ZOOSYSTEMATICA ROSSICA, 20(1): 62–70 28 JULY 2011

First Mesozoic representative of the subfamily Liparochrinae (Coleoptera: Hybosoridae) from the Lower Lebanese amber Первый мезозойский представитель подсемейства Liparochrinae (Coleoptera: Hybosoridae) из нижнемелового ливанского янтаря

A.G. KIREJTSHUK, D. AZAR & O. MONTREUIL

А.Г. КИРЕЙЧУК, Д. АЗАР, О. МОНТРЕЙ

A.G. Kirejtshuk, Zoological Institute of Russian Academy of Sciences, 1 Universitetskaja Emb., St. Petersburg, 199034, Russia; CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 57 Rue Cuvier, F-75231, Paris, France. E-mail: [email protected], [email protected] D. Azar, Lebanese University, Faculty of Sciences II, Natural Sciences Department, Fanar-Matn, P.O. box 26110217, Lebanon; CNRS 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France. E-mail: [email protected] O. Montreuil, CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France. E-mail: [email protected]

One specimen among coleopterous inclusions recently recovered in Lebanese amber is de- scribed as Libanochrus calvus gen. et sp. nov. and assigned to the subfamily Liparochrinae of the Hybosoridae. This specimen is incomplete but a large part of its head with appenda- ges, prothoracic segment with anterior legs, remains of the median part of the pterothoracic underside and the lateral base of the of the right elytron make possible the conclusion on the subfamily attribution and diagnose it among the rest of fossil and recent taxa of this family. At present it is the oldest representative of the subfamily. Один экземпляр среди инклюзов жуков, недавно найденный в ливанском янтаре, описан как Libanochrus calvus gen. et sp. nov. и отнесен к подсемейству Liparochrinae из семей- ства Hybosoridae. Этот экземпляр неполный, но большая часть его головы с придатками, переднегрудной сегмент с передними ногами, остатки медиальной части низа птерото- ракса и основание бока правого надкрылья позволяют cделать вывод о его подсемей- ственной принадлежности и определить его среди представителей семейства. В настоя- щее время он является древнейшим представителем подсемейства. Key words: Lower Cretaceous, fossil, Lebanese amber, Coleoptera, Hybosoridae, Leparochri- nae, new genus, new species Ключевые слова: нижний мел, ископаемые, ливанский янтарь, Coleoptera, Hybosoridae, Leparochrinae, новый род, новый вид

INTRODUCTION by members of the subfamily Hy- bosorinae (Bajan-Teg, Uver-Khangay aj- The family Hybosoridae Erichson, 1847 mag, Mongolia ‒ Jurahybosorus mongolicus is a worldwide distributed family within Nikolajev, 2005 and Karatau, Kazakhstan ‒ , being most diverse in the areas with tropical climate and currently Protohybosorus karatavicus Nikolajev, 2010; comprising more than 220 species belong- P. grandissimus Nikolajev, 2010 and P. me- ing to 35 genera within five subfamilies sasiaticus Nikolajev, 2010). This subfamily (Ocampo, 2006). It is represented in the is known also from the Lower Cretaceous fossil record since the Middle or Upper of Baissa (Buryatskaya Autonomous Repu-

© 2011 Zoological Institute, Russian Academy of Scienсes 64 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER blic, Transbaikalia, Russia; Middle Neoco- MATERIAL AND METHODS mian: Cretohybosorus buryaticus Nikolajev, 1999 and C. striatulus Nikolajev, 1999; and Till present day the oldest amber with also Leptosorus zherikhini Nikolajev, 2006), many inclusions is from Lebanon from the Lower Cretaceous of Chifeng (Azar, 1997). The Lebanese amber ranges in (Inner Mongolia, China: Leptophorus for- age from the Late Jurassic to Cenomanian. tus (Ren, Zhu & Lu, 1995) (Geotrupoides)), The fossiliferous outcrops are all approxi- from the Oligocene of Kučlin [Kutschlin] mately of the same age and are mainly late (Czech Republic: Phaeochrous tertiarum to lowermost (Azar et (Deichmüller, 1881) (Bolboceras)), from al., 2003a). The precise dating of the Leba- the Lower Dominican amber nese material was possible after the disco- (Dominican Republic: Procoilodes adrastus very of stratigraphical marker fossil pollen Ocampo, 2002 and Tyrannasorus rex Rat- trapped in the amber (Dejax pers. com.). In cliffe & Ocampo, 2001) and from the Upper the majority of amber outcrops in Lebanon, Miocene of Oeningen (Baden-Württem- the amber is found in its primary deposits. berg, Germany: lividus Heer, The material studied herein comes from 1862). Besides, from Baissa Cretanaides the Lower Cretaceous outcrops of Bouarij trogopterus Nikolajev, 1996 and Protanaides (Caza Zahleh, Beqaa, Central Lebanon) sibiricus Nikolajev, 2010 from the subfa- (for details see Azar et al., 2010). The ma- mily Anaidinae Nikolajev, 1996 and Mima- terial has been prepared (cut and polished), phodius pusillus Nikolajev, 2007 from the then included in Canada balsam medium subfamily Mimaphodiinae Nikolajev, 2007 between two glass cover slips as described are known. Recently Mesoceratocanthus by Azar et al. (2003b). Observations were tuberculifrons Nikolajev, Wang, Liu & Zi- made with usual optic equipment; in partic- iang, 2010 from the of the ular, the stereomicroscope Olympus SCX9 Upper Jurassic – Lower Cretaceus was de- and inverted microscope Olympus CK 40 é scribed in the subfamily . in Mus um National d’Histoire Naturelle, Finally, Coprologus gracilis Heer, 1847 with Paris, and also the stereomicroscope micro- unclear subfamilian attribution was de- scope Leica MZ 16.0 in Zoological Insti- scribed from Oeningen. In spite of the fact tute, St. Petersburg, were used. that some mentioned indications could be The material under consideration is regarded as somewhat doubtful, they give a temporally deposited in the Laboratory of é preliminary imagination on the distribution entomology, Mus um National d’Histoire of this family through time. Naturelle, Paris, until a national natural his- This paper constitutes the sixth contri- tory museum in Lebanon is established. bution to the knowledge on fauna of Co- leoptera from Lebanese amber (Kuschel & RESULTS Poinar, 1993; Lefebvre et al., 2005; Kirejt- shuk & Azar, 2008; Kirejtshuk et al., 2009a, Order COLEOPTERA 2009b), which is devoted to the families, Suborder oldest representation of which in fossils is Superfamily SCARABAEOIDEA frequently put namely in Lebanese inclu- sions. The specimen described here is also Family HYBOSORIDAE Erichson, 1847 the oldest representative of the hybosorid Subfamily LIPAROCHRINAE subfamily Liparochrinae Ocampo, 2006. A Ocampo, 2006 more detailed overlook on representation of Libanochrus gen. nov. the family Hybosoridae in the fossil record can be taken from the catalogue by Pon- Type species Libanochrus calvus gen. et omarenko & Kirejtshuk (2011). sp. nov.

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 65

Diagnosis. Body of comparatively small notum projecting posteriorly and armature size. Head and pronotum comparatively of protibia (intermixed by more prominent subflattened and without any horn or ca- and small teeth). rina. Head feebly transverse, its outline In contrast to most other members of evenly curved, including the short but wide the Hybosoridae, this genus is mostly char- eye canthus and clypeus; clypeo-frontal su- acterised by the comparatively short head ture not visible; labrum transverse, short and mandibles not exposed dorsally. The and rather wide, flat, prominent, exposed members of the subfamily Anadiinae dif- from above beyond apex of clypeus, located fer from the new species in their markedly in an evenly curved excision of the latter, longer head with frons not dilated later- its anterior edge forming a gentle curved ally, projecting and not very wide procoxae, line, its side forming a discontinuity with coarse sculpture of integument etc.; and the curve of clypeal side; eyes surrounded Cretanaides Nikolajev, 1996 from Anaidinae by fronto-clypeal folds; mandibles not ex- cannot be compared with the new species, posed dorsally. Antenna short and with first because it was described after one elytron. club segment suboval cupular-shaped. Pro- The shape of mesocoxae and very smooth notum widest at the distinct posterior an- integument of this specimen are similar to gles, with deeply excised anterior edge and those of some Ceratocanthinae rather than widely explanate sides; lateral margins cili- to those of and Liparochri- ate. All pairs of coxae narrowly separated nae, although other characters, including or conjoined; mesocoxae feebly transverse the outline of the anterior part of its head and not very wide. Integument of dorsum and an exposed labrum are very similar to smooth and unpunctured. Protibia triden- those in some extant representatives of the tate on outer edge, with distinct crenula- last two subfamilies. Libanochrus gen. nov. tions from the base and between the main is particularly distinguished from the mem- teeth, and bearing long setae between each bers of Ceratocanthinae by the exposed pair of teeth in the crenulation. Femora and labrum from under clypeus, much wider tibiae with brushes of long hair at apices. procoxae, shape of anterior legs and arma- Comparison. The genus Libanochrus ture of protibiae. In particular, the Meso- gen. nov. showing explanate sides of pro- zoic Mesoceratocanthus Nikolajev, Wang, notum, first antennal club segment cupu- Liu & Ziiang, 2010 from this subfamily, in lar-shaped, position of coxa, clypeo-frontal contrast to the new genus, has elongate and suture not visible, shape of protibia (with larger body (12 mm) with subquadrangu- setae between crenulations) coincides with lar prothorax, oval labrum, exposed and far the extant members of the family Hybosori- projecting mandibles, and also narrow pro- dae. A particular evidence of the subfamily tibiae without prominent teeth. attribution of the new genus is the position In fossils, the new genus differs from of transverse labrum, exposed from above, Mimaphodius Nikolajev, 2007 (Mimapho- at the same level as the dorsal surface of diinae) in the not very convex body, ex- head, fitting an excavation of the clyp- posed labrum, lack of clypeo-frontal suture, eus (as observed in Liparochrus Erichson, outline of head and pronotum, rather even 1848). Libanochrus gen. nov. differs from dorsal surface of head and widely explanate other genera in the anterior outline of its pronotal sides. It is distinct also from the rather short head (evenly curved, not sinu- Mesozoic Cretohybosorus Nikolajev, 1999, ate at front base of the eye canthus), mark- Leptosorus Nikolajev, 2006 and Protohypo- edly narrower mesocoxae and type of the ar- bosorus Nikolajev, 2010 in the not exposed mature of protibia, and also from Antiochrus mandibles, laterally extended head sides Sharp, 1873, in the not so convex body, lack making the head transverse, widely ex- of hairs on dorsum, posterior angles of pro- planate pronotal sides, posterior angles of

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70 66 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER pronotum slightly projecting backwards bles with the outer anterior edge somewhat (posterior edge of pronotum subsinuate shortly elevated and their elevated edge is at sides) [although the pronotum of Cre- closely clasped to the anterior edge of the tohybosorus could be somewhat similar]; labrum. However most species of the lat- and also Protanaides sibiricus in the ante- ter genus have the mandibles far projecting rior part of head strongly extended later- anteriorly and pronotal sides only subex- ally and surrounding the eyes, narrower planate or narrowly explanate. Besides, the head, pronotum more arcuately narrowing new species has a short labrum similar to anteriorly and rounded anterior angles and members of Seleucosorus Kuijten, 1983 (the also different armature of protibia. Since later have the mandibles also short, but dis- Jurahybosorus Nikolajev, 2005 (Hybosori- tinctly projecting anteriorly). Finally, the nae) was described after the metathoracic specimen examined is comparatively small, ventral sclerites and posterior legs only, it however many groups of the generic rank cannot be compared with the new species. include the members with small body size. Libanochrus gen. nov. also differs from the Libanochrus calvus gen. et sp. nov. shows Caenozoic Procoilodes Ocampo 2002 from the most similarity to some species of Phae- Hybosorinae (probably with somewhat ochrous and Kuijtenous Paulian, 1981, but in similar pronotum and small body size) in addition to the mandibles which are not ex- the flatly depressed to excavate (and not posed from under labrum, a comparatively tuberculate) frons, not pointed labrum and short head and gently outlined head sides smoothed surface of head and pronotum; (not subsinuate at the base of eye canthus), and from Tyrannasorus Ratcliffe & Ocampo the new species has no trace of border along 2001 in the completely different structure edges of its pronotum. of the transverse head (concealed man- Etymology. The name of the new genus dibles, not projecting labrum, laterally ex- is formed from Greek ‘Libanos’ (Lebanon) tended anterior part of the head). Finally, and a usual root of generic names in Hy- the new genus is very distinct from Copro- bosoridae descended from the Greek ‘chros’ logus gracilis Heer, 1847 in the strongly di- (skin, surface of body; skin colour). Gender lated sides of the head, pronotum more nar- masculine. rowing anteriorly and concealed mandibles. All genera of Hybosorinae with later- Libanochrus calvus sp. nov. ally expanded clypeus have more or less ex- (Figs 1–7) pressed sinuation at the base eye canthus, while Libanochrus calvus gen. et sp. nov. is Material examined. Holotype. ‘TAR 39’, sex characterised by the even outline of the lat- unknown; flat piece of amber, the inclusion bad- ly preserved but with characters that allow the eral edges of its head. Besides, the head of description and placement of the fossil, such as all recent Hybosorinae is markedly longer the head and most of its appendages, most of the than that in new species and the dorsum of prothoracic segment with anterior legs and most the recent Hybosorinae is clearly punctured of the ventral median surface of pterothorax, (not infrequently rather coarsely), but the left meso- and metafemora, a trace of the dorsal head and pronotum of the new species are surface of right meso- and metafemora is located smooth and nearly unpunctured. Hypsolo- in the epoxy between round microscope glasses. deres Fairmaire, 1893 has an appearance Pieces of yellowish to very dark organic matter dorsally somewhat similar to Libanochrus and very small gas vesicles surround the inclu- sion. The remains of the body seemed to be some- gen. nov., because its rather long labrum what pressed before entombing in the fresh resin. and mandibles are turned ventrally. Some Most parts of integument are masked by very species of Phaeochrous Laporte, 1840 small pieces of organic matter and gas vesicles. demonstrate comparatively short exposed The specimen examined seemed to be rather soft labrum (as in the new species) and mandi- when it became stuck to the resin but still alive

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 67

Fig. 1. Libanochrus calvus gen. et sp. nov., ho- lotype. Head and pronotum, dorsal view. Head width 2.6 mm. Fig. 2. Libanochrus calvus gen. et sp. nov., holo- type. Head and thorax, ventral view. Head width (being evidence of the traces of moving in the 2.6 mm. amber), then it was destroyed and decayed before fossilization (as a result its remains including ap- pendages became additionally depressed). Head transverse, somewhat depressed Description of the holotype. Length of medially and widely explanate along frons, head 1.4, width of head 2.6, width of avail- much narrower than pronotum; median part able part of pronotum 3.4 mm (complete of frons apparently partly isolated by fine body length of this specimen before its de- and oblique swellings at each side. Eyes not struction could be about 5.0 mm). Head clearly outlined (because the layer of small and pronotum without hairs; pronotal and pieces of organic matter and very small gas elytral sides, and legs with moderately long bubbles covers them) apparently finely fac- and fine hairs (particularly dense on entire eted eyes (although facets scarcely visible surface of femora and they are arranged in and eyes seeming translucent) and with lat- a long row along the anterior edge and two eral edge surrounded by fold of frons. Ulti- shorter ones located along posterior edge of mate maxillary palpomere narrowest and as meso- and metafemora; with brushes of long long as two previous ones combined. Apices setae at apices of all femora and very dis- of maxillary lobes (narrow galea and rather tinct brush of very long setae along anterior wide lacinia) exposed from under labrum edge of profemur; besides longitudinal rows and bearing extremely dense setae. First of setae are visible on protibiae). Dorsum of antennal club segment somewhat dorsoven- head and pronotum as well as prohypomera trally compressed. Pronotum widest at base apparently without puncturation. Along and compressed at right posterior angle, lateral edges of frons there are dense trans- however it was apparently not very con- verse strips looking like strengthening and vex, its anterior edge strongly trapezium- lightening of sclerotization. likely excised, its sides narrowly explanate

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70 68 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER

Antennal club of the specimen examined represented by the basal wall of the first seg- ment, but remains seem to be moved away. Etymology. The epithet of the new spe- cies refers to the mostly smooth and hairless integument of dorsum of the specimen.

DISCUSSION

The composition and placement of the family Hybosoridae are in great discussion during the last decade. The most gene- ralised viewpoint, which is close to the traditional one, was formulated by Browne and Scholtz (1999) and Howden and Gill Fig. 3. Libanochrus calvus gen. et sp. nov., ho- lotype. Head with exposed maxillary lobes (nar- (2000) according to which Hybosoridae, row galea and rather wide lacinia) and palpi, , Ochodaeidae Mulsant & dorsal view. Head width 2.6 mm. Rey, 1871 and Geotrupidae Latreille, 1802 can be treated as most relatives putting the latter as closer to their common ances- and with undulate edges, posterior angles tor. Krell (2006) after a review of the fossil with distinct top, posterior edge apparently records also regards the Hybosoridae and widely and evenly convex and with a slight Ceratocanthidae as sister groups with the sinuation at posterior angles. All coxae ap- closest link also to Ochodaeidae and joining parently (sub)contiguous; procoxae appar- all three families with Geotrupidae. Ocam- ently suboval to subquadrangular, meso- po (2006) and Ocampo and Hawks (2006) coxae suboval and metacoxae transverse. on molecular and morphological compari- Protibia comparatively short (slightly son put Hybosoridae (including Ceratocan- longer than head) and rather compressed, thidae) as a sister group of Ochodaeidae, its outer edge with clear not quite regu- both of which are linked with Glaphyridae lar and oblique crenulations from the base MacLeay, 1819. Nikolajev (2007) having and between three more prominent teeth analysed some fossil families considers as disposed with subequal distances between sister groups the Hybosoridae (including them, the proximal tooth rather small. Spur Belohinidae Paulian, 1959) and Trogidae of anterior legs very long (about as long MacLeay, 1819 and approaching both to as protarsomeres 1 and 2 combined) and Geotrupidae and some subfamilies of Scara- slightly curved. Profemora gently curved baeidae Latreille, 1802. The new species along posterior edge and strongly subangu- demonstrates that the subfamily Liparo- larly curved along the anterior one, about chrinae has a Mesozoic root in addition to twice as wide as probibia. Left meso- and the Middle Jurassic – Cretaceous Anaidi- metafemora are visible only from posterior nae Hybosorinae and Ceratocanthidae side and they look like strongly compressed. (Krell, 2007; Ponomarenko & Kirejtshuk, Protarsus rather thin and apparently about 2011; Nikolajev et al., 2010). On the other three quarters as long as protibia; protarso- hand, the comparatively great representa- mere 1 apparently longer than each of pro- tion of this family in fossils from the Asian tarsomeres 2–4 and protarsomere 5 longest outcrops could be evidence that this family (about as long as protarsomeres 1–4 com- was rather diverse on this continent during bined; one claw available apparently about the late Mesozoic, compared with diversity two fifths as long as protarsomere 5. of Geotrupidae, Ochodaeidae and Glaphy-

© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 69 ridae beginning from the Cretaceous (Krell, Browne, J. & Scholtz, C.H. 1999. A phylogeny 2007; Kirejtshuk et al., 2010; Ponomarenko of the families of Scarabaeoidea. Systematic & Kirejtshuk, 2011). Entomology, 24(1): 51–84. Howden, H.F. & Gill, B.D. 2000. Tribes of New World Ceratocanthinae, with keys to genera ACKNOWLEDGEMENTS and descriptions of new species (Coleoptera: Scarabaeoidea). Sociobilogy, 35: 281–329. The authors greatly appreciate the gener- Kirejtshuk, A.G. & Azar, D. 2008. New taxa ous assistance of A. Nel during the study. The of (Insecta, Coleoptera) from Leba- two first co-authors have a pleasant duty to ex- nese amber with evolutionary and systematic press their recognition to the Muséum National comments. Alavesia, 2: 15–46. d’Histoire Naturelle for providing them a pos- Kirejtshuk, A.G., Azar, D. Beaver, R.A., Man- sibility to work in this museum as visiting pro- delstam, M.Yu. & Nel, A. 2009a. The most fessors. The study of the first co-author was also ancient bark known: a new tribe, ge- supported by the Programme of the Presidium nus and species from Lebanese amber (Cole- of the Russian Academy of Sciences ‘Origin of optera, Curculionidae, Scolytinae). System- Biosphere and Evolution of Geo-Biological Sys- atic Entomology, 34(1): 101–112. tems’ and by a grant of the Russian Foundation Kirejtshuk A.G., Azar D., Tafforeau P., Bois- of Basic Research (09-04-00789). This paper is tel R. & Fernandez V. 2009b. New beetles a contribution to a team project ‘Biodiversity: of Polyphaga (Coleoptera, Polyphaga) from Origin, Structure, Evolution, and Geology’ al- Lebanese amber. Denisia, 26 (Zugleich Kata- lotted to the second co-author and his team by loge der OberösterreichischenLandesmu- the Lebanese University. An essential help was seen, Neue Serie. 86): 119–130. obtained from G.V. Nikolajev (Al-Farabi Kazakh Kirejtshuk, A.G., Ponomarenko, A.G., Pro- University, Almaty) and Wang Bo (Nanjing In- kin, A.A., Chang, H., Nikolajev, G.V., stitute of Geology and Palaeontology of Chinese Ren, D. 2010. Current knowledge on Me- Academy of Sciences) provided with their re- sozoic Coleoptera from Daohugou and Lia- cent publications. The authors greatly appreci- oning (North East China). Acta Geologica ate valuable comments on an earlier version of Sinica, 84 (4): 783–792. the manuscript received from F.T. Krell (Denver Krell, F.-T. 2006. Fossil Record and Evolution of Museum of Nature and Science) and S. Hisamat- Scarabaeoidea (Coleoptera: Polyphaga). Co- su (Ehime University, Matsuyama). leopterists Society Monograph, 5 (Supplement to The Coleopterists Bulletin, 60): 120–143. REFERENCES Krell, F.-T. 2007. Catalogue of fossil Scarabaeoi- dea (Coleoptera: Polyphaga) of the Mesozoic Azar, D. 1997. A new method for extracting and Tertiary. Version 2007. Technical Report. vege tal and fossils from the Lebanese Denver Museum of Nature & Science, Den- amber. Palaeontology, 40(4): 1027–1029. ver, 2007-8: 1–79. Azar, D., Nel A. & Gèze, R. 2003a. Use of am- Lefebvre, F., Vincent, B., Azar, D. & Nel, A. ber fossil inclusions in palaeoenvironmental 2005. The oldest beetle of the Euaestheti- reconstruction, dating and palaeobiogeogra- nae (Staphylinidae) from Early Cretaceous phy. Acta Zoologica Cracoviensa, 46 (suppl. Lebanese amber. Cretaceous Research, 26(2): Fossil ): 393–398. 207–211. Azar, D., Perrichot, V., Néraudeau, D., & Nel, Nikolajev, G.V. 2007. Mezozoyskiy etap evolyu- A. 2003b. New psychodid flies from the Cre- tsii plastinchatousykh (Insecta: Coleoptera: taceous ambers of Lebanon and France, with Scarabaeoidea) [Mesozoic stage of evolution a discussion about Eophlebotomus connectens of scarabaeoid beetles (Insecta: Coleoptera: Cockerell, 1920 (Diptera, Psychodidae). An- Scarabaeoidea)]: 1–221. Almaty, Kazakh nals of the Entomological Society of America, universiteti. 96(2): 117–127. Nikolajev, G.V., Wang Bo, Liu Yu & Zhang Azar, D., Gèze, R. & Acra, F. 2010. Lebanese Hai-chun 2010. First record of Mesozoic amber. In: Penney, D. (Ed.) Biodiversity of Ceratocanthinae (Coleoptera: Hybosori- fossils in amber from the major world deposits: dae). Acta Palaeontologica Sinica, 49(4): 271–298. Siri Scientific press, Manchester. 443–447.

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© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70