New Genus and Species of Subfamily Trinodinae (Coleoptera, Polyphaga

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ZOOSYSTEMATICA ROSSICA, 19(1): 54–69 15 JULY 2010

New genus and species of subfamily Trinodinae (Coleoptera, Polyphaga, Dermestidae) from Lowermost Eocene French amber Новый род и вид подсемейства Trinodinae (Coleoptera, Polyphaga, Dermestidae) из нижнеэоценового французского янтаря

A.G. KIREJTSHUK, J. HÁVA & A. NEL

А.Г. КИРЕЙЧУК, Ю. ХАВА, А. НЕЛЬ

A.G. Kirejtshuk, Zoological Institute of the Russian Academy of Sciences, 1 Universitetskaya Emb., St. Petersburg, 199034, Russia. E-mail: [email protected], [email protected] J. Háva, Private Entomological Laboratory and Collection, 37 Rýznerova St., CZ-252 62 Únětice u Prahy, Prague- west, Czech Republic. E-mail: [email protected] A. Nel, CNRS UMR 5202, Muséum National d’Histoire Naturelle, CNRS UMR 5202, Entomologie, 45 Rue Buffon, F- 75005 Paris, France. E-mail: [email protected]

In the paper six new species of the genus Oisenodes gen. nov. (Dermestidae, Trinodinae, Trino- dini) are described: O. azari sp. nov., O. clavatus sp. nov., O. gallicus sp. nov., O. metepisternalis sp. nov., O. oisensis sp. nov. and O. transversus sp. nov. A new tribe Trinoparvini Hava, trib. nov. is established for the recent genus Trinoparvus Háva, 2004. Short review of known fossil records of the subfamily Trinodinae is given. В статье описываются шесть новых видов рода Oisenodes gen. nov. (Dermestidae, Trino- dinae, Trinodini): O. azari sp. nov., O. clavatus sp. nov., O. gallicus sp. nov., O. metepisternalis sp. nov., O. oisensis sp. nov. и O. transversus sp. nov. Устанавливается новая триба Trinoparvini Hava, trib. nov. для рецентного рода Trinoparvus Háva, 2004. Дается краткий обзор ископаемых находок жуков подсемейства Trinodinae. Key words: Lowermost Eocene French amber, taxonomy, Coleoptera, Dermestidae, Trinodi- nae, new tribe, new genus, new species Ключевые слова: нижнеэоценовый французский янтарь, систематика, жесткокрылые, кожееды, Coleoptera, Dermestidae, Trinodinae, новая триба, новый род, новые виды

INTRODUCTION atics of fossil record and historical develop- ment by Kirejtshuk et al. (2009). More de- The paper is the eighth contribution to tailed information on representation of this the knowledge on fauna of Coleoptera from coleopterous family in the fossil record can Lowermost Eocene French amber collected be got in the catalogue by Ponomarenko & in Oise falls (Batelka et al., 2006; Bílý & Kirejtshuk, 2007; Kirejtshuk & Nel, 2008, Kirejtshuk (2010). This subfamily is known 2009; Moseyko et al., 2010, in press; Kirejt- in fossils from Baltic amber (genus Trinodes shuk et al., 2010a, 2010b), which is devoted Dejean, 1821: Larsson, 1978; Spahr, 1981; to the families remained unknown from this Háva, 2003; Háva & Prokop, 2004, 2006; resource and poorly known in fossils at all. Háva et al., 2006, 2008) and Lebanese amber The family Dermestidae was provided with (Kirejtshuk et al., 2009). Among inclusions a short necessary review of data on system- of the Lowermost Eocene French amber

© 2010 Zoological Institute, Russian Academy of Scienсes A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER 55 about a hundred specimens of Dermestidae Tribe Trinodini Casey, 1900 and, among them, more than 30 specimens of the subfamily Trinodinae were recently Type genus: Trinodes Dejean, 1821. found and all of them belong to the genera or one genus, apparently rather related to Oisenodes gen. nov. those represented in the Recent fauna. Type species: Oisenodes gallicus sp. nov. Etymology. The name of this new genus MATERIAL AND METHODS is formed from the name of locality with Many specimens recovered among inclu- amber deposits and generic root ‘nodes’ of sions from the Lowermost Eocene French the type genus of the subfamily. Gender: amber are deposited in the Laboratoire de masculine. Paléontologie, Muséum National d'Histoire Description. Adult males. Body elon- Naturelle, Paris (MNHN). Ordinary optic gated oval, strongly convex dorsally and equipment was used for the examination, moderately convex ventrally; dorsum dif- in particular, a stereomicroscope Olympus fusely punctured, with long and stout hairs, SCX9 and an inverted microscope Olympus frequently of two types: longer and thicker CK 40 in MNHN, and also the stereomicro- suberect ones and others shorter, thinner scope Leica MZ 16.0 in the Zoological Insti- and subrecumbent; thoracic and abdominal tute, St. Petersburg. All holotypes and most underside with comparatively sparse and paratypes of the new species deposited in moderately long recumbent hairs; length MNHN, part of paratypes of Oisenodes gal- 1.3–1.7 mm. Head oval and strongly de- licus sp. nov. and O. metepisternalis sp. nov. clined, with very large and coarsely faceted are deposited in the Zoological institute of eyes, labrum exposed, antennae eleven- the Russian Academy of Sciences (St. Pe- segmented with three-segmented anten- tersburg) and one paratype of O. gallicus nal club; median ocellus on head present. sp. nov. in the Entomologické oddělení Pronotum with convex anterior and pos- (Národní museum, Prague, coll. J. Háva). terior bi-sinuate edges, paralateral carinae Type strata. Lowermost Eocene, in am- distinct. Elytra with separately rounded ber, circa 53 Myr, Sparnacian, level MP7 of apices and usually leaving apex of pygidi- the mammal fauna of Dormaal. um not covered. Prosternal process rather Type locality. Farm Le Quesnoy, Chev- wide and widely rounded at apex. Posterior rière, region of Creil, Oise department edge of metaventrite between coxae almost (north of France). straight. Metacoxae moderately separated. Tarsomeres 1–4 very narrowly lobed to RESULTS simple. The posterior wing are shown in Fig. 19 and 20. Order COLEOPTERA Adult females. External characters are Family DERMESTIDAE corresponding with those in males. Diagnosis. The new genus is similar to Subfamily Trinodinae Casey, 1900 the genera Trinodes Dejean, 1821, Apsectus This subfamily can be diagnosed by the LeConte, 1854 and Evorinea Beal, 1961, (sub) erect, long and stout hairs on dorsum, but differs from them in the comparatively 11-segmented antennae with 1–6-segment- widely separated pro- and mesocoxae, and ed club, pronotum with paralateral striae, also truncate posterior edge of the metaven- prohypomera without depressions for re- trite between metacoxae. Besides, the char- ception of antennal club, prosternum some- acters given in the key below can be used for what projecting anteriorly. the diagnostics of the new genus.

Key to genera of the subfamily Trinodinae 9(8)b. Prosternal process incomplete, truncate or emarginate apically and abutting ante- 1a. Antennae ten-segmented; body form very rior mesoventral process; posterior edge of small, subcircular; median lobe of male geni- metaventrite between coxae with small me- talia not separated from parameres (tribe dian emargination (recent) ...... Trinoparvini Háva, trib. nov.) (recent) ...... Apsectus LeConte, 1854 ...... Trinoparvus Háva, 2004 1b. Antennae nine- or eleven-segmented; median Notes. Tribe Trinoparvini Háva trib. nov. lobe of male genitalia separated from para- (type genus: Trinoparvus Háva, 2004 recent), meres ...... 2 except the diagnostic features mentioned in 2(1)a. Antennae filiform (tribe Thylodriadini the above diagnosis, is characterised by the Mroczkowski, 1954) (recent) ...... 3 very small body (1.1–1.2 mm) and three- 2(1)b. Antennae with club ...... 4 segmented antennal club. The name of the 3(2)a. Antennae nine-segmented (recent) ...... Thylodrias Motschulsky, 1839 tribe has been already mentioned at various 3(2)b. Antennae eleven-segmented (recent) . . . . web-sites. . . . .Trichodryas Lawrence & Slipinski, 2005 4(2)a. Body elongate ...... 5 Key to the Oisenodes species 4(2)b. Body oval ...... 7 5(4)a. Body form very elongate and strongly 1a. Ultimate antennomere at most slightly longer convex; antennal club three-segmented than two previous ones combined; integu- (tribe Cretonodini Kirejtshuk & Azar in ment more smoothed and more finely punc- Kirejtshuk et al., 2009) (Lebanese amber) . . . tured, and comparatively less pubescent . . . 2 ...... Cretonodes 1b. Ultimate antennomere 1.5–2.0 times as long Kirejtshuk & Azar in Kirejtshuk et al., 2009 as two previous ones combined; integument 5(4)b. Body slightly elongate; antennal club more coarsely sculptured and punctured, and four-six-segmented (tribe Trichelodini Pea- also comparatively more pubescent ...... 4 cock, 1978) (recent) ...... 6 2(1)a. Metepisterna strongly widened posteri- 6(5)a. Procoxae strongly projecting, without orly; prosternal process with more convex plates; anterior mesoventral process very posterior edge (Figs 6, 21–23) ...... short and acute (recent) ...... O. clavatus sp. nov...... Hexanodes Blair, 1941 2(1)b. Metepisterna subparalell-sided or slightly 6(5)b. Procoxae not strongly projecting, with widened posteriorly; prosternal process with well developed plates; anterior mesoventral only slightly convex posterior edge ...... 3 process long and narrowly rounded (recent) 3(2)a. Head slightly narrower than pronotum; ...... Trichelodes Carter, 1935 antennae not longer than head width; dis- 7(4)a. Antennal club one- or two-segmented in tance between procoxae more than width of male, two-segmented in female; posterior femora; pronotal base about twice as long as margin of metaventrite without small me- its apex (Figs 14, 24) . . . O. oisensis sp. nov. dian emargination (recent and Baltic amber) 3(2)b. Head moderately narrower than prono- ...... Evorinea Beal, 1961 tum; antennae somewhat longer than head 7(4)b. Antennal club three-segmented ...... 8 width; distance between procoxae less than 8(7)a. First abdominal ventrite usually with width of femora; pronotal base markedly less two short oblique lines; posterior edge of than twice as long as its apex (Figs 7, 9, 13, metaventrite between coxae with small me- 15, 29–30 ...... O. transversus sp. nov. dian emargination (recent and Baltic amber) 4(1)a. Posterior edge of pronotum about four ...... Trinodes Dejean, 1821 times as long as lateral ones; prosternal pro- 8(7)b. First abdominal ventrite usually without cess with shallowly emarginated posterior two short oblique lines ...... 9 edge; metepisterna subparallel-sided (Figs 9(8)a. Prosternal process complete, compara- 3–5, 19) ...... O. azari sp. nov. tively wide and fitting into cavity at ante- 4(1)b. Posterior edge of pronotum about three rior edge of mesoventrite; posterior edge of times as long as lateral ones; prosternal pro- metaventrite between coxae subtruncate and cess with more or less convex posterior edge; without small median emargination (Lower- metepisterna more or less widened at poste- most French amber) . . .Oisenodes gen. nov. rior edge ...... 5

5(4)a. Metepisterna about four times as long as different size. wide; ultimate antennomere more than 1.5 Paratype ‘PA 135’, probable male with times as long as two previous ones combined; widely rounded hypopygidium; the com- pronotum somewhat less than 2.5 times as plete specimen is included in a small irregu- wide as long (Figs 1–2, 17–18) ...... lar amber parallelepiped with some small gas ...... O. gallicus sp. nov. 5(4)b. Metepisterna about three times as long as vesicles and “milky cover” along pubescent wide; ultimate antennomere not more than integument; the inclusion is put in a circu- 1.5 times as long as two previous ones com- lar reservoir with Canada Balsam fixed on a bined; pronotum about 2.5 times as wide as microscope glass. long (Figs 8, 10–12, 16, 25–28) ...... Paratype ‘PA 15552’; the complete speci- ...... O. metepisternalis sp. nov. men is included in a small subcylindrical Notes on bionomy. Recent species of amber bar with some small gas vesicles and Trinodes seem to have the bionomy more or “milky cover” along pubescent integument, less similar to that in extant species of the and also with some very small pieces of or- new genus under consideration. Their adults ganic matter and irregular crack along body and larvae live amongst webs of tube- and plane; the amber bar with beetle is glued on sheet-web building spiders beneath loose a microscope glass by epoxy and on this glass dry bark on old trees, mainly of oak Quer- another bar with Diptera inclusion is glued cus, where they feed on the dead remains of at another side of this microscope glass. insects and spider exuviae. Adults of recent Paratypes ‘PA 12256’, male and female; species have been also found on blossom the complete two specimens are included in and foliage. a irregular bar forming a trapezium in cross section; specimens visible clearly but with optic aberration; the bar has clear consis- DESCRIPTIONS OF SPECIES tence, 9 mm of length, width of widest plane Oisenodes gallicus sp. nov. 7 mm and 2 mm of narrowest plane; with (Figs 1–2, 17–18) very small complete Nematocera and some pieces of organic matter and cracks. Material examined. Holotype ‘PA 2307’ Paratype ‘PA 12222’, female with mod- (MNHN), probable male with slightly erately rounded hypopygidium and partly exposed anal sclerite; the complete speci- exposed ovipositor; the complete specimen men is included in a small irregular amber is included in a somewhat irregular bar parallelepiped with some small gas vesicles forming a trapezium in cross section; speci- and “milky cover” along pubescent integu- men visible dorsoventrally; the bar has clear ment; the inclusion is put in a circular res- consistence, 7 mm of length, width of widest ervoir with Canada Balsam fixed on a mi- plane 6 mm and 2 mm of narrowest plane; croscope glass. with some pieces of organic matter. Paratype ‘PA 160’, probable female with Paratypes ‘PA 438’, male and female; the moderately rounded hypopygidium; the complete two specimens are included in a complete specimen is included near the sur- irregular bar forming a semicircle in cross face of one facet of the subcylindrical bar section; specimen visible at one cross side with visible underside faced to the surface (underside) and approached to facets along and surrounded by many gas vesicles, par- length (dorsum); the bar has clear consis- ticularly at head and prosternum; dorsum tence, 8–10 mm of length, width of plane invisible, because many of not transparent 6 mm; with a mummy of Nematocera and gas vesicles are located behind the beetle; some pieces of organic matter. the bar has 17 mm of length and 4–7 mm Etymology. The name of the new spe- of cross-section in different places and in- cies is derived from the ancient name of the cludes also many crashes and gas vesicles of country of origin of the inclusion.

Figs 1–2. Oisenodes gallicus sp. nov., holotype ‘PA 2307’. Body, ventral view (1) and dorsal (2) view. Length 1.3 mm

Diagnosis. This new species is very simi- – O. transvesus sp. nov. in the widely lar to O. metepisternalis sp. nov. and clearly separated procoxae, somewhat shorter an- differs from the latter only in the somewhat tennae with much longer ultimate antenno- longer pronotum and shape of metepisterna. mere, somewhat smoothed microsculpure Its antennal club has the ultimate anten- of integument with distinct puncturation, nomere somewhat longer than that in O. more raised pubescence and markedly nar- metepisternalis sp. nov. Oisenodes gallicus rower metepisterna. sp. nov., in contrast to O. metepisternalis Description. Length 1.3, width 0.7, height sp. nov., has the lateral sides of pronotum apparently about 0.4 mm. Elongate oval, more arcuate and also has somewhat less strongly convex dorsally and moderately raised and less conspicuous pubescence of convex ventrally; subunicolourous dark integument. Oisenodes gallicus sp. nov. dif- brown to blackish; appendages brownish; fers also from: dorsum with long and stout yellow, brown, – O. azari sp. nov. in the shape and pro- yellow greyish or greyish hairs (colouration portion of pronotum, comparatively smaller depending on lighting) of two types: longer head with smaller eyes, somewhat longer ones nearly 1.5 times as long as ultimate hairs (setae) on the dorsum and denser antennomere and more than twice as long and more conspicuous hairs on underside, as femora wide, thicker and erect; others coarser puncturation and microsculpture of somewhat shorter than ultimate antenno- integument; mere, thinner and subrecumbent; thoracic – O. clavatus sp. nov. in the shape and underside with comparatively sparse and proportions of antennomeres in its club, rather long recumbent hairs, but somewhat somewhat longer hairs (setae) on the dor- shorter than the shorter ones on dorsum; sum and denser and more conspicuous hairs abdominal ventrites with slightly conspicu- on underside, coarser puncturation and mi- ous recumbent hairs about half as long as crosculpture; those on thoracic underside. – O. oisensis sp. nov. in the somewhat Head and pronotum with distinct dense smaller head, shape of pronotum, markedly punctures, apparently slightly larger than more robust and much less convex body, eye facets in diameter, interspaces between and also in the longer ultimate antennom- them somewhat narrower than a puncture ere, less raised and less conspicuous pubes- diameter and smoothed. Elytra with larg- cence of underside; er and sparser punctures and completely

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 54–69 A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER 59 smooth interspaces with distance between rounded at apex. Mesocoxae apparently punctures as great as a puncture diameter. transversely oval, slightly projecting and Metaventrite and abdominal ventrites with very widely separated. Ratio between pro, extremely sparse and extremely small dis- meso- and metacoxae about as following: tinct punctures, interspaces between them 1.0:2.3:0.8. Metaventrite rather medially more or less smoothed, but punctures be- convex, posterior edge between coxae shal- coming larger distally. lowly arcuately excised. Distance between Head oval and strongly declined (hypog- metacoxae markedly greater than antennal nathous), slightly exposed dorsally, much club wide. Metepisterna moderately wide narrower than pronotum, with very large, and subparallel-sided at most length, but vertical and coarsely faceted eyes, somewhat widened at posterior end. Metacoxae more convex and frons strongly turned ventrally. or less strictly transverse (apparently not Labrum well exposed, short and transverse. oblique), with moderately raised and exter- Antennae slightly shorter than head wide, nally rather approaching to sides. First ab- apparently eleven-segmented with three- dominal ventrite about as long as 2nd–4th segmented club composing almost one-third ventrites combined and each of the latter of total antennal length, scape apparently comparable in length; hypopygidium some- rather small and elongate, ultimate antenno- what longer than previous ventrite. Epi- mere about one and two-third as long as two pleura rather narrow and elevated laterally. preceding antennomeres combined; anten- Legs well developed, moderately narrow nomeres covered with sparse, rather short and long, diffusely covered with compara- and very stout hairs. Pronotum somewhat tively short and stout setae. Tibiae rather narrower than combined elytral base, more compressed and scarcely widened apically, than twice as wide as long, strongly vaulted about four sevenths as long as antenna and at disk and rather steeply sloping at sides; its about a half as wide as antennal club. Femora anterior edge gently convex and posterior of usual shape and moderately compressed, one distinctly bordered and bi-sinuate, sides 2.0–2.5 times as wide as tibiae. Tarsi moder- looking nearly like continuation of anterior ately long, about as long as tibiae (althought edge, about one-third as great as posterior anterior ones somewhat shorter than the rest edge, paralateral carinae distinct. Scutellum ones), tarsomeres 1–4 very narrowly lobed transverse and subpentagonal, somewhat to simple, comparable in length, ultimate wider than long. Elytra about one and one- tarsomere somewhat longer; claws simple, third as long as wide combined, longest at narrow and not long, about one third as long suture and arcuate in posterior along sides, as ultimate tarsomere. rather convex along the middle and steeply Variations. Length 1.3–1.7, width 0.7– sloping and somewhat declined on ventral 0.9 mm. The paratype ‘PA 135’ is darker sides (from below visible lateral sides more than the holotype (body nearly uni- widely separated than distance between lat- colourous blackish), although because of eral edges), with weak shoulders, adsutural ‘milky cower’ the underside of it can not lines not visible. Pygidium with widely sub- be seen clearly. Most specimens examined truncate apex. have both dorsal and ventral integument Most part of head underside, proster- masked by pubescence with ‘milky’ cover, num and mesoventrite not visible because although the the paratype ‘PA 160’ demon- of declination of head and pubescence strates comparatively clear metaventrite with “milky cover”. Mouthparts not vis- and abdominal ventrites. Finally the para- ible. Procoxae apparently moderately large; types ‘PA 438’ look more slender and have distinctly transverse, slightly projecting more narrowing elytral apices, although downwards and comparatively widely sepa- this impression could be a sequence of op- rated; prosternal process wide and widely tic aberration.

Oisenodes azari sp. nov. (Figs 3–5, 19)

Material examined. Holotype ‘PA 2501’ (MNHN), probable female with moderately rounded hypopygidium; the nearly complete specimen with open left elytron and exposed unfolded posterior wings, but with missing antennae is included in a small irregular clear amber parallelepiped. The inclusion is put in a circular reservoir with Canada Bal- sam fixed on a microscope glass. Paratype ‘PA 1581’, probable female with moderately rounded hypopygidium; the complete specimen is included in an irregular bar forming a semicircle in cross section with a flat plane along whole length; specimen well visible laterally; the bar has clear consistence, 8–10 mm of length, width of longitudinal plane 6 mm. Etymology. The new species is named for Dany Azar who prepared many specimens in amber for this study. Diagnosis. The new species under consideration is very distinct among the congeners due to its emaginate posterior edge of prosternal process. This new species differs Figs 3–5. Oisenodes azari sp. nov., holotype ‘PA 2501’. Body, length 1.3 mm, dorsal (3) and ven- from O. gallicus sp. nov. and O. metepister- tral (4), view and antennal club, lateral view (5). nalis sp. nov. mostly in the shape and pro- Scale bar: 0.1 mm (5). portions of pronotum, comparatively larger head with larger eyes, somewhat shorter hairs (setae) on the dorsum and sparse and species of the genus, the most characters less conspicuous hairs on underside, much shared by both are omitted in the descrip- finer puncturation and more smoothed in- tion below. tegument. It differs from O. oisensis sp. nov. Description. Length 1.3, width appar- in the much longer ultimate antennomere, ently 0.7 mm. Elongate oval, strongly con- narrower metepisterna, very smoothed in- vex dorsally and ventrally; subunicolourous tegument with distinct puncturation and reddish; dorsum with long and stout yel- less raised pubescence. Besides, O. azari lowish to reddish hairs of two types: lon- sp. nov. differs from O. transversus sp. nov. ger ones nearly 1.5 times as long as femora in the much longer ultimate antennom- wide, thicker and erect; shorter ones some- ere, narrower metepisterna, smoothed in- what thinner and about as long as femora tegument with distinct punctures and less wide; thoracic underside with very sparse, raised pubescence, and also from O. clavatus thin, moderately long and very slightly con- sp. nov. in the shape and proportions of an- spicuous recumbent hairs, but much shorter tennomeres in its club, comparatively larger than the shorter ones on dorsum; abdominal head with somewhat larger eyes. ventrites with slightly conspicuous recum- Notes. Taking into consideration a great bent hairs less than half as long as those on similarity of the new species with the type thoracic underside. Dorsum with distinct

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 54–69 A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER 61 and very sparse punctures, about as large as matter, some small cracks and some small eye facets in diameter, interspaces between gas vesicles. them four–five times as great as a puncture Paratype ‘PA 1842, 1/3’, probable male diameter and smoothed. Metaventrite and with widely rounded hypopygidium; the ventrites with very small and sparse punc- complete specimen is included in a irregular tures, much smaller than facets in diameter, bar with two subparallel flat planes along interspaces between them rather great and whole length; specimen visible mostly lat- smooth. erally and ventrally and approached to the Pronotum somewhat narrower than bar facets at one of ends; the bar has clear combined elytral base, much more than consistence, 10–13 mm of length, width of twice as wide as long, its anterior edge widest plane 10 mm; at the specimen there subrectilinear and posterior one markedly is a crack along body plane and a crack in less than twice as wide as anterior one dis- sagittal plane at posterior edge. tinctly bordered and bi-sinuate, sides nearly Paratype ‘PA 164, 2/3’; the complete spec- straight and rather short (about one quarter imen is included in a irregular bar with two of posterior edge), paralateral carinae dis- subparallel flat planes along whole length; tinct. Elytra about one and one third as long specimen with partly exposed posterior wing as wide combined. Pygidium with widely visible laterally and disposed between the flat subtruncate apex. Prosternal process wide planes; the bar has different layers and some and widely emarginate at apex. Ratio be- cracks, 8–9 mm of length, width of widest tween pro-, meso- and metacoxae about as plane 9 mm; at ventral side of posterior part following: 1.0:2.0:0.8. Metepisterna moder- of body there is a gas vesicle. ately wide and subparallel-sided, about four Etymology. The name of the new species times as long as wide. First abdominal ven- refers to the structure of its peculiar anten- trite about as long as 2nd and 3rd ventrites nal club. combined; hypopygidium about 1.5 times as Diagnosis. This new species differs from long as previous ventrite. Femora of usual all congeners in the shape of metepisterna, shape and moderately compressed, 2.0–2.5 and also from: times as wide as tibiae. – O. gallicus sp. nov. and O. metepister- Variation. Paratype with body length nalis sp. nov. in the shape and proportions of 1.3 and height 0.6 mm; brownish, with antennomeres in its club, somewhat shorter somewhat coarser puncturation and more hairs (setae) on the dorsum, sparse and less conspicuous pubescence on underside. An- conspicuous hairs on underside, much finer tennae apparently shorter than head wide, puncturation and smoothed sculpture, and eleven-segmented with three-segmented from the latter also in the shape and width club composing almost one third of total an- of the metepisterna; tennal length, ultimate antennomere about – O. azari sp. nov. in the shape and pro- one and one third as long as two preceding portion of antennomeres in its club, com- antennomeres combined. paratively smaller head with somewhat smaller eyes, convex posterior edge of pros- Oisenodes clavatus sp. nov. ternal process; (Figs 6, 21–23) – O. oisensis sp. nov. in the metepisterna strongly widened posteriorly, more convex Material examined. Holotype ‘PA 8813’ apex of prosternal process, smoothed integ- (MNHN), probable female with moder- ument with coarser puncturation and more ately rounded hypopygidium; the complete raised pubescence; specimen visible laterally and ventrally – O. transversus sp. nov. in the shape of is included in a flattened irregular amber antennomeres 9 and 10, more convex apex piece with many pieces of dark organic of prosternal process, shape of metepisterna,

Figs 6–10. Oisenodes gen. nov. 6, O. clavatus sp. nov., holotype ‘PA 8813’, length 1.2 mm; 7, O. transversus sp. nov., holotype ‘PA 6520’, length 1.5 mm; 8, O. metepisternalis sp. nov., paratype ‘PA 382’; 9, O. transversus sp. nov., holotype ‘PA 6520’; 10, O. metepisternalis sp. nov., paratype ‘PA 12377’. Body, ventral view (6, 7), metacoxa and metepisternum, ventral view (8), antennal club, lateral (9) and ventral view (10). Scale bar: 0.1 mm (8–10).

smoothed integument with coarser punc- slightly conspicuous recumbent hairs less turation and more raised pubescence. than half as long as those on thoracic under- Notes. Taking into consideration a great side. Dorsum with distinct dense punctures, similarity of this new species to the type about half as large as eye facets in diame- species of the genus, the most characters ter, interspaces between them 3–5 times shared by both are omitted in the below de- as great as a puncture diameter and rather scription. smoothed. Metaventrite and ventrites with Description. Length 1.2, width 0.8, height clear very small punctures, interspaces be- about 0.5 mm. Elongate oval, strongly con- tween them very wide and smooth. vex dorsally and ventrally; subunicolourous Antennae slightly shorter than head dark brown to blackish; appendages brown- wide, eleven-segmented with three-seg- ish; dorsum with long and stout brownish mented club composing almost three eights greyish or greyish hairs (depending on light- of total antennal length, scape apparently ing) of two types: longer ones more than 3 rather small and elongate oval, ultimate an- times as long as ultimate antennomere and tennomere slightly shorter than two preced- about twice as long as femora wide, thicker ing antennomeres combined; antennomeres and erect; shorter ones somewhat thinner; covered with sparse, rather short and very thoracic underside with very sparse, long stout hairs. Pronotum somewhat narrow- and very slightly conspicuous recumbent er than combined elytral base, more than hairs, but much shorter than the shorter twice as wide as long; its anterior edge gen- ones on dorsum; abdominal ventrites with tly convex and posterior one somewhat less

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 54–69 A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER 63 than twice as great as anterior one, distinct- many layers making difficult to observe the ly bordered and bi-sinuate, sides about one specimen clearly (mostly ventrally), 9 mm third as great as posterior edge, paralateral of length, width of widest plane 7 mm and carinae distinct. Elytra about one and one 2 mm of narrowest plane; with two com- fifths as long as wide combined. Prosternal plete Nematocera of different species and process wide and widely rounded at apex. some pieces of organic matter. Ratio between pro-, meso- and metacoxae Paratype ‘PA 2262’ male with exposed about as following: 1.0:2.3:0.8. Distance penis trunk and parameres; the complete between metacoxae markedly greater than specimen is included in a small irregular antennal club wide. Metepisterna moder- clear amber parallelepiped. The inclusion is ately wide and rather widened at posterior put in a circular reservoir with Canada Bal- edge, nearly three times as long as wide at sam fixed on a microscope glass. posterior edge. Femora of usual shape and Etymology. The name of the new species moderately compressed, 2.0–2.5 times as refers to the structure of its metepisterna. wide as tibiae. Diagnosis. This new species is very simi- Variations. Paratype ‘164’ with body lar to O. gallicus sp. nov. and clearly differs length 1.5 mm. Some variability is observed from the latter only in the somewhat short- in pubescence. er pronotum and shape of metepisterna. The antennal club of the new species under Oisenodes metepisternalis sp. nov. consideration includes the ultimate anten- (Figs 8, 10–12, 16, 25–28) nomere somewhat shorter than that in O. gallicus sp. nov. Besides, O. metepisternalis Material examined. Holotype ‘PA 336 sp. nov. has the lateral sides of the prono- 2/7’ (MNHN); probable female with mod- tum nearly rectilinear (not arcuate) and erately rounded hypopygidium; the com- also more raised and more conspicuous pu- plete specimen with partly exposed both bescence of integument making the study of posterior wings is included in a subparal- the body surface very complicated. lel-sided irregular bar of clear consistence Notes. Taking into consideration a great forming a semicircle in cross section with similarity of this new species to the type one flat plane along the whole length with species of the genus, the most characters “milky cover” along pubescent integument shared by both are omitted in the below (although the dorsum partly clear); length description. of bar 12–15 mm and width of longitudinal Description. Length 1.3, width 0.8, plane 8 mm. height 0.6 mm. Elongate oval, strongly con- Paratype ‘PA 382’; the nearly complete vex dorsally and ventrally; subunicolourous specimen with partly exposed right poste- dark brown to blackish; appendages brown- rior wing and with missing right anterior ish; dorsum with long and stout brown- and intermediate legs is included in a small ish greyish or greyish hairs (depending on subparallel-sided irregular bar forming a lighting) of two types: longer ones nearly semicircle in cross section with one flat 1.5 times as long as ultimate antennomere plane along the whole length with many gas and about as long as femora wide, thicker vesicles of different sizes and “milky cover” and erect; shorter ones different from the along pubescent integument (particularly longer ones only in length; underside with at abdominal apex); length of bar 9.0 mm rather dense, long and strongly conspicuous and width of longitudinal plane 6 mm. recumbent hairs, but much shorter than the Paratypes ‘PA 12377’, male with exposed shorter ones on dorsum. Puncturation and anal sclerite; the complete specimen is in- sculpture of integument masked by pubes- cluded in a irregular and partly polished bar cence, ‘milky’ cover and many gas vesicles. forming a triangle in flat view; the bar has Antennae slightly shorter than head wide,

three times as long as wide. First abdominal ventrite about twice as long as 2nd ventrite, 3rd and 4th ventrites somewhat shorter than the latter; hypopygidium somewhat longer than 2nd ventrite. Variations. Length 1.2–1.4 mm. The paratype ‘PA 382’ has the slightly arcuate pronotal sides and longer elytra. The paratype ‘PA 2262’ shows the aedeagus with very narrow apex of penis trunk and also short and wide parameres. The surface of its pronotal base and elytra demonstrating clear punctures nearly as great as facets, interspaces between them 2–3 puncture di- ameters and rather smoothed.

Oisenodes oisensis sp. nov. (Figs 14, 24, 29–30)

Material examined. Holotype ‘PA 1387’ (MNHN), probable male; the nearly complete specimen with missing left posterior leg and with strong ‘milky’ cover along underside is included in a small sub-parallel- sided bar of clear consistence with one flat plane along the whole length and semicir- Figs 11–14. Oisenodes gen. nov. 11, 12, Oise- cular in cross-section; the beetle is visible nodes metepisternalis sp. nov., holotype ‘PA 336’, mostly ventrally because its dorsum is rath- length 1.3 mm (11), paratype ‘PA 382’, length 1.4 mm (12); 13, O. transversus sp. nov., holotype ‘PA 6520’, length 1.5 mm; 14, O. oisensis sp. nov., holotype ‘PA 1387’, length 1.6 mm. Body, dorsal (11–13) and ventral (14) view. eleven-segmented with three-segmented club, ultimate antennomere about one and one third as long as two preceding antennomeres combined. Pronotum slightly narrower than combined elytral base, about three times as wide as long; its anterior edge slightly convex and posterior one distinctly bordered and bi-sinuate, paralateral carinae distinct. Scutellum is clear and looking like transverse and subtriangular rather than subpentagonal, somewhat wider than long. Elytra about one and one third as Figs 15–16. Oisenodes gen. nov. 15, O. transver- long as wide combined. Ratio between pro, sus sp. nov., holotype ‘PA 6520’; 16, O. metepis- meso- and metacoxae about as following: ternalis sp. nov., holotype ‘PA 2262’. Ovipositor 1.0:2.3:0.8. Metepisterna comparatively (15) and aedeagus (16), ventral view. Scale bar: wide and widened posteriorly and about 0.1 mm.

, holotype ‘PA 2501’, , holotype ‘PA sp. nov. Oisenodes azari , 20 , 19 , paratype ‘PA 164’, length 1.5 mm (21), holotype ‘PA 8813’, length 1.2 mm (22), 164’, length 1.5 mm (21), holotype ‘PA , paratype ‘PA , holotype ‘PA 2307’, length 1.3 mm; , holotype ‘PA sp. nov. , holotype ‘PA 1387’, length 1.6 mm. Body, dorsolateral (21, 23), ventral (22) and ventrolateral 1387’, length 1.6 mm. Body, , holotype ‘PA sp. nov. clavatus sp. nov.

Oisenodes Oisenodes gallicus O. oisensis , , , 18 24 , 21–23 17

gen. nov. gen. nov.

Oisenodes Oisenodes

Figs 17–20. (24) view. type ‘PA 1842’, length 1.3 mm (23); type ‘PA length 1.3 mm. Body, ventral (17, 19) and dorsal (18, 20) view. length 1.3 mm. Body, Figs 21–24.

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 54–69 66 A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER holotype ‘PA 336’, length 1.3 mm (25, 26); paratype ‘PA 382’, length 1.4 mm (27, 28). Body, dorsal (25), 382’, length 1.4 mm (27, 28). Body, 336’, length 1.3 mm (25, 26); paratype ‘PA holotype ‘PA

, holotype ‘PA 6520’, length 1.5 mm. Body, dorsal (29) and ventral (30) view. 6520’, length 1.5 mm. Body, holotype ‘PA

, sp. nov. sp. nov. metepisternalis transversus

Oisenodes Oisenodes

Figs 25–28. ventral (26), dorsolateral (27) and ventrolateral (28) view. Figs 29–30.

© 2010 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 19(1): 54–69 A.G. KIREJTSHUK ET AL. NEW TRINODINE GENUS AND SPECIES FROM LOWERMOST EOCENE AMBER 67 er closely approached to the unpolished sur- though apparently rather coarse and sparse face of bar; the amber bar with length 8 mm puncturation visible on elytra and also and width of longitudinal plane 7 mm. comparatively dense and coarse punctura- Etymology. The name of the new spe- tion on the middle of metaventrite. Anten- cies refers to the name of locality, where the nae rather shorter than head wide, eleven- specimens were collected. segmented with three-segmented club com- Diagnosis. This new species is about as posing about three eights of total antennal pubescent as O. gallicus sp. nov. and O. length, ultimate antennomere only slightly metepisternalis sp. nov., but clearly differs longer than two preceding antennomeres from both mostly in the more slender and combined. Pronotum much narrower than much more convex body, and also in the combined elytral base, more than twice as shorter ultimate antennomere. At the same wide as long; its anterior edge slightly con- time O. oisensis sp. nov. has short sides of vex, posterior one less than twice as long pronotum somewhat like those in O. azari as anterior one, sides about one quarter as sp. nov., although it differs from the latter great as posterior edge and nearly rectilin- in the much shorter ultimate antennomere, ear. Elytra about one and one third as long wider metepisterna, not smoothed integu- as wide combined. Metepisterna compara- ment with coarser puncturation and more tively wide and subparallel-sided, about 3 raised pubescence. Finally, the new spe- times as long as wide. First abdominal ven- cies under consideration is distinct from trite about as long as 2nd and 3rd ventrites O. transversus sp. nov. in the somewhat combined and each of the latter and 4th more slender and convex body as well as ventrite comparable in length; hypopygid- in the shape of pronotum as well as from ium somewhat longer than previous ven- O. clavatus sp. nov. in the metepisterna trite, moderately rounded at apex. Femora subparallel-sided, only slightly convex of usual shape and moderately compressed, apex of prosternal process, not smoothed 2.5–3.0 times as wide as tibiae. integument with finer puncturation and less raised pubescence. Oisenodes transversus sp. nov. Notes. Taking into consideration a great (Figs 7, 9, 13, 15) similarity of this new species to the type species of the genus, the most characters Material examined. Holotype ‘PA 6520’ shared by both are omitted in the below de- (MNHN), female with exposed ovipositor; scription. the specimen with missing right protibia Description. Length 1.6, width 0.6, height and protarsus, right mesotibia and mesotar- about 0.7 mm. Elongate oval, strongly con- sus, left posterior leg, and also with broken vex dorsally and ventrally; subunicolour- and partly removed left part of thorax and ous brownish; appendages rather yellow- abdomen is included in a very small amber ish; dorsum with long and stout yellowish piece put between two glass covers for mi- hairs of two types: longer ones markedly croscopy. more than twice as long as femora wide, Etymology. The name of the new species thicker and erect; shorter ones intermixed refers to its transverse pronotum. on each dorsal sclerite; thoracic under- Diagnosis. The new species is distinct side with moderately dense, long and very among the congeners due to more narrowly slightly conspicuous recumbent hairs, but separated procoxae. Nevertheless, it is rather much shorter than the shorter ones on dor- similar to O. oisensis sp. nov., differing from sum; abdominal ventrites with rather con- the latter only in the somewhat more robust spicuous and fine recumbent hairs less than and apparently not so strongly convex body half as long as those on thoracic underside. as well as in the shape of pronotum. Besides, Integument masked with ‘milky’ cover, al- it differs also from:

– O. azari sp. nov. in the much shorter posterior edge with distinct border and ultimate antennomere, shape of pronotum, about three times as great as lateral ones. wider metepisterna, not smoothed integu- Elytra about one and one fifths as long as ment with coarser puncturation and more wide combined. Metepisterna moderately raised pubescence. wide and subparallel-sided, about 3.5 times – O. clavatus sp. nov. in the shape of an- as long as wide. First abdominal ventrite tennomeres nine and ten, less convex apex about as long as 2nd and 3rd ventrites com- of prosternal process, shape of metepister- bined, the each of latter and 4th ventrite na, not smoothed integument with coarser comparable in length; hypopygidium some- puncturation and more raised pubescence; what longer than previous ventrite, widely – O. gallicus sp. nov. and O. metepister- rounded at apex. Femora of usual shape and nalis sp. nov. in the longer antennae with moderately compressed, 2.5–3.0 times as shorter ultimate antennomere, coarser mi- wide as tibiae. crosculpure of integument with indistinct puncturation, less raised pubescence, and ACKNOWLEDGEMENTS from the second also in the narrower metepisterna. The authors thank the company Lafarge- Granulat for the help with the sampling of the Notes. Taking into consideration a great fossils and the family Langlois-Meurinne for the similarity of this new species to the type authorization of working on their property. The species of the genus, the most characters authors also appreciate to G. De Ploëg (MNHN) shared by both are omitted in the below de- and D. Azar (Lebanon University) for the care- scription. ful preparation of the material. Gilbert Hodebert Description. Length 1.5, width 0.7 mm. (MNHN) assisted to the authors in preparation of Elongate oval, strongly convex dorsally and illustrations for this and other papers. A.G. Kire- ventrally; subunicolourous dark brown; ap- jtshuk has a pleasant duty to express his recogni- pendages reddish; dorsum with long and tion to the administration of MNHN for provid- ing him a possibility to work in this museum in stout yellowish hairs of two types: longer 2006–2010 as visiting professor. This study was ones about twice as long as femora wide, also supported by the grants of Russian Founda- thicker and erect; shorter ones intermixed tion of Basic Research No. 09-04-00789-а and on each dorsal sclerite; underside with also Programme of the Presidium of the Russian dense, long and very slightly conspicuous Academy of Sciences “Origin and Evolution of recumbent hairs, but much shorter than the Biosphere”. The authors greatly appreciate also shorter ones on dorsum. Integument show- both anonymous reviewers for their important ing rather comparatively coarse microsculp- contribution improving the manuscript. ture to alutation on each sclerites and indistinct puncturation, although on abdominal REFERENCES ventrites traces of coarse and shallow indis- Batelka, J., Collomb, F.-M. & Nel, A. 2006. tinct punctures with interspaces somewhat Macrosiagon deuvei n. sp. (Coleoptera: Ripi- greater than puncture diameter visible. An- phoridae) from the French Eocene amber. tennae somewhat longer than head wide, Annales de la Société Entomologique de Fran- eleven-segmented with three-segmented ce. (N.S.), 42(1): 75–78. club composing about three eights of to- Bílý, S. & Kirejtschuk, A.G. 2007. Philanthaxo- tal antennal length, ultimate antennomere ides gallicus gen. nov., sp. nov. from the only slightly longer than two preceding an- Lowermost Eocene French amber (Co- leoptera: Buprestidae). Folia Heyrovskyana, tennomeres combined. Pronotum slightly Ser. A., 14(4): 181–186. narrower than combined elytral base, much Háva, J. 2003. World Catalogue of the Dermes- more than twice as wide as long; its anterior tidae (Coleoptera). Studie a Zprávy Oblast- edge slightly convex, sides nearly somewhat ního Muzea Praha-východ v Brandýse nad arcuately narrowing anteriorly, bi-sinuate Labem a Staré Boleslavi, Suppl., 1: 1–196.

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