Trogossitidae Latreille, 1802

Kolibac, Jiri, 2013, Trogossitidae: A review of the beetle family, with a catalogue and keys, ZooKeys 366, pp. 1-194 : 4-7

Family Trogossitidae Latreille, 1802

Trogossitidae Latreille, P. A. 1802: 110.

Ostomidae Hallan, J. 2007-2012: http://insects.tamu.edu/research/collection/hallan/test/Arthropoda/Insects/Coleoptera/Family/Trogossitidae.txt (check-list). Barron, J. R. 1971: 14. Boosten, G. 1983: 290 (biology). Bouchard, P. et al. 2011: 56 (review of higher taxa). Burakowski, B. et al. 1986: 116. Chûjô, M. & Lee, C. E. 1994: 187 (Korean species). Crowson, R. A. 1955: 82. Crowson, R. A. 1967: 211. Gourves, J. 2006: 56 (biology). Gray, D. W. 2002: 1583 (systematics). Hayes, J. L. J. et al. 2008: 206 (biology). Hieke, F. & Pietrzeniuk, E. 1984: 315 (Baltic amber). Hunt et al. 2007: 1915 (molecular phylogeny). Kireichuk, A. G. & Ponomarenko, A. G. 1990: 79 (Mesozoic fossils). Klimaszewski, J. & Watt, J. C. 1997: 43 (key). Kohnle, U. & Vite, J. P. 1984: 504 (biology). Kolibáč, J. 1993a: 20. Kolibáč, J. 1993b: 89. Kolibáč, J. 2004: (phylogeny). Kolibáč, J. 2005: 39 (morphology of adults). Kolibáč, J. 2006: 117 (morphology of larvae, phylogeny). Kolibáč, J. 2007a: 363 (Palaearctic beetles catalogue). Kolibáč, J. 2009: 127 (nomenclatory). Kolibáč, J. et al. 2005: 25, 129 (Central Europe, key). Kolibáč, J. & Leschen, R. A. B. 2010: 241 (review). Larsson, S. G. 1978: 150 (Baltic amber). Lawrence, J. F. 1982: 519. Lawrence, J. F. et al. 2011: 72 (phylogeny). Lawrence, J. F. & Britton, E. B. 1994: 118. Lawrence, J. F. & Newton, A. F., Jr. 1982: 281 (phylogeny). Lawrence, J. F. & Newton, A. F., Jr. 1995: 867 (review of higher taxa). Lawrence, J. F. et al. 1993: CD ROM (identification of larvae). Lawrence, J. F. et al. 1999a: CD ROM (identification of adults). Lawrence, J. F. et al. 1999b: CD ROM (identification of larvae). Lawrence, J. F. in Stehr F. W. 1991: 448. Leschen R. A. B. 2002: 263 (review, USA). Léveillé, A. 1910: 1. Lucht, W. 1981: 35. Luna de Carvalho, E. 1979: 80 (key). Majer, K. 1994: 384 (phylogeny, morphology). Merkl, O. 1993: 7 (key). Mitter, H. 1983: 52 (distribution). Nikitsky, N. B. 1980: 43 (key), 92 (larvae). Nikitsky, N. B. 1992: 80 (key). Ponomarenko, A. G. & Kireichuk, A. G. 2004-2008: http://www.zin.ru/animalia/Coleoptera/rus/paleosy2.htm (list of fossils). Paulian, R. 1998: 120 (key). Reitter, E. 1876: 3 (review, key). Reitter, E. 1922 (key). Schmied, H. et al. 2009: 23 (list of fossil taxa). Ślipiński, S. A. 1992: 440 (key to subfamilies, review of classification). Spahr, U. 1981: 74 (, list of fossils). Stresemann, E. et al. 1989: 270 (key). Tsinkevich, V. A. 1997: 27 (review). Valcarcel, J. P. & Prieto Pilona, F. 2001: 109. Waltz, R. D. 2002: 177. Weidner, H. 1993: 126 (key). Winkler, A. 1927: (Palaearctic beetles catalogue). Zherichin, V. V. 1978: 29 (Mesozoic fossils, Baysa).

Morphology

( Figs 1 , 2 ). Adults ( Fig. 1 ) (according to Kolibáč and Leschen 2010). Body size: 1.0-35.0 mm. Body wide, flattened in most Peltinae ; elongate in most Trogossitinae ; small and broadly oval in Larinotini and Decamerini ; convex in Thymalini . Body mostly bare or sparsely pubescent but sometimes also with tufts of setae, scales, dense pubescence or long hairs. Head usually not declined, although many members of Thymalini are moderately to strongly conglobate (the Rentonium group). Anterior margin of frons straight to very deeply emarginate ( Nemozoma ). Gular sutures widely or narrowly separated or strongly convergent. Posterior edge of epicranium with two incisions or evenly rounded. Frontoclypeal suture present or absent. Antennal insertions partly covered by edge of frons or visible in dorsal view. Subantennal groove conspicuous (receiving from one to three antennomeres), reduced or absent. Eyes not emarginate or (rarely) posteriorly emarginate, flat to elevated; sometimes divided by canthus into dorsal and ventral eyes (as in Gyrinidae and some Cerambycidae ). Labrum broadly oval to oblong, mostly slightly emarginate; epipharynx consisting mainly of a cordate sclerite; tormae connected, or not, at centre by tormal bridge. Antennae 8- to 11-segmented, with conspicuous 1- to 3-segmented club or with widened distal antennomeres that may be asymmetrical. Three to five apical antennomeres often bear sensorial fields. Mandible with one or two, rarely three, apical teeth; usually with ventral ciliate furrow; prostheca well-developed (brush of setae), reduced to absent; mola present, reduced or absent. Maxilla with distinct galea and lacinia; galea sometimes with ciliate or denticulate setae; lacinia with one to three apical hooks or with spine-like setae or with dense soft pubescence; basistipes more or less coalescent with lacinia or free; palpifer denticulate along outer margin (in Trogossitinae ); apical palpomere conical or cylindrical. Male submentum with tuft of setae in some Trogossitinae ; ligula membranous, rigid or coalescent with prementum; apex of ligula deeply or shallowly emarginate, often with ciliate setae; palpi conical or cylindrical, rarely weakly securiform. Tentorial arms connected by bridge or bridge reduced. Cervical sclerites present. Pronotum usually transverse; elongate only in some Trogossitinae . Lateral carinae almost always present, often denticulate; reduced only in some Trogossitinae ( Corticotomus ). Prosternal process apically dilated or narrowed. Procoxa slightly to strongly transverse. Coxal cavities internally open and externally closed or widely open. Notosternal suture complete. Trochantin elongate, exposed. Mesonotum distinct, with transverse scutum and well-developed scutellar shield. Elytra usually regularly punctate, with or without conspicuous carinae; epipleura well-developed or reduced in posterior half; elytra of Trogossitinae with interlocking mechanism along apical part of suture. Mesoventrite wide, with distinct prepectus. Mesocoxae usually projecting. Mesanepisternum triangular, not extending to mesocoxal cavity. Mesepimeron triangular, usually reaching coxal cavities, so that these are laterally open (cavities closed by meeting of mesoventrite and metaventrite in Egoliini ). Metaventrite more or less flattened, with distinct discrimen and transverse katepisternal suture; subcoxal lines present in Colydiopeltis , Larinotus , and Thymalus . Metanepisternum longitudinal, often with carina at centre. Metacoxae extending laterally to meet elytral epipleura, often with longitudinal furrow at centre. 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. Tarsi 5-5-5; 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 pre- and 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 ( Kolibáč 2006) 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 Kolibáč 2006). 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 Crowson (1964a) and synonymized with Peltinae by Kolibáč (2006) because of possible paraphyly of the latter subfamily. Later, in response to new observations, we suggested ( Kolibáč 2008, Kolibáč and Zaitsev 2010) 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)	Trogossitinae
-	Adult: labium with membranous ligula; epipharynx without cordate sclerite; antennal club weakly asymmetrical or symmetrical, terminal antennomeres (antennal club) without distinct sensorial fields; front coxal cavities mostly externally open (except for Lophocaterinae : Decamerini ); body often oval and flat (but sometimes also convex or conglobate); elytral suture without distinct interlocking mechanism. Larva: head capsule mostly without endocarina; gular sutures and hypostomal rods reduced; frontal arms often curved; gular region without paragular sclerites	2
2	Adult: frontoclypeal suture absent or inconspicuous; gular sutures wide, subparallel; eyes moderate, not distinctly elevate; antennal club symmetrical; radial cell oblong, moderate; tibial spines along sides mostly reduced; body flat, convex or conglobate. Larva: lacinia mandibulae tridentate, absent or minute. Fungivorous or phytophagous	Peltinae
-	Adult: frontoclypeal suture present, sometimes distinctly emarginate (or concave); gular sutures wide, convergent at apex; eyes almost elevate, laterally situated; antennal club weakly asymmetrical; radial cell moved downwards, towards wing centre, sometimes small or reduced; tibial spines along sides present; body always flat. Larva: lacinia mandibulae plumose, always distinct. Primitive members fungivorous or phytophagous, advanced ones floricolous or predatory	Lophocaterinae

Figures

Figure 1. Adult morphology: A Alindria sp. from Laos, ventral surface B Leipaspis lauricola, labium C Airora cylindrica, antennal club D Corticotomus cylindricus, mandible dorsally E Airora cylindrica, mandible ventrally F Acalanthis quadrisignata, labium G Acalanthis quadrisignata, maxilla H Acalanthis quadrisignata, wing I Peltonyxa deyrollei, tegmen composed of 3 parts J Airora cylindrica, tegmen composed of 2 parts.

Figure 2. Larval morphology: A Tenebroides " fuscus ", dorsal surface B Tenebroides " fuscus ", mandible ventrally C Lophocateres pusillus, mandible ventrally D Tenebroides " fuscus ", head ventrally E-F Peltis ferruginea, head capsule (E dorsally F ventrally) G Ancyrona diversa (1 dorsally 2 ventrally) H Peltis ferruginea (1 dorsally 2 ventrally).