and a very short, straight medial spur; medial field with four free veins and no medial fleck; wedge cell well developed and apically truncate; anal lobe well developed, without embayment; AP3+4 forked. Legs short and slender; tibia subequal in length to femur; protibia with small setose excavation at basal third in males (except O. mastersi); tibial spurs well developed, usually unequal and paired on all legs. Tarsi 5–5–5; tarsomeres without ventral lobes; pretarsal claws simple; empodium bisetose. Abdomen with five ventrites, which are flattened, more or less connate, separated by very narrow grooves and all on the same plane, with edges abutting. Functional spiracles on abdominal segments I–VIII. Anterior edge of segment IX in male subtruncate; apex broadly rounded; tergite IX apically rounded; segment X free, forming a cylindrical protrusion bearing the anus. Aedeagus of modified trilobate type, with parameres completely fused to phallobase and partially fused together, each with a mesal notch near apex. Testes of the tubular type in Tetraphalerus (Sarmiento 1969; Hünefeld et al. 2011) but not yet examined in Omma. Sternite VIII in female with long, articulated spiculum ventrale. Ovipositor narrowly elongate, lightly pigmented except for baculi; paraprocts much longer than gonocoxites, each of which with well developed apical gonostylus.

Larvae. Elongate, slightly flattened, widest at level of mesothorax and gradually narrowed posteriorly to apex of tergum IX. Head prognathous and slightly retracted; posterior edge of head capsule deeply emarginate; epicranial stem absent and frontal arms not apparent; endocarina thick and V-shaped with each arm forked at apex. Stemmata, four on each side, well separated. Frontoclypeal suture indistinct or absent. Labrum free, strongly transverse. Antennae moderately long, 4-segmented, with distinct antennifer; sensorium on antennomere 3 about half as long as antennomere 4. Mandibles tridentate, with large, slightly asymmetrical and irregular molae, which lack asperities or transverse ridges; prostheca absent. Ventral mouthparts retracted forming maxillolabial complex, with well developed maxillary articulating area. Galea articulated and palpiform, lacinia fixed and apically rounded with several stout spines and setae; maxillary palp 3-segmented with distinct palpifer. Labium consisting of prementum and postmentum, the former with a well developed ligular sclerome; labial palps 2-segmented and widely separated. Hypostomal rods long, converging for anterior third, then diverging and extending almost to posterior edge of head capsule; ventral epicranial ridges absent. Gular region reduced. Meso-and metatergum with strongly transverse ambulatory ampulla; thoracic venter without distinct armature. Legs reduced, 6-segmented, with two pretarsal claws; coxae widely separated. Abdomen with transverse tergal ampullae on first three segments; ampullae paired or reduced on posterior segments. Tergum IX subequal in length to VIII, apically rounded, without armature; sternite IX exposed, simple, not distinctly separated from segment X, which is posteriorly oriented. Anal opening Y-shaped with dorsal and paired ventrolateral lobes. Spiracles annular, with closing apparatus.

Classification of the Australian genera

Subfamily Ommatinae Sharp and Muir, 1912

Omma Newman, 1839 (Pl. 14F–J, M)

Type species. Omma stanleyi Newman, 1839.

Australian species. Four

Distribution. Omma stanleyi occurs in eastern Australia from VIC and SA to central QLD. O. mastersi Malceay is known from NSW and QLD but usually occurs inland from the coast. O. rutherfordi Lawrence is known from the mallee of SA and in the Stirling Range, WA, and O. sagitta Neboiss has been recorded from south-western WA and southern SA.

References. Neboiss (1968, 1989); Lawrence (1999).

Key to the species

1. Vestiture consisting of slender, recumbent setae ( Fig. 2.4 ), which are yellowish-brown in colour; elytral window-punctures irregularly transversely oval ( Fig. 2.4 , 2.6 ); length greater than 12 mm (Pl. 14F–G); eastern Australia……………………………… O. stanleyi Newman

–Vestiture including scale-like setae ( Fig. 2.2 – 2.3 ), which may be yellowish-brown or white in colour, sometimes mixed with spine-like processes; elytral window-punctures circular ( Fig. 2.3 , 2.5 ); length less than 12 mm……………………………… 2

2(1). Body slender; antennae longer, usually extending beyond basal third of elytra (Pl. 14M); head behind eyes gradually narrowed; elytral apices independently subacute; vestiture consisting of both scale-like setae and stout, erect, spine-like projections ( Fig. 2.2 ); hind wings in male highly reduced; SA and WA……………………………… O. rutherfordi Lawrence

–Body robust; antennae shorter, extending at most to basal third of pronotum (Pl. 14H–J); head behind eyes abruptly narrowed; elytral apices independently rounded; vestiture consisting of scale-like setae, without spines ( Fig. 2.3 ); hind wings well developed in both sexes……………………………… 3

3(2). Body shorter and broader, elytra less than twice as long as wide; protibia without setose excavation at basal third in males; apical third of elytra with white scales bordering suture ( Fig. 2.1 ; Pl. 14I–J); eastern and central Australia……………………………… O. mastersi Macleay

–Body more elongate, elytra more than twice as long as wide; protibia with clear setose excavation at basal third in males; apical third of elytra without white scales bordering suture (Pl. 14H); WA……………………………… O. sagitta Neboiss

Figs 2.2–2.6. Australian Ommatidae. 2, Omma rutherfordi Lawrence, portion of elytron; 3, O. mastersi Macleay, portion of elytron; 4, O. stanleyi Newman, portion of elytron; 5, Omma mastersi Macleay, elytral window puncture; 6, O. stanleyi Newman, elytral window puncture.

References

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Beutel RG, Ge SQ, Hörnschemeyer T (2007) On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics 23, 1–29.

Cai C, Huang D (2016) Omma daxishanense sp. nov., a fossil representative of an extant Australian endemic genus recorded from the Late Jurassic of China (Coleoptera: Ommatidae). Alcheringa: An Australasian Journal of Palaeontology 3, 1–7.

Crowson RA (1962) Observations on the beetle family Cupedidae, with descriptions of two new forms and a key to the recent general. Annals & Magazine of Natural History 13(51), 147–157. doi:10.1080/00222936208651227

Grebennikov VV (2004) Review of larval morphology of beetles of the suborder Archostemata (Insecta: Coleoptera), including first-instar chaetotaxy. European Journal of Entomology 101, 273–292. doi:10.14411/eje.2004.038

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Hünefeld F, Marvaldi SE, Nüller B, Lawrence JF, Beutel RG (2011) The male postabdomen of the ancestral archostematan beetle Tetraphalerus bruchi Heller, 1913 (Ommatidae) and its phylogenetic significance. Arthropod Structure & Development 40, 146–158. doi:10.1016/j.asd.2010.12.003

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Jarzembowski EA, Wang B, Zhang H, Fang Y (2015) Boring beetles are not necessarily dull: New notocupedins (Insecta: Coleoptera) from the Mesozoic of Eurasia and East Gondwana. Cretaceous Research 52(B), 431–439.

Jarzembowski EA, Wang B, Zheng D (2017) A new ommatin beetle (Insecta: Coleoptera) with unusual genitalia from mid-Cretaceous Burmese amber. Cretaceous Research 71, 113–117. doi:10.1016/j.cretres.2016.10.010

Kirejtshuk AG, Nel A, Collomb FM (2010a) New Archostemata (Insecta: Coleoptera) from the French Paleocene and early Eocene, with a note on the composition of the suborder. Annales de la Société Entomologique de France (N.S.) 46(1–2), 216–227.

Kirejtshuk AG, Ponomarenko AG, Prokin AA, Chang H, Nikolajev GV, Ren D (2010b) Current knowledge of Mesozoic Coleoptera from Daohugou and Liaoning (northeast China). Acta Geologica Sinica 84(4), 783–792[English Edition]. doi:10.1111/j.1755-6724.2010.00253.x

Kirejtshuk AG, Nel A, Kirejtshuk PA (2016) Taxonomy of the reticulate beetles of the subfamily Cupedinae (Coleoptera: Archostemata), with a review of the historical development. Invertebrate Zoology 13(2), 61–190.

Lawrence JF (1999) The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebrate Taxonomy 13, 369–390. doi:10.1071/IT99008

Martin SK (2010) Early Jurassic coleopterans from the Mintaja insect locality, Western Australia. Acta Geologica Sinica 84(4), 925–953 [English Edition]. doi:10.1111/j.1755-6724.2010.00276.x

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Neboiss A (1968) The type of Omma mastersi Macleay (Coleoptera: Cupedidae). Memoirs of the National Museum of Victoria 28, 21. doi:10.24199/j.mmv.1968.28.05

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3. CUPEDIDAE LAPORTE, 1836

John F. Lawrence and Hermes E. Escalona

Fig. 3.1. Adinolepis matthesonae (Neboiss).

Common name. Reticulated beetles

Introduction. The family Cupedidae is a relictual group, containing 31 extant species in nine genera. According to Lawrence (1999), the modern cupedid fauna includes two subfamilies: Priacminae (Crowson 1962) with Priacma serrata (LeConte) from north-western North America and Paracupes brasiliensis (Kolbe) from Brazil and eastern Ecuador, and Cupedinae with Cupes Fabricius (eastern North America), Prolixocupes Neboiss (Chile-Argentina and south-western North America), Tenomerga Neboiss (Holarctic, Afrotropical and Oriental), Rhipsideigma Neboiss (Afrotropical), Ascioplaga Neboiss (Australia and New Caledonia) and the endemic Australian genera Distocupes Neboiss and Adinolepis Neboiss. Relationships of some of these genera are under scrutiny and some authors (Kirejtshuk 2005; Kirejtshuk et al. 2010a, 2010b, 2016) would dispense with the Priacminae, as well as the genera Distocupes and Tenomerga (= Cupes) and consider the families Cupedidae and Ommatidae to be subfamilies of Cupedidae extending back to the Lower Triassic. There have been two cladistic analyses of modern cupedids: one based primarily on head morphology (Hörnschemeyer et al. 2006) and another using a variety of morphological features (Hörnschemeyer 2009). In the first, a monophylum was recognised containing Distocupes, Adinolepis and Ascioplaga based mainly on the type of cuticular scales and the presence of a deep ventral mandibular cavity. This clade was sister to one containing Rhipsidiegma, Prolixocupes, Cupes and Tenomerga, which in turn, formed a clade with two basal lineages for Priacma and Paracupes. In the later study, there was a succession of clades for Priacmini, Prolixocupes and Ascioplaga + Adinolepis, with a large derived clade with Distocupes at the base and two main clades: the South African Tenomerga leucophaea + Cupes capitatus + Rhipsideigma sister to the remaining eleven Tenomerga species (Ge & Yang 2004; Hörnschemeyer & Yavorskaya 2016). Most references on Mesozoic fossils currently placed in this family may be found in the section on Archostemata; more recently, two new genera have been described from Cretaceous amber from Myanmar (Jarzembowski et al. 2017a, 2017b, 2017c).

Biology. Cupedid larvae are known to bore into dead wood that has been attacked by wood-rotting fungi. Snyder (1913) found larvae of the North American Tenomerga cinerea (Say) in wood infested with Daedalea quercina (Polyporaceae), while those of the Asian T. mucida (Chevrolat) were associated with a species of Stromatoscypha (Meripilaceae) (Fukuda 1941). Larvae of the Australian Distocupes varians (Lea) have been collected in structural timber and also in old rotten pine logs on the ground (Neboiss 1960, 1968, 1984, 1987; Lawrence 1999).

Characteristics. Adults. Elongate, strongly flattened, from 6 to 17 mm in length. Body surfaces entirely or mostly covered with scale-like setae. Head prognathous, constricted posteriorly to form a narrow neck and distinct temples. Eyes relatively large and more or less protuberant, entire, very finely facetted, without interfacetal setae. Antennal insertions dorsal and relatively narrowly separated, not concealed by frontal ridges. Head dorsally with one (Adinolepis and Ascioplaga) or two (Distocupes) pairs of subconical tubercles. Frontoclypeal suture and subantennal grooves absent. Labrum free, very small, apically truncate. Antennae moderately long, 11-segmented and filiform, with basal two to several antennomeres covered with scales. Mandible short and broad, unidentate with one or two subapical teeth; without mola or prostheca. Maxilla highly reduced and partly concealed by mentum, galea and lacinia hyaline and setose, the former very slender, the latter broad, without uncus. Terminal maxillary palpomere expanded and securiform with patch of sensilla lying on surface and not in deep cavity. Mentum distinctly transverse, concealing palpal insertions and prementum, the latter consisting of two trilobed, membranous processes; terminal labial palpomere expanded and secufirom. Gular sutures well separated; gula longer than wide. Corpotentorium weakly developed or absent. Pronotum widest at about middle, distinctly narrower at base than combined elytral bases. Lateral pronotal carinae complete, anterior angles usually slightly to moderately produced laterally (strongly produced forward in Ascioplaga). Prosternum in front of coxae moderately long and convex, with paired oblique crural impressions extending mesally almost to midline, where separated by a short carina, and laterally across the pleuron and onto the pronotal hypomeron. Notopleural and pleurosternal sutures separated posteriorly on either side of elongate, slender, anteriorly acute pleuron, the sutures uniting just before the anterior edge and continuing as a short notosternal suture. Prosternal process complete, narrowed posteriorly with narrowly rounded apex. Procoxae subglobular, not strongly projecting, without concealed lateral extensions, with distinct coxal plates; trochantin large, well sclerotised, broadly exposed and abutting sternum and pleuron. Procoxal cavities slightly transverse, narrowly separated, broadly open externally and internally. Scutellar shield abruptly elevated, anteriorly simple, posteriorly expanded and rounded to truncate. Elytra complete, usually with 5 longitudinal carinae (including lateral and sutural edges); epipleura narrow and complete. Mesoventrite with small cavity and short discrimen. Mesoventral process divided into two short, acute processes. Mesocoxae conical and projecting, with broadly exposed trochantins. Posterior portion of mesanepisterna, and entire mesepimeron concave, forming part of oblique crural impression for mid leg. Mesocoxal cavities contiguous, broadly closed laterally by mesepimera and metanepisterna. Metaventrite strongly narrowed anteriorly; discrimen more than half as long as ventrite and transverse (metakatepisternal) suture well developed. Metanepisternum completely exposed with anterior concavity and longitudinal groove forming part of crural impression for mid leg. Exposed portion of metepimeron narrowly elongate. Metacoxae strongly transverse with subconical mesal projections, contiguous, extending laterally to meet elytra, with well developed coxal plates. Metendosternite with a long, slender stalk, short, narrow ventrolateral processes, short, broad lateral arms, and a very long and slender anterior process bearing approximate anterior tendons. Hind wing moderately broad, with very short bending zone near apex of radial bar (anterior wing strut); apical field very short, with three radial extensions; pterostigma absent; RP complete to wing base, usually joined to RA3+4 by two cross-veins; RP and MP joined by two cross-veins forming the oblongum cell; MP1+2 with hinge crossed by fold and with short, straight medial spur; medial field with four or five free veins and no medial fleck; wedge cell well developed and apically truncate; anal lobe well developed, without embayment; AP3+4 forked. Legs short and slender; tibia distinctly longer than femur; tibial spurs small and paired on all legs. Tarsi 5–5–5; tarsomere 4 lobed beneath and tarsomeres 1–4 densely setose beneath; pretarsal claws simple; empodium not apparent. Abdomen with five ventrites, which are flattened but slightly overlapping; ventrite 1 distinctly shorter than 2, with well developed, narrowly acute intercoxal process. Functional spiracles on abdominal segments I–VIII. Anterior edge of segment IX in male broadly rounded; posterior edge narrowly rounded or acute; tergite IX at least partly fused to X. Aedeagus of modified trilobate type, with parameres completely fused to phallobase and partially fused together, each with a small hook near apex; penis with short basal struts. In Distocupes, testes are of the follicular type. Male diploid chromosome number is 2n = 19, with a male sex chromosome system of the XO type and achiasmatic meiosis (Galián & Lawrence 1993). Sternite VIII in female with long, fixed spiculum ventrale. Ovipositor narrowly elongate, lightly pigmented except for bacula; paraprocts much longer than gonocoxites, which bear well developed gonostyli. (Hörnschemeyer & Yavorskaya 2016).

Larvae. Elongate, more or less parallel-sided, lightly pigmented except for mandibles and apex of tergum IX; surfaces smooth, clothed with scattered long setae. Head prognathous and protracted. Posterior edge of head capsule deeply emarginate; epicranial suture not apparent; median endocarina simple, not forked, extending as far as clypeus. Stemmata usually absent (one on each side in first instar of Distocupes Neboiss). Frontoclypeal suture indistinct. Labrum free, strongly transverse. Antennae in late instars moderately long and usually 4-segmented, with sensorium on antennomere 3 about half as long as antennomere 4; antennae of first instar 2-segmented with sensorium almost as long as apical antennomere. Mandibles tridentate; molae large, slightly asymmetrical and irregular with no asperities or transverse ridges; prostheca absent. Ventral mouthparts retracted forming maxillolabial complex, with well developed maxillary articulating area. Galea articulated and palpiform, lacinia fixed and apically rounded with several stout spines and setae. Maxillary palp 3-segmented with distinct palpifer. Labium consisting of prementum and postmentum, the former with a well developed ligular sclerome. Labial palps 2-segmented and widely separated. Hypostomal rods long and diverging; ventral epicranial ridges absent. Gular sutures separated; gula transverse. Meso- and metatergum with strongly transverse ambulatory ampulla. Thoracic venter with large field of asperities. Legs reduced, 6-segmented, with two unequal pretarsal claws or with claws fused together. Abdomen with transverse tergal ampulae present on most abdominal segments. Tergum IX slightly longer than VIII, apically narrowed to form narrow truncate process. Sternite IX exposed, simple; segment X posteroventrally oriented, with pair of large, globular projecting lobes. Anal opening Y-shaped, between these lobes and a weak transverse dorsal lobe. Spiracles annular, with closing apparatus. (Neboiss 1968; Grebennikov 2004; Yavorskaya et al. 2015).

Key to the Australian genera and species

1. Six or more basal antennomeres clothed with scales ( Fig. 3.4 , 3.7 ); antennae shorter, extending at most to basal fourth of elytra (Pl. 14A–D. N–O); head with cylindrical, apically rounded scales intermixed with flattened scales ( Figs. 3.2 , 3.4 , 3.7 ); length usually less than

10 mm…Adinolepis………………………………2

–Fewer than six basal antennomeres clothed with scales ( Figs. 3.5 , 3.8 – 3.9 ); antennae longer, usually extending beyond basal third of elytra; head without cylindrical, apically rounded scales ( Fig. 3.3 , 3.5 ); length almost always greater than 10 mm 4

2(1). Eyes much larger than temples and extending laterally beyond the edges of temples ( Fig. 3.4 ; Pl. 14A–B); anterior edge of lateral pronotal carina with distinct tooth, so that pronotal width at this point is as great as maximum pronotal width; 6–9 mm (Pl. 14A–B); VIC and NSW to southern QLD……………………………… A. yuanga (Neboiss)

–Eyes not or only slightly longer than temples and not extending laterally beyond their edges ( Fig. 3.2 , 3.7 ; Pl. 14C–D, N–O); anterior edge of pronotal carina without or with very small tooth, so that width at this point is always less than maximum pronotal width……………………………… 3

3(2). Elytral scales more generally distributed, usually extending onto window punctures ( Fig. 3.6 ); colour darker with white scales forming three irregular transverse bands; 6.3–7 mm (Pl. 14N–O); VIC, SA, and NSW……………………………… A. eumana (Neboiss)

Figs 3.2–3.9. Australian Cupedidae. 2, Adinolepis eumana (Neboiss), head and prothorax, dorsal, showing less prominent eye and lack of distinct lateral pronotal tooth; 3, Distocupes varians (Lea), head and prothorax, dorsal, showing lack of apically rounded scales and presence of conical tubercles over antennal bases and eyes; 4, Adinolepis youanga (Neboiss), head and prothorax, dorsal, showing promient eyes and well developed lateral pronotal tooth; 5, Ascioplaga scalena (Neboiss), head and prothorax, dorsal, showing the first 5 antennomeres clothed with scales and a single conical tubercle over the antennal base; 6, Adinolepis eumana (Neboiss), anterior portion of elytra, dorsal, showing the more extensive covering of scales; 7, Adinolepis eumana (Neboiss), head and prothorax, dorsal, showing more than 5 antennomeres clothed with scales; 8, Ascioplaga scalena (Neboiss), head and prothorax, lateral, showing the first 5 antennomeres clothed with scales and a single conical tubercle over the antennal base; 9, Distocupes varians (Lea), head and prothorax, lateral, showing conical tubercles over both antennal bases and eyes.

–Elytral scales distributed mainly on the longitudinal costae and not or rarely extending onto the window punctures; colour lighter, except for dark areas alternating with white ones on costae ( Fig. 3.1 ; Pl. 14); 7–10 mm (Pl. 14C–D); VIC, NSW and ACT……………………………… 4

4(3). Elevated elytral interval 3 with three sections of pale scales; VIC, NSW and ACT……………………………… A. mathesonae (Neboiss)

–Elevated elytral interval 3 with two sections of pale scales; WA A. apodema Neboiss

5(1). Two basal antennomeres clothed with scales ( Fig. 3.9 ); head with two pairs of conical tubercles, one above antennal bases and one above eyes ( Fig. 3.9 ); 10–14.5 mm (Pl. 14K–L); VIC, ACT, NSW, southern QLD……………………………… Distocupes varians (Lea)

–Five basal antennomeres clothed with scales ( Fig. 3.5 , 3.8 ); head with one pair of conical tubercles above antennal bases ( Fig. 3.8 ); 16.2 mm (Pl. 14D); northern QLD……………………………… Ascioplaga scalena (Neboiss)

Classification of the Australian genera

Distocupes Neboiss, 1984 (Pl. 14K–L; Figs 3.3, 3.9)

Type species. Cupes varians Lea, 1902.

Australian species. One.

Distribution. Distocupes varians (Lea) is known from TAS, VIC, NSW and QLD.

Biology. Larvae of this species have been taken in structural timber and in partly buried pine logs in a garden. As in Tenomerga mucida (Chevrolat) (Yavorskaya et al. 2015), the first instar of D. varians has 2-segmented antennae.

References. Neboiss (1960, 1968, 1984).

Adinolepis Neboiss, 1984 (Pl. 14A–D,

N–O; Figs. 3.2, 3.4, 3.6, 3.7)

Type species. Cupes eumana Neboiss, 1960.

Australian species. Four.

Distribution. Three of the four species of Adinolepis occur in the south-eastern part of the continent (VIC, NSW and QLD), while A. apodema Neboiss is known from southern WA. Hörnschemeyer (2009) suggested that A. apodema might be a colour variant of A. mathesonae Neboiss, which is known from the south-eastern part of the continent.

References. Neboiss (1960, 1984, 1987)

Ascioplaga Neboiss, 1960 (Pl. 14E; Fig. 3.5, 3.8)

Type species. A. mimeta Neboiss, 1960.

Australian species. One.

Distribution. The genus Ascioplaga occurs in New Caledonia and Australia. Ascioplaga scalena (Neboiss) was described from northern QLD.

References. Neboiss (1984); Hörnschemeyer (2009).

References

Crowson RA (1962) Observations on the beetle family Cupedidae, with descriptions of two new forms and a key to the recent genera. Annals & Magazine of Natural History 5(51), 147–157. doi:10.1080/00222936208651227

Fukuda A (1941) Some ecological studies on Cupes clathratus. Transactions of the Natural History Society of Formosa 31, 394–399.

Galián J, Lawrence JF (1993) First karyotypic data on a cupedid beetle (Coleoptera: Archostemata) showing achiasmatic meiosis. Entomological News 104(2), 83–87.

Ge SQ, Yang XK (2004) Two new Chinese species of Tenomerga Neboiss (Coleoptera: Cupedidae), with a world catalog of the genus. Proceedings of the Entomological Society of Washington 106(3), 631–638.

Grebennikov VV (2004) Review of larval morphology of beetles of the suborder Archostemata (Insecta: Coleoptera), including first-instar chaetotaxy. European Journal of Entomology 101, 273–292. doi:10.14411/eje.2004.038

Hörnschemeyer T (2009) The species-level phylogeny of archostematan beetles – where do Micromalthus debilis and Crowsoniella relicta belong? Systematic Entomology 34, 533–558. doi:10.1111/j.1365-3113.2009.00476.x

Hörnschemeyer T, Yavorskaya M (2016) 5.2. Cupedidae Laporte, 1832. Introduction and phylogeny. In Handbuch der Zoologie/Handbook of Zoology. BandVolume IV Arthropoda: Insecta. Teilband/Part 38. Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2nd edn. (Eds RG Beutel and RAB Leschen) pp. 44–47. Walter de Gruyter, Berlin.

Hörnschemeyer T, Goebbels J, Weidemann G, Faber C, Haase A (2006) The head morphology of Ascioplaga mimeta (Coleoptera: Archostemata) and the phylogeny of Archostemata. European Journal of Entomology 103, 409–423. doi:10.14411/eje.2006.055

Jarzembowski EA, Wang B, Zheng D (2017a) The first cupedine beetle from Burmese amber (Coleoptera: Cupedidae). Comptes Rendus. Palévol 16, 241–247. doi:10.1016/j.crpv.2016.08.004

Jarzembowski EA, Wang B, Zheng D (2017b) A new reticulated beetle (Coleoptera: Cupedidae) with aedeagus preserved from mid-Cretaceous amber of Myanmar. Cretaceous Research 80, 86–90. doi:10.1016/j.cretres.2017.08.015

Jarzembowski EA, Wang B, Zheng D (2017c) A new spiny rerticulated beetle (Coleoptera: Cupedidae) from Cretaceous Burmese amber. Proceedings of the Geologist’s Association. doi:10.1016/j.pgeola.2017.07.003.

Kirejtshuk AG (2005) A revision of the genus Cupes Fabricius, 1801 from Baltic amber and some notes on taxonomy and composition of the family Cupedidae (Coleoptera, Archostemata). Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg 89, 55–84.

Kirejtshuk AG, Nel A, Collomb FM (2010a) New Archostemata (Insecta: Coleoptera) from the French Paleocene and early Eocene, with a note on the composition of the suborder. Annales de la Société Entomologique de France (N.S.) 46(1–2), 216–227.

Kirejtshuk AG, Ponomarenko AG, Prokin AA, Chang H, Nikolajev GV, Ren D (2010b) Current knowledge of Mesozoic Coleoptera from Daohugou and Liaoning (northeast China). Acta Geologica Sinica 84(4), 783–792[English Edition]. doi:10.1111/j.1755-6724.2010.00253.x

Kirejtshuk AG, Nel A, Kirejtshuk PA (2016) Taxonomy of the reticulate beetles of the subfamily Cupedinae (Coleoptera: Archostemata), with a review of the historical development. Invertebrate Zoology 13(2), 61–190.

Lawrence JF (1999) The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebrate Taxonomy 13, 369–390. doi:10.1071/IT99008

Neboiss A (1960) On the family Cupedidae, Coleoptera. Proceedings of the Royal Society of Victoria 72, 12–20.

Neboiss A (1968) Larva and pupa of Cupes varians Lea, and some observations on its biology (Coleoptera; Cupedidae). Memoirs of the National Museum of Victoria 28, 17–19. doi:10.24199/j.mmv.1968.28.04

Neboiss A (1984) Reclassification of Cupes Fabricius (s. lat.), with descriptions of new genera and species (Cupedidae: Coleoptera). Systematic Entomology 9, 443–477. doi:10.1111/j.1365-3113.1984.tb00520.x

Neboiss A (1987) A new species of Adinolepis Neboiss from Western Australia (Cupedidae: Coleoptera). Records of the Western Australian Museum 13, 323–325.

Snyder TE (1913) Record of the rearing of Cupes concolor Westw. (Coleoptera). Proceedings of the Entomological Society of Washington 15, 30–31, pl. 1.

Yavorskaya MI, Kojima K, Machida R, Beutel RG (2015) Morphology of the first instar larva of Tenomerga mucida (Chevrolat, 1829) (Coleoptera: Archostemata: Cupedidae). Arthropod Systematics & Phylogeny 73(2), 239–258.

4. MYXOPHAGA CROWSON, 1955

John F. Lawrence

The suborder Myxophaga contains four small families of minute beetles inhabiting aquatic or riparian habitats: Lepiceridae (Mexico to northern South America, Cretaceous of South-east Asia), Torridincolidae (South America, southern and central Africa, Madagascar, Japan and China), Sphaeriusidae (North to South America, Asia, Madagascar, Africa, Australia) and Hydroscaphidae (western North America, Mexico to South America, Eurasia, North Africa, South-east Asia and Madagascar) (Beutel 1999; Navarrete-Heredia et al. 2005; Anton & Beutel 2006; Kirejtshuk & Poinar 2006, 2013; Flowers et al. 2010; Ge et al. 2010; Sampaio & Ferreira 2013; Shepard et al. 2013; Short et al. 2015).

Adult subordinal autapomorphies include the absence of a maxillary galea, a movable tooth on the left mandible, a broad connection between the mesoventrite and metaventrite and long fringe-hairs lining the margins of the hindwings. The exposure of the propleuron, which extends anteriorly to the edge of the prothorax and posteriorly behind the procoxae, is a feature shared with both Archostemata and Adephaga, but in myxophagan adults, unlike those of the other two suborders, prothoracic trochantin is solidly attached to the coxa, separated by a suture in Lepicerus but solidly fused in the other myxophagan families to form a trochantinopleuron, as in Polyphaga. Wing venation and folding in Myxophaga, although highly reduced due to small size, is similar to that in Archostemata and Adephaga and very different from that in Polyphaga: (1) RA1+2 and RA3+4 diverge but never converge again to form a radial cell; and the latter extends into the apical field; (2) the posterior wing strut (MP1+2) has a distinct medial hinge through which a fold passes; (3) an oblongum cell is almost always formed between RP and MP1+2; (4) there is usually more than one cross-vein between RA and RP. The spiral rolling of the apical field is a character shared with Archostemata but not with Adephaga. Other features of adult Myxophaga include the small size (less than 2.8 mm but usually much smaller), antennae usually less than 10-segmented (11-segmented in Delevea, Satonius and Sphaerius), with the terminal antennomere more or less enlarged (with a compact 3-segmented club in Sphaerius), mandibular mola present; mesoventrite very short, mesocoxal cavities partly closed laterally by the mesepimeron and a narrow portion of the metanepisternum in Torridincolidae, but by the mesepimeron only in the other families. Internal features include the six free Malpighian tubules, tube-like, coiled testes and a telotrophic ovary (Hydroscapha).

Myxophagan larvae resemble those of basal Polyphaga in having the tibia and tarsus fused to form a tibiotarsus and microphagous mouthparts including a well developed, tuberculate or asperate mandibular mola. Although both of these features have been used to support a sister-group relationship between the two suborders, there are several structural differences in the mouthparts that suggest an independent development of microphagy in the two groups. Basal polyphagan larvae, for instance, have structures such as the hypopharyngeal bracon, hypopharyngeal sclerome and accessory ventral process on the mandible, which are absent in Myxophaga, while myxophagan larvae have sensory papillae on the labium. A distinctive feature of myxophagan larvae is the presence of three to six relatively large stemmata borne on a distinct lateral protuberance. The most prominent feature of almost all myxophagan larvae is the presence of spiracular gills, but the segmented structures of Torridincolidae differ significantly from the balloon-like gills of Hydroscaphiae and Sphaeriusidae. Tracheal gills are absent in the presumed larva of Lepiceridae (Lawrence et al. 2013), but there is some indirect evidence that this larva may have been misidentified. Antennae in most myxophagan larvae are 2-segmented, with the long, slender and more or less acute sensorium arising from antennomere 1 in Hydroscaphidae but from the middle or near the apex of antennomere 2 in Sphaeriusidae and Torridincolidae. Additional information on both larval and adult morphology may be found in Arce-Peréz et al. 2016, Beutel 1998, Beutel 2016, Beutel & Arce-Peréz 2016, Beutel & Haas 1998, Beutel & Vanin 2016, Beutel et al. 1999, Büning 2005, Vanin et al. 2016.

Crowson (1960) considered this taxon to be the sister group of Polyphaga based mainly on the fusion of the larval tibia and tarsus, and this conclusion was supported by Beutel & Haas (2000) and several other studies, but not by McKenna et al. (2015), who concluded that Archostemata + Myxophaga + Adephaga were sister to Polyphaga. Ponomarenko (1969) suggested that Recent Myxophaga may be descendants of the schizophoroid Archostemata (Ponomarenko 1969), especially the primarily Triassic and Jurassic Catiniidae, but at least two catiniid genera from the Jurassic and early Cretaceous, Cervicatinus Tan & Ren (2007) and Forticatinius Tan & Ren (2007), have been shown to be members of the family Trogossitidae (Polyphaga: Cucujiformia) (Kirejtshuk et al. 2010; Yu et al. 2012, 2015; Kirejtshuk 2017). Within the Myxophaga, morphological cladograms based on adults of all families and larvae of all except Lepiceridae (Beutel 1999) indicated that Lepiceridae was sister to Sphaeriusidae, Hydroscaphidae and Torridincolidae, but in cladograms based on molecular data (McKenna et al. 2015), Hydroscaphidae was sister to the remaining families and Sphaeriusidae + Lepiceridae sister to Torridincolidae. Fossil Lepiceridae from Burmese amber are in a confused state due in part to the poor condition of the types leading to morphological misinterpretations; at present, the fossils are placed in two genera, Haplochelus and Lepiceroides, by Kirejtshuk and Poinar (2006, 2013) but considered to be species of Lepicerus by Jałoszyński et al. (2017).

Myxophaga occur in various riparian or aquatic situations, such as in mud and sand in and at the edges of streams and ponds, in hot springs (some Hydroscapha) and on the surfaces of rocks which are covered with a moving film of water or constantly sprayed by waterfalls (hygropetric habitats). Although adults of Torridincolidae may have a ventral plastron (Hinton 1969) and larvae of Torridincolidae, Sphaeriusidae and Hydroscaphidae possess spiracular gills (tracheal gills) (Hinton 1967). The suborder is poorly represented in Australia by a single species of Sphaerius (see Chapter 5).

References

Anton E, Beutel RG (2006) On the head morphology of Lepiceridae (Coleoptera: Myxophaga) and the systematic position of the family and suborder. European Journal of Entomology 103, 85–95. doi:10.14411/eje.2006.014

Arce-Pérez J, Navarrete-Heredia JL, Beutel RG (2016) 6.1. Lepiceridae Hinton, 1936 (= Cyathoceridae). In Handbuch der Zoologie/Handbook of Zoology. BandVolume IV Arthropoda: Insecta. Teilband/Part 38. Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2nd edn. (Eds RG Beutel and RAB Leschen) pp. 65–67. Walter de Gruyter, Berlin.

Beutel RG (1998) Torridincolidae: II. Description of the larva of Satonius kurosawai (Satô, 1982) (Coleoptera). In Water Beetles of China. Vol. II. (Eds MA Jäch and L Ji) pp. 53–59. Zoologisch-Botanische Gesellschaft in Österreich & Wiener Coleopterologenverein, Vienna.

Beutel RG (1999) Phylogenetic analysis of Myxophaga (Coleoptera) with a redescription of Lepicerus horni (Lepiceridae). Zoologischer Anzeiger 237(1998/1999), 291–308.

Beutel RG (2016) 6. Myxophaga Crowson, 1955. Introduction, phylogeny. In Handbuch der Zoologie/Handbook of