Trogossitidae: A review of the beetle family, with a catalogue and keys

Abstract The family Trogossitidae (Coleoptera: Cleroidea) is reviewed to species level. Keys to its genera, tribes and subfamilies are presented for the first time. All known species and subspecies are listed, together with complete taxonomic references back to 1910, the date of issue of the last catalogue of Trogossitidae. Higher taxa reviews are accompanied by remarks on phylogeny, distribution and biology as well as a brief description of adults and larvae. All known fossil records of Trogossitidae are reviewed and discussed. The work includes maps of distribution, colour photographs of generic representatives, morphological illustrations, SEM photographs and phylogenetic trees.


Introduction
The main purpose of the work is to introduce some modern order into current knowledge of the family Trogossitidae and extend knowledge of this relatively small but fascinating group of beetles, especially to both amateur entomologists and professional "non-cleroid" workers. It is deliberately written as a "compilation" of papers on the topic to date, especially because some of them were published in journals and books that are not easily accessible to all, and to bring various fragmented sources together.
Because of the character and purpose of the work, I have tried to avoid introducing any new thoughts and systematic changes, apart from a few minor ones mentioned in the "New taxonomic acts" section. A catalogue of species lies at the core of the work. I have not repeated references included in Coleopterorum Catalogus of Temnochilidae by ; however, Léveillé's reference always takes first place in any particular reference list. Taxonomic references follow, just as they have been excerpted from Zoological Records after 1910. The catalogues for some species are, without doubt, incomplete. Some references for biology and local distribution must also, perforce, be lacking -I beg, therefore, the kind reader's patience and leniency.
The systematics of Trogossitidae is still in its infancy. There remains a great deal of work to be done in the higher taxonomy, as well as with regard to generic limits, especially in widespread, species-rich genera. Ancyrona is a good example of such a genus, distributed from tropical Africa, the Palaearctic, south-eastern Asia to Australia. Tenebroides and Temnoscheila are further complex taxa, each with more than a hundred described species distributed in both North and South America. On the other hand, there also exists a relatively rich modern material of trogossitids to be collected in various parts of the world, certainly containing plenty of new species. Unfortunately, only a few people are seriously interested in the family and only a few of them, in turn, try to gather and publish further information. Therefore, another purpose of the book is to encourage interest in this highly interesting group of beetles.

Brief review of classification
The superfamily Cleroidea was established by Böving and Craighead (1931). Until that time, Trogossitidae had been classified within Clavicornia together with Nitidulidae (usually as Ostomidae but also as Ostomatidae, Peltidae and Temnochilidae). The names Trogositidae (from Trogositae Fabricius, 1801) and the correct spelling Trogossitidae (from Trogossitarii Latreille, 1802) are the most modern forms of the name (see Kolibáč and Leschen 2010 for details). The family Phloiophilidae is mentioned in Pic's (1926) catalogue of Melyridae sensu lato (the family is sometimes referred to as Phloeophilidae as well).  discussed the classification of its only species in detail and classified it within modern Cleroidea. I have suggested a classification of Phloiophilidae as a tribe in Peltinae ) but this has not found wide acceptance. Reitter (1876) published an excellent world-wide review that is basic to the study of Trogossitidae. Similarly, the world catalogue by  is among the classic works. The number of genera and species has only slightly increased since the publication of the latter list, even on a world scale. Crowson ( , 1966Crowson ( , 1970 has changed the rank or status of subfamilies and families classified within Trogossitidae sensu lato several times. Subsequently,  and later Ślipiński (1992) integrated Crowsons's families into the single family Trogossitidae. Several years ago, I established tribes in all trogossitid subfamilies on the basis of the morphological characters of adults and larvae . The 2008 system is employed throughout this work.  is always the first reference in catalogues. Reitter (1876), as the most important reference in some taxa, is also listed. The note "synonymized by author" refers to the author of the preceding reference.
More than seven years have passed since I formulated theses on the higher classification of Trogossitidae. Although some opinions about the phylogeny have changed and the systematic placement of some genera has recently been called into question, the main purpose of the keys is confined to identification of the trogossitid genera. The keys are given for extant subfamilies, tribes and genera. Extinct taxa are listed in relevant sections, together with their descriptions and remarks on their classification. Generic names in parentheses in particular descriptions denote a similar character state occuring in another genus or genera.
The morphological descriptions of particular genera are largely based on several hundred detailed ink-drawings that have already been published by myself . PDF files of the papers that include them, as well as all other relevant publications since 2000, are available on request (see author's address). All scale bars in plates (Figs 1-12) express one millimetre. Beetles in colour plates  are pictured in approximate proportion ("large species" are larger than "small species"). White arrows in SEM photographs  denote important characters further mentioned in relevant captions. Numbers in parentheses in maps of distribution (Maps 1-13) denote the number of species within the given genus. Metendosternite with conspicuous lateral arms. Wings usually present, but missing in some species. Apical field sometimes with one or more small sclerites just beyond radial cell; RP2 sometimes present. Radial cell as long as wide or shorter than wide, sometimes very reduced or absent; cross-vein r3 usually absent. Basal portion of RP short or sometimes absent. Medial field with as many as four free veins, a wedge cell and no medial fleck (fewer veins and no wedge cell in smaller species); anal embayment usually notch-like, absent in some Trogossitini. Trochanters triangular. Femora sometimes clavate. Tibiae often with row of spines along outer side; apex of tibia with row of spines and two hooked spurs or only one spur hooked or spurs reduced (spines reduced in smaller taxa); tibial spurs pattern varies from 2-2-2 to 0-0-0.  tarsomere 1 sometimes partially fused with 2 but always with conspicuous suture between them or very small and tarsal pattern seemingly 4-4-4 or 4-4-5; tarsomeres 1-4 never with membranous lobes; apical tarsomere usually as long as combined length of tarsomeres 1-4; claws large, without denticles (with the exception of some Decamerinae); empodium bisetose, strongly projecting. Abdomen with five or six ventrites. Ventrites I-III fused. Intercoxal process small, narrow. Spiculum of ventrite VIII sometimes present in males and always in females. Segment IX well-developed or reduced to "spicular fork". Aedeagus sheath-like cucujiform type, with fixed or articulated parameres that may be partly or entirely fused together or absent. Tegmen usually with anterior ventral strut and two opposing dorsal struts ("double tegmen" of Crowson 1964a), but usually inverted (rarely uninverted or placed laterally) and often composed of two or three parts (undivided in some members of the Rentonium group). Penis with two anterior struts. Ovipositor lightly sclerotized, except for baculi, moderate in size with sparsely pubescent coxites and styli. Bursa copulatrix large, spherical. Spermatheca elongate or oval, with gland. Vagina without sclerites. Six malpighian tubules present in Tenebroides and Lophocateres. Larvae ( Fig. 2) (according to Kolibáč and Leschen 2010). Five to seven larval instars observed in Trogossitinae (Temnoscheila, Tenebroides), four instars in Lophocaterinae (Lophocateres), all beetles reared in laboratory conditions. Body elongate, only weakly flattened. Colour white or pale, but sclerotised areas distinctly pigmented (head capsule, thoracic and abdominal terga, and urogomphi). Vestiture consisting of setae; rarely with bristles or expanded setae; sometimes body with short and sparse pubescence or only with setae on the last segment. Head protracted. Posterior edge of capsule slightly emarginate. Epicranial stem absent or present and of variable length. Median endocarina usually present, of variable length and usually extending between frontal arms (absent in Thymalus and/ or coincident with epicranial stem and frontal arms in Peltinae); paired endocarinae present or absent. Frontal arms V-shaped, straight or curved (nearly S-shaped). Five stemmata usually present and arranged in a pattern with two anteriorly and three in a posterior row; sometimes reduced to four, three, two or none. Frontoclypeal suture usually absent (distinct in the Rentonium group and Thymalus). Labrum free; epipharynx membranous; shape of tormae variable and lacking posterior extensions. Antennae 3-segmented, with short sensorium present at apex of segment 2. Mandibles with one or two apical teeth (serrate in the Rentonium group); mola absent or present; mesal edge of mandibular base with brush of hairs or rigid denticulate processes that may be hylaline. Ventral mouthparts retracted. Maxillary articulating area present or absent. Cardo typically undivided (divided in Calitys). Mala with apex usually simple, with large pedunculate seta in predatory species (Trogossitini and Egoliini); inner apical angle usually lacking small teeth (present in Protopeltus and Larinotini); palps with four, three (e.g., Ancyrona and Lophocateres), or two (Rentonium group) palpomeres. Labium consisting of prementum, mentum, and submentum, or preand postmentum (Thymalus and Parapeltis); mentum or postmentum free or connate with base of maxillae; prementum sclerotized and elongate; mentum mostly unsclerotized in some taxa; ligula absent or present; if present apex emarginate or not, or divided apically; palps usually 2-segmented (1-segmented in the Rentonium group). Gular region longer than wide, or wider than long; fused to labium or not. Hypostomal rods present, reduced or absent; sometimes extending to posterior edge of head; subparallel or diverging posteriorly. Ventral epicranial ridges present or absent. Prothorax usually with one large sclerite dorsally and one elongate sclerite ventrally. Protergum with or without sclerotized plate with a longitudinal median ecdysial line. Meso-and metathorax usually with pair of sclerites dorsally (absent in some taxa) and one weakly sclerotised, pale plate ventrally. Sometimes all thoracic sclerites indistinct. Coxae widely separated. Thoracic legs 5-segmented, including claw-like pretarsus with single seta. Nine abdominal segments visible from above. Abdominal ampullae present or absent. Segment IX shorter than or subequal to VIII. Segment X almost always concealed by segment IX (visible from above only in Larinotus). Urogomphi usually well-developed (sometimes reduced) and dorsally or posteriorly oriented; large, hook-shaped or nearly straight; strongly sclerotized and pigmented; often with spines or secondary processes; apically bifurcate or not; pit present between urogomphi in Parapeltis; median process present between urogomphi in Lophocaterini and urogomphi located at apex of median process in some members of the Rentonium group. Anal region posteriorly or posterioventrally oriented; paired pygopods on segment X absent. Six malpighian tubules in Tenebroides; four in Lophocateres.

Key to subfamilies
Identification of the trogossitid subfamilies using the various determination keys published by a range of authors tends to be a complicated and frustrating process. Unfortunately, my "lumping" of nine former subfamilies (e.g., Ślipiński 1992) in two  rather complicated the identification of the individual specimen. In the traditional system for the trogossitids used in the 19th century (e.g. Erichson, Reitter, Léveillé), Peltinae were flat and fungivorous whereas Trogossitinae were cylindrical and predatory. Further study of such modified taxa as the rentoniins, decamerins or colydiopeltins revealed huge morphological and biological diversity within Peltinae (sensu . The same situation holds in Trogossitinae, in which superficially different taxa (such as Calitys, Larinotus, the gymnochilins and egoliins) are classified together in one subfamily. The subfamily Lophocaterinae was established by  and synonymized with Peltinae by  because of possible paraphyly of the latter subfamily. Later, in response to new observations, we suggested  that Peltinae be split once more into Lophocaterinae and Peltinae. The latter is the system used in this book.
Similar ways of life (members of the both subfamilies tend to be predatory), reductions of morphological structures common to the whole order Coleoptera (e.g. wing venation, lateral edge of pronotum, mola), mosaic character patterns and probably some underlying synapomorphies complicate the definition of subfamilies even other higher taxa in Trogossitidae, in much the same way as they do in the related family Cleridae. The key that follows is therefore not based on absolutely inclusive synapomorphies. The most important, clearly-visible characters appear in bold type.

1
Adult: labium with rigid ligula; epipharynx mostly with cordate sclerite along apex of labrum; antennal club mostly conspicuously asymmetrical, terminal antennomeres (antennal club) mostly with sensorial fields; front coxal cavities externally closed; body cylindrical or oval but not conglobate; end of elytral suture with distinct interlocking mechanism ("elytral lock"). Larva: head capsule with distinct endocarina, gular sutures and hypostomal rods; frontal arms mostly straight; gular region mostly with paragular sclerites. Mainly predatory, rarely fungivorous or phytophagous (e.g. feeding on grains Reitter, E. 1922: 66. Type genus. Calitys Thomson, 1859Bouchard, P. et al. 2011: 57. Crowson, R. A. 1970: 13 (referred as Calitinae subfam.nov.). Ślipiński, S. A. 1992: 442 (Calitinae). Lawrence, J. F. & Newton, A. F., Jr. 1995: 869 (Calitinae). : 117 (Calityni Winkler, 1922; sic!) (diagnosis, new status). : 364. Kolibáč, J. & Leschen, R. A. B. 2010 Remarks. The position of the single genus Calitys within the trogossitid system has changed many times over the past century or so. It has been classified within either Peltinae or Trogossitinae (compare, for example, vs. Crowson 1970, then treated as a separate subfamily (Crowson 1970in Trogossitidae s.str. without Peltidae, Ślipiński 1992 in Trogossitidae s.lat.). Reitter's early idea (1876) that it might be classified within the former Leperini or Leperinae (i.e. Gymnochilini herein) is also interesting and worthy of review. Calitys belongs among the primitive fungivorous groups and has several features shared with Peltinae, for example: robust mandibles with mola, flat body, wide pronotum, weakly asymmetrical antennal club, and absence of elytral interlocking mechanism. However, it also has bizarre sculptures on dorsal surface of body, wax scales, and tufts of rigid setae that together differentiate the genus from all other trogossitids. Its basal position in the trogossitine branch is based chiefly on procoxal cavities perfectly externally closed, presence of paragular sclerites in larval cranium and concave larval tergite IX. Wax scales and tufts of setae on head, antennae, elytra, and pronotum make it resemble Gymnochilini. Nonetheless, it remains possible to imagine Calitys also as a derived member of Peltinae or even Lophocaterinae.  Biology. Fungivorous. Live under bark of old coniferous trees (fir, pine) and on tree fungi. Calitys scabra was observed together with its larvae, for example, in the old stump of a fir Abies alba in Slovakia (J. Vávra, pers. observ.). Brustel (2009) recorded it from Antrodia sp. polypore fungi in Pyrenées Mts.
Distribution. Two species Holarctic. Two more species also reported from South Africa, of which C. spinifera is unknown to me. I studied a single C. africana non-type specimen in the Musée d'Histoire Naturelle in Geneva in 2003. It does not belong to Cleroidea. Map 1. A distribution of the tribe Calityini. agreement with my observations . Finally, character analyses (l.c.) also resulted in a close relationship between Larinotini and Egoliini. The relationship of adults appears to be so close that one might speculate about the paraphyly of Larinotini; it is, however, hard to justify such a hypothesis if one considers the monotypy of the latter (all derived features found in Larinotus are only autapomorphies).

Genus
Nebophilus Crowson, 1970[type species: Nebophilus hirsutus Crowson, 1970; designated by author] Crowson, R. A. 1970: 14. Lawrence, J. F. 1980 Description (see also Crowson 1970Crowson , Ślipiński 1992 Biology. The larvae and adults were found together by J. Doyen "in rotten wood beneath resupinate fungi" (Ślipiński 1992 : 4. Arias, E. et al. 2009: 40 quadrisignata Erichson, 1844Chile (AL, varA) Léveillé, A. 1910: 4. Arias, E. et al. 2009: 39. Kolibáč, J. 1999b: 12. Kolibáč, J. 2005 : 293. Arias, E. et al. 2009: 38. Kolibáč, J. 2005 shangrila Arias, Ślipiński, Lawrence & Elgueta, 2009; Chile (AD) Arias, E. et al. 2009: 39. Some remarks about the independent status of the tribe Gymnochilini with regard to Trogossitini are made below, in the section relating the latter tribe. The inclusion of Phanodesta from Juan Fernandez Isl. was more or less confirmed by two separate character analyses . However, the phylogeny of the genus was rendered unclear by the number of autapomorphies (e.g. winglessness, characteristic elytral structure) and separated distribution. I postulated a a sister group of Leperina with a "Gondwanan" distribution (Australia-Chile). Recently, a phylogeny and distribution of Phanodesta, together with descriptions and combinations of some more species from New Zealand and its vicinity have been addressed by  in detail. Two more genera occured in the gymnochiline clade in my second analysis : Seidlitzella and Melambia. The first genus, Seidlitzella, was considered related to the Palaearctic species of Leperina, as also pointed out by Schawaller (1993). However, his formal synonymization of Seidlitzella was not confirmed in a recent study by  who established the new genus Kolibacia for Leperina tibialis and L. squamulata instead.
In both analyses by , the genera Seidlitzella and Melambia were considered primitive or basal among the Gymnochilini or Trogossitini. They were included in the trogossitins in my original tribe definition . A comparison made specifically for the current shows the heterogenity of Melambia species and the need for revision of the genus with respect to the systematic position of particular species. It cannot be excluded that there are some species of Melambia congeneric with Alindria. Biology. Predatory. Adults run rapidly on logs and branches of fallen trees, hunting for prey. If disturbed, they fly very quickly. Some beetle collectors remark (P. Pacholátko, pers. comm.; experience from southern India) that some species can even jump(!) before they fly off.

Key to genera
Distribution. South-eastern Asia including Indonesia, Laos, Vietnam (numerous modern unpublished records). Often recorded together with species of Xenoglena.
Biology. The adults and larvae are probably predatory. An adult K. squamulata was, for example, found in rotten birch (Schawaller 1993).  remarked that K. squamulata develops under bark and sometimes within the trunks of various deciduous trees infested with the larvae of Melandrya, Tremex fuscicornis, Mesosa, Plagionotus and others. On the other hand,  consider both species mycophagous, citing Kryzhanovskij (1965) and Nikitsky (1992). The former source is unknown to me, but the latter author considers K. squamulata predatory. However, K. squamulata differs from Leperina in the presence of the mandibular mola which is definitely a feature of mycophagous cleroids, although it may also be considered a primitive (rudimentary) character, a trait seen in similar fashion in certain Gymnocheilis species (see figures in   . Unfortunately, swayed by various sources, I mistakenly changed the name Leperina to Lepidopteryx in the Errata to Volume 4 of the "Catalogue" (Kolibáč 2009, Löbl andSmetana 2010). Schawaller (1993: 2) considered Lepidopteryx a synonym of Leperina because of "invalid description" of the genus.
Most recently, Leschen and Lackner (2013: 289)   Lepidopteryx Hope, 1840, are objective synonyms because they share type species (Dejean, 1835: 314 (Fabricius, 1801) according to Léveillé, 1910: 22], although Reitter (1876: 37) placed Lepidopteryx as a synonym of Gymnochila. Note that White (1846) misspelled Gymnocheilis as Gymnocheila and wrongly attributed the name to Gray." Description (according to . Body size: 5.5-15.6 mm. Colour of body black and red-brown, unicolorous to multicoloured. Dorsal vestiture consisting of scales. Head extending beyond anterior angles of pronotum. Frons more or less horizontal with mandibles visible dorsally. Median lobe of clypeus absent. Edge of labrum weakly emarginate or straight. Eyes entire. Gena acute. Supraocular scales present or absent. Antenna 11-segmented with loose antennal club, lengths of antennomeres II and III equal or not; antennomere XI distinctly longer than wide and circular, about as long as wide. Prothorax with lateral carinae simple, weakly or unevenly crenulate; anterior angles projecting or acute; posterior angles of prothorax angulate. Pronotal surface generally uneven, punctation uniform or not with or without median glabrous areas; centre of disc usually bearing scales. Procoxae visible in lateral view. Hypomeron with or without scales, setose or glabrous; anterior portion rugose. Length of elytra 2.5-4× as long as pronotum or greater; disc with three simple carinae that are not beaded; sublateral keel absent; intercarinal space multipunctate; window punctures present and tuberculate; intercarinal scales of elytral disc variable, from very short and oval that may be countersunk within punctures to elongate with lengths at least 2.5× longer than wide; lateral carina simple; epipleuron hidden in lateral view. Hind wings present, fully developed; MP3 spur present (Kukalová-Peck and Lawrence 1993). Aedeagus  with parameres apically rounded to acute, inner outline between parameres bisinuate, weakly sinuate, or straight, length of parameres variable, median strut acute. Protibial edge spinate, mucro absent or weakly developed, spurs longer than tarsomere 2 with anteriormost spur greatly enlarged.
A description by   Biology. Both larva and adult are predatory. L. decorata and L. monilata were found in stems of Eucalyptus obliqua, where they were preying on larvae of Epithora dorsalis (Bashford 1994). The latter author notes a single-year life-cycle for the Leperina species (referred as Lepidopteryx). Adults have also been reared on Acacia dealbata (Bashford 1991 Remarks. The genus is apparently related to Anacypta and Xenoglena. The body is larger than in Anacypta but not so slender as in Xenoglena, moreover perfectly covered in scales. Further to the three species described, I have also encountered some undescribed species, all of them distributed in the Indonesian islands. Description. Body size: about 7.0-9.0 mm. Body shape flat. Gular sutures narrow, subparallel at apex. Frontoclypeal suture absent. Frons: longitudinal groove or depression absent. Cranium ventrally: tufts of long setae at sides absent. Submentum: ctenidium present. Antennal groove present. Eyes: size large, dorsal. Eyes number: four. Epicranial acumination absent. Lacinial hooks absent. Galea: shape clavate. Galea: ciliate setae absent. Mediostipes-Lacinia fused together. Palpifer: outer edge even. Mandibular apical teeth number: two, vertically situated. Mola absent. Penicillus (at base) long setae. Pubescence above mola or cutting edge absent. Ventral furrow absent. Basal notch shallow or absent. Labrum-Cranium not fused. Epipharyngial sclerite present. Lateral tormal process: projection curved downwards, processes not connected (Airora Remarks. As mentioned previously, six Phanodesta species from Juan Fernandez Island have been considered peculiar wingless beetles, with a form of elytral sculpture that is unknown in other trogossitids. Two analyses have shown a relationship for the genus within Gymnochilini, perhaps as a strongly-derived descendent of Australian Leperina . Most recently,  revised and redescribed the genus, established several new species from New Zealand, and combined some Leperina with Phanodesta species. They also discovered an uncommon distribution pattern comprising New Zealand (and adjacent islands) and Juan Fernandez Island.
Description (according to . Body size: 5.4-11.5 mm. Colour of body black or red-brown. Dorsal vestiture consisting of setae, scales or both. Head extending beyond anterior angles of pronotum. Frons more or less horizontal with mandibles visible in dorsal view. Median lobe of clypeus absent. Edge of labrum weakly to strongly emarginate or straight. Eyes entire. Gena acute. Supra-ocular scales present or absent. Antenna 11-segmented with loose club; rela-tive lengths antennomeres II and III variable; antennomere XI distinctly longer than wide and circular, about as long as wide or shorter. Prothorax with lateral carinae simple, or weakly to strongly and evenly or unevenly crenulate; anterior angles projecting or acute; posterior angles angulate. Pronotal surface even or uneven with impressions or shallow grooves; punctation uniform or not, with or without median area glabrous. Procoxae visible in lateral view. Hypomeron setose, glabrous, or bearing scales; anterior portion of hypomeron weakly to strongly rugose. Length of elytra 2.5-4× or less than 2.5× as long as pronotum; carinae present and usually beaded, with punctures located centrally within it, or adjacent and contacting carina; number of carinae variable, but usually 7-9; sublateral keel absent or present; intercarinal space apunctate; window punctures absent; intercarinal scales of disc never countersunk within puncture, elongate with lengths 2.5× longer than wide or ovate to circular with lengths less than 2.5× longer than wide; scales erect or not overlapping to strongly overlapping and adpressed; lateral carina simple; epipleuron visible or hidden in lateral view. Hind wings present and fully developed with MP3 spur present or vestigial and in the form of small buds, or absent. Aedeagus with parameres apically angulate, rounded or acute; inner outline between parameres bisinuate, weakly sinuate or straight; length of parameres and shape of median strut variable. Protibial edge smooth or crenulate; mucro absent or weakly developed; spurs longer than tarsomere 2 with anteriormost protibial spur greatly enlarged. Bursa and spermatheca bulbous, spermatheca about one fifth the size of the bursa and bearing a small tubulate spermathecal gland. A description by  based on P. cribraria: Body shape elongate. Gular sutures reduced. Frontoclypeal suture absent. Frons: longitudinal groove or depression absent. Cranium ventrally: tufts of long setae at sides present. Submentum: ctenidium absent. Antennal groove present. Eyes: size flat. Eyes number: two. Epicranial acumination deep. Lacinial hooks absent. Galea: shape clavate. Galea: ciliate setae absent. Mediostipes-Lacinia fused together. Palpifer: outer edge even. Mandibular apical teeth number: two, vertically situated. Mola reduced but present. Penicillus (at base) present (fine, often membranous). Pubescence above mola or cutting edge absent. Ventral furrow ciliate. Basal notch shallow or absent. Labrum-Cranium not fused. Epipharyngial sclerite present. Lateral tormal process: projection curved downwards, processes not connected (Airora Remarks. Schawaller (1993) synonymized Seidlitzella with Leperina. However, he based his observations on a comparison between Seidlitzella procera and two Palaearctic Leperina species, L. squamosa and L. tibialis. Australian species are often very different, although some their morphological details may also be similar (L. decorata is the type species of Leperina). Recently,  have established the new genus Kolibacia for the Palaearctic Leperina squamosa and L. tibialis. However, as their paper centred chiefly on Phanodesta, a differential diagnosis between Seidlitzella and Kolibacia was not addressed in detail. The main differences are explained in "A key to the species" of Gymnochilini, below.
Biology. Predatory. Adults found on logs of various trees (e.g. the fir Abies cilicia), larvae found under pine bark (Schawaller 1993 : 364. Nikitsky, N. B. 1992 Remarks. Outer habitus resembles the jewel beetles (Buprestidae), especially those of the genus Chrysobothris. Anacypta asahinai Kono, 1938 was combined with Xenoglena by . The combination is in accord with the opinion of Nikitsky (1992) and also my own independent study of the species. Nikitsky (l.c.) moreover suggested its synonymization with Xenoglena quadrisignata.
Biology. Predatory. Adults dwell on fallen trees and dry branches, hunting for xylophagous insects. They fly and run at great speed and appear very like some jewel beetles in body shape.
Distribution. Indonesia, Malayan Peninsula, Russian Far East, Japan, northern China. A large body of material of perhaps-undescribed species is known to me from northern Laos.
There are also some genera that are not included in the two character analyses because of insufficient data sets, namely Dupontiella, Elestora, Eupycnus, Euschaefferia, and Parallelodera. The classification of all these rather advanced genera within Trogossitini is undeniable, apart from the monotypic Elestora which is obviously related to Melambia, for which the systematics are quite complicated and in need of revision.
Most of the members of Trogossitini lead the kind of life typical of predatory Cleridae, especially of the subfamilies Clerinae and Tillinae. Adults hunt for xylophagous insects (e.g. Curculionidae: Scolytinae, Bostrichidae) on branches and logs while larvae dwell and hunt under bark or in galleries. However, some trogossitine adults live in insect galleries together with their larvae (e.g. Nemozoma). The trogossitins are not as efficient in the air as the gymnochilins, and neither do they move so swiftly on the ground.  horizontally situated. Lacinia mandibulae absent. Mola absent. Maxillary palpi 3-segmented. Cardo: size much smaller than stipes. Labial palpi 2-segmented. Prementum in single part. Antennal joints 1 and 2 elongate. Sensory appendix very small. Thoracic sclerites pattern (dorsally) 1-2-2. Thoracic sclerites pattern (ventrally) 3+1+1. Abdominal segment IX not divided. Tergite IX flat. Urogomphi present, hooked; median process absent.
Distribution Remarks. The phylogeny of the genus is unclear and in need of revision, together with Melambia. Elestora could be related to some species of the latter genus (see "Remarks" on Melambia). I presume a classification of Elestora within the trogossitins rather than with the gymnochilins.
Biology. Predatory. It was recorded from the galleries of the bark beetle Pseudothysanoes burtoni and has also been reared on the plant Vachellia farnesiana   : 364. Mamaev, B. M. 1976: 1652. Reitter, E. 1876: 24. Remarks. The placement of the genus in Trogossitini should be revised because my analysis of 2008 disclosed a possible relationship of some its species with Gymnochilini. There are distinct differences in the body shape among the numerous species of Melambia, for example between M. grandis (a robust species with a cordate pronotum) and M. orientalis (an elongate species with pronotum shaped somewhat like that of Tenebroides). Consideration of a re-classification of Seidlitzella within Gymnochilini, similar in habitus to Melambia, needs species revision and new phylogenetic analysis with reference to Trogossitini and Gymnochilini, including special attention to the related trogossitine genus Alindria. As a preliminary opinion, I assume that both genera, Melambia and Alindria, form a basal group of Trogossitini.
Description Biology. Predatory. According to , M. tekkensis and M. cardoni larvae prey on larvae of jewel beetles and longhorn beetles (under the bark of, for example, apricot trees or Grewia).
Distribution. South-eastern, southern and central Asia; also several species in Africa from Egypt to South Africa. Such a disjunctive distribution is possible, but the African species need to be checked because of possible confusion with the similar genus Alindria.

Description (T. dubius): "Ein 8 mm langer Käfer von ähnlicher Gestalt wie Parnidium Geinitzi. Der Prothorax is aber anders geformt un nähert sich mehr der Kreisform, Auch der Kopf scheint anders gewesen zu sein. Flügeldecken punktiert, 3,5 mal so lang als breit. Geinitz vergleicht diese Form mit Latridiites Schaumi, mit dem sie allerdings auch einige Ähnlichkeit hat."
This is potentially the oldest fossil record of Trogossitidae. Unfortunately, its description is inadequate and the illustration very poor. It is unclear to me why two more Mesozoic species from Russia were assigned to this dubious genus. It was not indicated in the original papers describing the two new species if the type of T. dubius was studied in situ (Ponomarenko 1985. The specimen appears to be housed in Ernst Moritz Arndt University of Greifswald, Germany. (A facsimile of the original description and illustration in Figs 24, 25.) Description (T. glabrus, translation from Russian). "Head sligthtly longer than wide, narrowed in front of eyes; eyes relatively large, situated at sides of head; cheeks [genae] short; temples [tempora] slightly shorter than eyes. The first antennomere large, second transverse, third 1.5 times longer than second, fourth to seventh as long as wide, eighth longer than wide, nineth to eleventh asymmetric. Pronotum with lateral margins rounded, its corners not acute, 1.5 times longer than wide. Front coxae not large, spherical, with exposed trochantins. Prosternal 3.0, width 1.0 mm)." (Ponomarenko 1990: 74 Remarks. This fossil differs from all other Mesozoic Trogossitidae described to date. If Martynov (1926) interpreted the shapes of the head and antennal segments well, it is the first known member of Trogossitidae without a distinct antennal club and with the head narrowed towards its base. The following features of Trogossitidae appear in the fossil: (1) general shape and size of body, (2) distinctly flattened sides of pronotum and elytra, (3) double rows of punctures/tubercles among elytral carinae, (4) robust bidentate mandibles, (5) extremely large scapus, and (6) dilated antennal segments with what are perhaps sensorial fields in the enlarged parts of each segment. The classification within Trogossitinae is based on the presence of the sensorial fields in the enlarged parts of the antennomeres alone. Small tubercles occurring in pronotum and elytra are known in trogossitine genera Calitys and Phanodesta only; no peltine representative possesses such structures. The tribe differs from the recent and fossil members of Trogossitinae in broadly oval body (this occurs in some Gymnochilini only), pronotum narrowed anteriad, antennae without conspicuous club and asymmetrical segments in flagellum, head narrowed towards base. The shape of antennal segments 10 and 11 is unknown because they are missing (only a trace of segment 10 is visible). The antennae may be only 10-segmented with the last segment enlarged (as in e.g. Egoliini). The new tribe is probably isolated from other tribes of Trogossitinae and may be considered a sister group to them. The ventral part of the fossil is unfortunately unknown, so a classification of Lithostoma remains uncertain, chiefly based on the distinctly enlarged scapus.
On the other hand, the concept of Lithostomatini was justifiably called into question by Yu et al. (2012) who argued that insufficient morphological information existed for the establishment of a higher taxon. † Genus Lithostoma Martynov, 1926 http://species-id.net/wiki/Lithostoma Map 6 Martynov, A. V. 1926: 13 (in Russian) (Martynov 1926: 32.) Original description of the species. English text: "Head and antennae as in generic description; sides of pronotum convex, points distinct; elytra broad, rounded at the posterolateral margins; marginal dilatations rather broad; the dividing stripes not elevated. Length of the body 6 mm." (Martynov 1926: 32) Translation of Russian text: "Head free, strongly projecting anteriorly (partly artificial condition in compressed specimen). Mandibles robust, left mandible bidentate (right inconspicuous). Antennae as described above, each joint weakly dilated at apex; each dilated area with thin [sic], dark, round rim. Final two joints torn off, only a trace of joint 10 present. Pronotum widened towards base, with shallow punctures interspersed with small tubercles. Elytra wide, rounded apically and dorsally [sic]; flattened sides well-developed and probably lighter than dark brown convex portion of elytra. Elytra, including flattened sides, with rows of well-developed small, black tubercles [orig.: "convex punctures"]; carinae among them not higher than tubercles. Length of body from anterior margin of labrum to apex of elytra -6 mm." (Martynov 1926: 13.) Note: The Russian and English texts vary somewhat from one another; the Russian is more comprehensive.
Distribution. Yu, Y., Leschen, R. A. B., Ślipiński, A., Ren, D. & Pang, H. 2012: 246. Type genus. Sinopeltis jurassica Yu, Leschen, Ślipiński, Ren & Pang, 2012 Remarks. This genus, containing two species, has only recently been established. The fossils are well preserved, with both part and counterpart. The species are relatively large, body shape perfectly appropriate to Peltinae or Lophocaterinae. The eyes are distinctly elevate, much more so than those in extant Ancyronini. The 3-segmented antennal club of S. jurassica is "weakly asymmetrical" (quite symmetrical in the original picture), that of S. amoena is "strongly asymmetrical" (inconspicuously so in the picture). The mesocoxae appear contiguous (unknown state in Trogossitidae) in S. jurassica, whereas they are narrowly separated in S. amoena. Both discrepancies mentioned may be the results of different positions of body parts (coxae, antennae) assumed during fossilization. Unfortunately, neither the ends of the tibiae nor the tarsi and mouthparts are described in either species, which leads to a lack of direct evidence for classification. Both S. jurassica and S. amoena are about 165 million years old and belong, if their classification is correct, together with species of Thoracotes, among the oldest known fossil members of Trogossitidae.
Remarks. The single genus Peltis exhibits a noteworthy mixture of the both advanced and primitive morphological features. However, some of the derived, especially larval, character states may considered as various kinds of reduction. A few species of Peltis are highly adapted to a hidden way of life under tree bark or in rotten wood: they are flattened and slow-moving, their robust mandibles have distinct mola, and the larval urogomphi are strongly reduced. The outer appearance is similar to that in Calitys, while some details of both adult and larval morphology (for example the gular appendages in the larval cranium) resemble Thymalus. The synonymization of Zimioma with Peltis is undeniable. Apart from body size, there is no morphological character to distinguish between the two genera.  Bacianskas, V. 2009: 30 (biology). : 24. Barron, J. R. 1996. : 76. Kolibáč, J. 2006: 111. Kolibáč, J. 2007a: 366. Noreika, N. 2009. Spahr, U. 1981: 74 (amber and copal fossils).

Genus
Larva: Frontal arms curved (cucujoid). Epicranial stem reduced. Endocarina present. Gular sutures inconspicuous. Gula: anterior apodemes present. Paragular sclerites absent. Hypostomal rods absent. Stemmata number: five. Mandibular apical teeth number: two, horizontally even, vertically situated. Lacinia mandibulae tridentate. Mola absent. Maxillary palpi 3-segmented. Pedunculate seta absent. Mala simple. Mala: bidentate protrusion present. Ligula present. Labial palpi 2-segmented. Antennal joints 1, 2 transverse. Sensory appendix larger than half of joint 3. Thoracic sclerites pattern Biology. Fungivorous. Crowson (1964b) noted adults and larvae from Great Britain: Adult and larva fungivorous, larvae feed beneath the thin and fleshy fruiting bodies of the basidiomycete Phlebia radiata Fr. of the Meruliaceae, which occurs on the dead wood of various deciduous trees (oak, beech, hazel), occasionally also conifers (pine). Adults are active in the warm days of autumn and winter (approximately, from late September to March). They have not been observed outside that period. They can be collected by sweeping from dry or decaying branches. Larvae may be found at all seasons, under the fruiting bodies of the fungus or under bark in spring and summer. Wielink et al. (2010) observed the species in the Netherlands and found adults active only by night. They live together with larvae on dead oak branches infested by the fungus Peniophora quercina.
"Maxilla with galea and lacinia largely fused; erect setae among pubescence of upper surface; metendosternite with elongate oblique arms, without lamina; elytra with pattern of light and dark patches" (ex Crowson 1966).
Biology. R. apterum was found in Nothophagus and Podocarpus leaf-litter at an altitude of 900 m (Crowson 1966 Kolibáč, J. 2006: 116 (phylogeny). Description (according to Crowson 1966, modified). Body size: 1.3 mm. Adult: Body conglobate. General form short ovate and very convex, Byrrhid-like; length not more than 2 mm. Head strongly transverse; front margin of clypeus with a marked emargination; galea and lacinia separate; lacinia without spines or hooks; no erect setae on upper body surface; metendosternite with slender lamina. Front coxal cavities internally closed, coxae very elongate, separated by a very narrow process; first ventrite with a median keel at least in front; ventrites II-IV each with a transverse line of strong backward-projecting setae near its hind border; tegmen undivided.
Biology. Collected in "male flower of Pinus insignis" (Crowson 1966 Biology. The single known larva described (Crowson speculated that it might be identified as Rentonium daldiniae) was extracted from a forest litter sample (mainly Nothofagus) at an elevation of about 500m (Crowson 1966). An adult R. daldiniae was found in Daldinia sp. fungus growing on a dead tree (Crowson 1966 Crowson, R. A. 1970: 7. Kolibáč, J. 2005: 81 (redescription) daldiniae Crowson, 1966; New Zealand (RAC) Crowson, R. A. 1966: 121, 123 (supposed larva). Crowson, R. A. 1970: 7. Kolibáč, J. 2005 : 32. Barron, J. R. 1971: 35. Crowson, R. A. 1964a: 296. Kolibáč, J. 2005. Kolibáč, J. 2006: 111 (phylogeny). : 366. Nikitsky, N. B. et al. 1998. : 83. Reitter, E. 1876: 64. Thymalops Iablokoff-Khnzorian,1962 [Type species: Cassida limbata Fabricius, 1787] : 35. Iablokoff-Khnzorian, S. M. 1962 Remarks. Comparing the larvae as well as adults of Thymalus and Protopeltis, I found some interesting similarities, which led me to consideration of their phylogenetic relationship. Later character analysis  showed a relationship of Thymalus and Protopeltis with the former Rentoniini Crowson, 1966. This in turn led to the establishment of the tribe Thymalini for the group. However, Crowson (1966Crowson ( , 1970 also associated the former monotypic tribe Protopeltini Crowson, 1966 with the rentoniins. Recently, such a classification was called into question by  and Leschen et al. (2012), who found Trogossitidae polyphyletic in their character analyses; however, both analyses were based on the same character states. Their model genera Thymalus and Rentonellum are classified outside Cleroidea in Lawrence et al. (2011) trees whereas Leschen et al. (2012), using a restricted character set, removed them only from Trogossitidae and/or suggested subfamily rank for rentoniins again, without necessarily believing in a mutual relationship between the two genera. Some more detail appears in "Remarks" with the tribe Thymalini. Léveillé (1877) described the Caucasian species Thymalus aubei as T. fulgidus var. aubei Léveillé, 1877. However, Thymalus fulgidus Erichson, 1844 was originally described from North America and  synonymized this species with T. marginicollis Chevrolat, 1842. That is perhaps why the latter author also synonymized the taxon aubei as a synonym of marginicollis, probably without examination of the holotype or even Caucasian specimens. Russian entomologists, for example Nikitsky et al. (1998) Biology. The beetles are not associated with any particular tree species and are found on both deciduous and coniferous trees. Thymalus limbatus is known from the trunks of birch, beech, linden, and spruce, mostly under bark. It is assumed that the larvae feed on fungi in rotten or decaying wood . Thymalus marginicollis has been collected from the fungi Polyporus betulinus, P. versicolor, Daedalea confragosa, on the trunks of birch and also on "wild flowers in plant press" .
Distribution. Holoarctic: Northern states of USA, Canada, Europe, North Africa, Siberia to China and Japan. Some specimens, probably a new species, have recently been collected in Chinese Sichuan and Yunnan and also in northern Thailand.

Key to tribes of Lophocaterinae
Remarks. The fossil is the oldest known record confirmed for the entire superfamily Cleroidea on the European continent. Due to the fine state of preservation, certain morphological character states of the fossil were inserted into a character matrix of Trogossitidae genera. The resulting tree reveals the basal position of Cretamerus vulloi within the lophocaterine clade. It may form an extinct branch of the recent Decamerini.
In view of its fine state of preservation, certain morphological character states of the fossil to be inserted into a character matrix of Trogossitidae genera. The resulting tree reveals the basal position of Cretamerus vulloi within the lophocaterine clade. It may form an extinct branch of the recent Decamerini.
Remarks. Since it was established , Decamerinae has always been connected with Peltidae (= Peltinae) in the works of a variety of authors. Unfortunately, the single larva described to date  was not associated with adults. It possesses some general features of Lophocaterinae (mandible with long lacinia mandibulae, cranium with characteristic gular area) but it differs in having curved (cucujoid) frontal arms and in the absence of a median process between the urogomphi. The larva of Eronyxa expansus is evidently lophocaterine ); however, the adults of Eronyxa and the present decamerins are similar in morphology and chiefly of floricolous habit. Two character analyses  placed Eronyxa on the border between Lophocaterini and Decamerini. Although the latter genus is classified within Lophocaterini herein, I feel its true relationship lies with Decamerini. This issue should be re-examined, although not before an indisputable larva of Chilean Decamerini is found and/or reared.  . Crowson, R. A. 1964a: 291 (key). : 47. Kolibáč, J. 2006.

Key to genera
Remarks. In consideration of a recent re-classification of Eronyxa within Lophocaterini , the systematic position of Antixoon needs to be checked. However, the bifid tarsal claws correspond with    : 28. Crowson, R. A. 1964a: 291. Kolibáč, J. 2005. Kolibáč, J. 2006: 111 (phylogeny : 27. Crowson, R. A. 1964a: 291. Kolibáč, J. 2005. Kolibáč, J. 2006: 111 (phylogeny Biology. The species occur on flowers. Pollen grains were found in the gut of Diontolobus punctipennis  Biology. The species were collected by beating branches and sifting rotten wood and litter. Detritus and some insect remnants were found in the gut of A. dwesae, whereas only insect fragments were found in the gut of A. ciskeiensis. The species are probably predatory and partly or occasionally fungivorous. Afrocyrona dwesae may be unable to fly.
Distribution. Three described species are known from South Africa: Eastern Cape and Transvaal. One or two more species have been recently found in the island of Sokotra.
Biology. Predatory. Adults can be beaten from dry branches or individually collected on fallen timber, where they hunt for other insects.
Distribution. Africa in the south of Sahara, eastern Asia (from India to Japan and Russian Far East), Australia: Queensland, New South Wales (types of A. aegra, A. amica, A. latebrosa, A. laticeps, A. vesca checked), New Guinea (including undescribed species). Strangely enough, one Japanese species, A. japonica, was introduced to Europe and first recorded in Hungary ("Visegrad 1904"). It has now been collected several times in adjacent Slovakia, on branches of decaying oak. Two Tertiary fossil species are known from European Eocene.
Species groups. In a review of Afrocyrona and Ancyrona, I divided the latter genus in several informal species groups . They are listed below together with a short diagnosis, although not all Ancyrona species are classified within the particular groups. This is only the first attempt to classify the rich and complex genus, so a monophyly of these groups should be checked and tried, then the groups may be established at the rank of subgenera). Biology.  found "only detrital material and vegetable fibres" in the gut of adult G. varians; however, there were "insect fragments as well as detrital material" in the gut of a larva of the same species. I assume that the adults and larvae are predatory.
Distribution. New Zealand. Biology. The species are probably predatory. According to Matthews (1992), they live in dry sclerophyll and Eremaean zones.

Species
Distribution. The genus is autochthonous in Australia; N. squamata from the Phillippines is probably mislabelled, misidentified or introduced (I did not examine the species). Recently, I studied a specimen of Neaspis cf. variegata collected in Brazil   Schmied, H. et al. 2009: 26. Remarks. Sinosoronia might be related to another Mesozoic genus, Peltocoleops. The latter genus was described as "Cleroidea incertae sedis" ) and classified within Lophocaterini by myself . The two genera differ distinctly in the shape of the antennal club. This is compact and 3-segmented, with segments weakly asymmetrical in Peltocoleops but loose, 2-or 3-segmented, with segments distinctly asymmetrical in Sinosoronia. It is therefore suggested that the latter genus be classified within the tribe Ancyronini, which has corresponding features in recent representatives. The large, elevated eyes observed in Sinosoronia also support such a classification (according to Kolibáč and Huang 2008).
The "posterior femur" in the original description is probably the hind coxa. The long antenna with a loose club resembles that of species of the Ancyrona gabonica species-group, while a similar shape of the pronotum may be found in the colobicoides species-group. Such an extremely small size of body is not known in recent Ancyronini but occurs in an concurrently described species from the late middle Eocene . Apart from body size, the two species share large, elevated eyes and similar shape of pronotum. The time difference between these two very similar species is about 100 million years, much more than between the Eocene and the present time. Round body and body size might appear indicative of a group of the rentoniine genera. However, the body is much smaller (about 1 mm) and the antennae shorter with a symmetrical club in the rentoniins . If the asymmetrical club is considered an apomorphy, Sinosoronia may well be an ancestor of Ancyronini rather than Thymalini (according to Kolibáč and Huang 2008 Barron, J. R. 1971: 11, 12 (syn. Lophocateridae = Peltinae). Barron, J. R. 1975: 1119: 119 (Lophocateridae). Kolibáč, J. 2006: 128 (diagnosis, stat. n.). : 365. Kolibáč, J. 2010: 35. Lafer, G. Sh. 1992 (key). Lawrence, J. F. & Newton, A. F., Jr. 1995: 868 (Lophocateridae). Lucht, W. 1998: 207 (key). Ślipiński, S. A. 1992: 442 (Lophocaterinae).
Remarks. The main issue to be addressed for Lophocaterini is their possible paraphyly in relation to Ancyronini. The whole clade (lophocaterins + ancyronins) is monophyletic but the lophocaterins might be paraphyletic (i.e., non-holophyletic in the traditional Hennigian meaning) because ancyronins can only be advanced members of more primitive lophocaterins. See also "Remarks" in the Ancyronini section. Further, more detailed study is required to resolve the question. The generic composition of Decamerini and its position within Lophocaterinae should be examined along -not, however, before an associated larva of the decamerins is known.
Floricateres Crowson, 1970[Type species: Floricateres pusillus Crowson, 1970] Crowson, R. A. 1970: 10. Kolibáč, J. 2005. Description. Body size: about 3.5 mm. Body shape flat. Gular sutures wide, subparallel. Frontoclypeal suture broadly emarginate. Frons: longitudinal groove or depression absent. Cranium ventrally: tufts of long setae at sides absent. Submentum of males: ctenidium present. Antennal groove present. Eyes: size moderate. Eyes number: two. Epicranial acumination moderate. Lacinial hooks absent. Galea: shape very small. Galea: ciliate setae absent. Mediostipes-Lacinia not fused. Palpifer: outer edge even. Mandibular apical teeth number: two, horizontally situated. Mola absent. Penicillus (at base) absent. Pubescence above mola or cutting edge absent. Ventral furrow absent. Basal notch moderate. Labrum-Cranium not fused. Epipharyngial sclerite absent. Lateral tormal process: projection projection reduced or absent (Promanus Biology. Peltonyxa pusillus was collected on the flowers of Bursaria (Crowson 1970). South Australian species occur in the sclerophyll and Eremaean zones (Matthews 1992 Similarities of Trichocateres with Lophocateres and Indopeltis: the mandible with membranous penicillus and distinct prostheca; the wing venation with cross-veins MP3-4 and AA1+2-3+4 absent; and the metendosternite with robust stalk and widely separated anterior tendons. It resembles Indopeltis in aedeagus with projecting phallobasic apodeme, eyes similarly shaped, relatively large and situated dorsally, and lateral edge of pronotum undulating, whereas Lophocateres parallels include labrum with tormal processes branched at base and maxilla with mediostipes not fused with lacinia. Excluding the characters mentioned, Trichocateres differs from all three abovementioned genera chiefly in tibial spur pattern 1-1-1, two sharp grooves in prosternal process and tufts of long hairs on elytra and pronotum. (Genus diagnosis after  Biology. The circumstances of collection are not exactly known; the specimens were knocked down from branches or fallen timber. Remnants of insect cuticle were found in the gut of the Assam specimen, the remains of an insect larva in the gut of the Laos specimen.
Distribution. India: Assam, northern Laos. Note: I studied only one non-type specimen determined as C. africana in the Museé d'Histoire Naturelle in Geneva. The specimen does not belong in Cleroidea.
Note. I have not studied any specimens of the species. The autochthonous distribution of Latolaeva or Ancyrona in South America is unprobable, although introduction is possible. Species of the two genera could be misidentified for an autochthonous or introduced Australian Neaspis or Peltonyxa (see "Distribution" for Neaspis). Remarks. This beetle was originally described in Trogossitidae and later removed from the superfamily as Coleoptera incertae sedis.

Ostoma australis
Original description of the genus. "A broadly elliptic beetle of small size; pronotum broadly hemiorbicular (hemispherical?); elytra short and broad, ornamented with many longitudinal striae; legs short, three pairs of coxae clearly separated, fore-coxa transverse, mid-coxa rounded, both posterior coxae transverse and connected with each other; abdomen with 5 visible sternites." (Lin 1985: 309) Original description of the species. "The body of a small beetle with head and legs missing, 3.5 mm long and 2.3 mm wide. Body broadly elliptic. Pronotum in hemiorbicular (hemispherical?) form, slightly broader than long; anterior margin of pro- Figure 19. Cretocateres mongolicus Ponomarenko, 1986. A reproduction of the original table.    notum concave, posterior margin of pronotum as wide as the anterior. Anterior coxa transverse and disjointed. Mid-thorax slightly smaller than metathorax. Both mid-coxae rounded and disjointed from one another. Posterior coxae transversely connected. Elytra much broader at base, gradually narrowing toward apex, rounded at apical angle; surface covered with several longitudinal striae. Abdomen with 5 visible sternites." (Lin 1985: 309) Species: † hyla Lin, 1985

phylogeny of the family Trogossitidae
Recently, several important studies have centred upon the phylogeny of Cucujiformia, including Cleroidea and Trogossitidae. Beutel and Pollock (2000) analyzed 20 larval head characters in 22 taxa (8 cleroid, 10 cucujiform, 4 outgroup) and advocated the monophyly of Cleroidea, but pointed out the paraphyly of traditional Cucujoidea. Trogossitidae were represented by Calitys and Temnoscheila, which genera were found in a sister relationship (Calitys was perhaps meant by the authors to be a representative of Peltinae). However, the inclusion of Phloiophilus into the trogossitids, as is presented herein, would be considered paraphyletic (but not polyphyletic) according to this study. Hunt et al. (2007) used an extensive molecular data set of dozens Coleoptera specimens. His Cleroidea are also monophyletic but with the inclusion of Byturidae/ Biphyllidae as a sister group of Phloiophilus. Trogossitidae are paraphyletic in the final tree: Trogossitinae + ((Lophocaterinae + Peltinae) + rest of Cleroidea). On the other hand, the most parsimonious tree of all 1880 studied beetle genera based on two mitochondrial genes shows the following phylogeny: [(Diplocoelus + Biphyllus) + (Byturus + (Phloiophilus + Priasilpha))] + [((Kolibacia + Temnoscheila + Nemozoma + Tenebroides) + (Ancyrona + Lophocateres + Grynocharis)) + (Peltis grossa + Peltis ferrugineum)] + (Cleridae + Melyridae sensu lato). This means that Trogossitidae (minus Phloiophilus) are perfectly monophyletic in the tree. It is also notable that Lophocaterinae are a sister group of Trogossinae there. (Note: Leperina is used for Kolibacia tibialis, Trogossita for Temnoscheila, Ostoma for Peltis ferrugineum in the mentioned paper.) The most recent system of Trogossitidae is based on my morphological studies of 2005 (adults) and 2006 (larvae). The most important result is a confirmation of both Crowson's ideas on the relationship of Calitys with Trogossitinae and Protopeltis with the Rentonium-group. On the other hand, his opinion about the relationship between the lophocaterins and the trogossitins is called in question. However, it is correct to say that larval characters actually connect the two groups while the majority of adult characters tend to defeat them in favour of a sister relationship of the lophocaterins and trogossitins. Both major clades were analyzed separately under equal weights and then with the use of successive reweighting . I employed 31 traditional morphological characters (16 adult, 15 larval) for 15 taxa (8 cleroid, 6 cucujoid, 1 derodontoid) in the analysis of 2008. Five cucujoid families (minus Phloeostichidae: Hymaea) were found to be monophyletic as well as Cleroidea. The group of cucujoid families (Cerylonidae, Coccinelidae, Endomychidae, Cucujidae, Byturidae, Phloeostichidae) were found to be paraphyletic with regard to the monophyletic Cleroidea. Trogossitidae were also found to be paraphyletic in this analysis. However, Phloiophilus was placed within a branch composed of trogossitid representatives, probably as a sister group of Thymalus. In the following analysis, Phloiophilus was analysed together with 43 trogossitid genera (88 characters of which 56 were adult and 32 larval). A strict consensus tree of 48 equally parsimonious trees indicates a possible relation of Phloiophilus with Peltinae and justifies independent subfamilial status for Lophocaterinae (the tree is reproduced herein, Fig. 26). It must be pointed out that 32 of 48 trees supported a sister relationship between the lophocaterins and the peltins. Only 16 trees supported a relationship between the lophocaterins and the trogossitins. This is perfectly consistent with the results of Hunt et al. (2007) and .
The most extensive modern paper on the beetle phylogeny based on morphological data is that by Lawrence et al. (2011). The authors used more than 500 characters and analyzed 366 genera of 165 beetle families. The final resulting tree is, in the section relevant to us, far different from that by Hunt et al. (2007) and it is beyond the scope of this contribution to analyze differences at superfamilial level. Trogossitidae are found to be paraphyletic (Acalanthis + Temnoscheila, Eronyxa + Grynoma are placed in separate branches), even polyphyletic because Thymalus is placed in a cluster together with Lamingtonium and the nitidulid families, while Rentonellum as a sister group of Smicrips together with Laemophloeus and Propalticus lie in a very distant part of the cladogram. Phloiophilus was not studied in the paper. Bocáková et al. (2011) used Peltis ferrugineum, Grynocharis oblonga and Temnoscheila japonica among outgroups in their molecular phylogeny of Melyridae sensu lato. Peltis and Grynocharis again form a sister group while the position of Temnoscheila is paraphyletic with regard to the rest of Cleroidea.
A work by Gunter et al. (2013) focuses on the molecular phylogeny of Cleridae. However, a relatively large set of trogossitid representatives were also studied. The resulting tree is very interesting: Phloiophilus is included in the trogossitine cluster Figure 26. A phylogenetic tree of Trogossitidae adapted and modified from original drawing by . Paraphyletic taxa denoted with the asterisk. Published with kind permission of Pro Entomologia, Basel, a publisher of Entomologica Basiliensia et Collectionis Frey.
between Larinotus, Leperina and Temnoscheila while Rentonellum, Peltis, Grynocharis, Ancyrona and Neaspis form a second cluster with a sister relationship to the rest of Cleroidea (similar to Hunt et al. 2007).
The most updated studies are those by the Tree of Life team and R. A. B. Leschen and co-authors, unpublished as yet. Their preliminary, mutually different, results were presented in the XXIVth International Congress of Entomology in Daegu (McKenna et al. 2012.
Although the outcomes of the works above differ in details, major ideas may be summarized as follows: 1) The superfamily Cleroidea is a monophyletic group.

2)
A part of the traditional Cucujoidea is probably paraphyletic with regard to Cleroidea. Cleroidea will probably, therefore, be extended to include several other "cucujoid" families.

3)
Lophocaterinae probably constitutes a sister group to Peltinae.

4)
Trogossitinae, as a basal group, may form a sister taxon to the major bulk of Cleroidea (although none of the molecular studies included the Metaxinidae-Chaetosomatidae-Thanerocleridae cluster, which might be in sister relationship with the trogossitines). This thesis, however, can hardly be justified by just the morphological evidence. If the subfamily Trogossitinae is actually confirmed paraphyletic, it should be classified at family rank again, as well as Peltidae composed of Peltinae and Lophocaterinae.

5)
The exact position of Phloiophilus is uncertain; it is a basal group of Cleroidea, probably related to Trogossitidae sensu lato.