The Oldest Representatives of the Family Coccinellidae(Coleoptera
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ZOOSYSTEMATICA ROSSICA, 21(1): 131–144 25 JULY 2012 The oldest representatives of the family Coccinellidae(Coleoptera: Polyphaga) from the Lowermost Eocene Oise amber (France) Древнейшие представители семейства Coccinellidae (Polyphaga: Coleoptera) из нижнеэоценового янтаря Уаз (Франция) A.G. KIREJTSHUK* & A. NEL А.Г. КИРЕЙЧУК*, А. НЕЛЬ A.G. Kirejtshuk, Zoological Institute of the Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia; CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France. E-mails: [email protected], [email protected]. *Corresponding author. A.Nel, CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France. E-mail: [email protected] In the paper two new species of the genus Rhyzobius Stephens, 1829 (R. antiquus sp. nov. and R. gratiosus sp. nov.) and one new species of the genus Nephus Mulsant 1846 (N. subcircularis sp. nov. without a certain subgeneric placement) from the Lowermost Eocene amber of Oise are described. A short review of known fossil records of the family Coccinellidae is given. В статье описаны два новых вида из рода Rhyzobius Stephens, 1829 (R. antiquus sp. nov. и R. gratiosus sp. nov.), а также один новый вид рода Nephus Mulsant 1846 (N. subcircularis sp. nov. без определенной подродовой принадлежности) из нижнемелового янтаря Уаз. Приводится краткий обзор сведений по ископаемым семейства Coccinellidae. Key words: Lowermost Eocene, amber, Oise, France, Coleoptera, Coccinellidae, Coccidulini, Scymnini, new species Ключевые слова: нижний эоцен, янтарь, Уаз, Франция, Coleoptera, Coccinellidae, Coc- cidulini, Scymnini, новые виды INTRODUCTION and Kubisz (2000) mentioned this family without any more precise attribution from The paper is the tenth contribution to the Eocene Baltic amber, and Grimaldi & the knowledge of the fauna of Coleoptera Engel (2005) published pictures of an adult from the Lowermost Eocene French amber and a larva from the Miocene Dominican collected in Oise falls (Batelka et al., 2006; amber, while Gersdorf (1969) reported a Bílý & Kirejtshuk, 2007; Kirejtshuk & Nel, compression print of this family among 2008, 2009; Moseyko et al., 2010; Kirejt- shuk et al., 2010a,b; Kovalev et al., 2012). sediments from the Upper Pliocene of Wil- The family Coccinellidae Latreille, 1807 lershausen (Niedersachsen, Germany). is known in fossils only from some Cainozoic Besides, among the inclusions from Bal- resources, and no amber specimen has been tic amber Berendt (1845), Menge (1856), described till now. All indications of Cocci- Klebs (1910), Larsson (1978) and Hieke nellidae in the Jurassic cannot be regarded & Pietrzeniuk (1984) indicated the genera as subject of consideration, because for now Coelopterus Mulsant, 1853 and Pharus Mul- the reliable cucujoids appear in the fossil re- sant, 1850 (Sticholotidinae Weise, 1901), cord not earlier than at the beginning of the Scymnus Kugelann, 1794 (Coccidulinae Cretaceous. Hieke & Pietrzeniuk (1984) Mulsant, 1846), Platynaspis L. Redten- © 2012 Zoological Institute, Russian Academy of Scienсes 132 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE bacher, 1843 (Chilocorinae Mulsant, 1846), MATERIAL AND METHODS and Coccinella Linnaeus, 1758 (Coccinel- linae Latreille, 1807). The subfamily Coc- Many specimens recovered among in- cidulinae was also recorded in the Lower/ clusions from the Lowermost Eocene Middle Miocene of Shanwang Basin [Shan- French amber are deposited in the Labora- dong, China: Scymnus cf. kawamurai Ohta, toire de Paléontologie, Muséum National 1929 (by Zhang, 1989) and Hippodamia ol- d’Histoire Naturelle, Paris (MNHN). For bia J. Zhang, Sun et X. Zhang, 1994] and in this study ordinary optic equipment was the Oligocene of Brunnstatt (Haut-Rhine, used, in particular the stereomicroscope Elsas, France: Scymnus angulatus Förster, Olympus SCX9 and inverted microscope 1891). The subfamily Chilocorinae was also Olympus CK 40 in the Paris museum, and recorded in the Earliest Oligocene of Flo- also the stereomicroscope Leica MZ 16.0 rissant (Colorado, USA: Chilocorus ulkei in the Zoological Institute, St Petersburg. Scudder, 1900) and also from Brunnstatt All the holotypes and most paratypes of the [Chilocorus foersteri Ukrainsky, 2010 (=pol- new species are deposited in the Paris muse- itus Förster, 1891, non Mulsant, 1850) and um, one paratype of Rhizobius antiquus sp. C. inflatus Förster, 1891]. nov. (“PA 245”) is deposited in the Zoolo- Finally, the subfamily Coccinellinae was gical Institute of the Russian Academy of mentioned for Florissant [Adalia subversa Sciences (St Petersburg). Scudder, 1900 (also by Wickham, 1912); An- The general classification of the fam- atis resurgens Wickham, 1917; Coccinella flo- ily Coccinellidae has several versions; the rissantensis Wickham, 1914; C. sodoma Wick- ideas developed by Sasaji (1968) and Kovár ham, 1913], the Oligocene of Brunnstatt (1996) were used in the paper. The recent [Aphidecta marginata (Foerster, 1891); Ada- monograph on the genus Rhyzobius (To- lia sp. (by Théobald, 1937)], the Latest Oli- maszewska, 2010) was rather an important gocene of Rott am Siebengebirge [Germany: source of information on the species of this Coccinella antiqua Heyden et Heyden, 1862; genus in addition to the recent specimens C. bituminosa Heyden et Heyden, 1866; C. from the collections of the Paris museum fossilis C. Heyden et L. Heyden, 1866; C. and the Zoological Institute, St Petersburg. krantzi C. Heyden et L. Heyden, 1866; C. Type strata. Lowermost Eocene, circa – prisca Schlechtendal, 1894; Sospita haagi (C. 53 Myr, Sparnacian, level MP7 of the mam- Heyden et L. Heyden, 1866); Lasia primitiva mal fauna of Dormaal. C. Heyden et L. Heyden, 1866], the Late Type locality. Farm Le Quesnoy, Chev- Miocene of Oeningen [Baden-Wurtemberg, rière, region of Creil, Oise department Germany: Coccinella amabilis Heer, 1865; (north of France). C. andromeda Heer, 1847; C. colorata Heer, 1865; C. heeri Ukrainsky, 2010 (=decempus- RESULTS tulata Heer, 1879, non Linnaeus, 1758); C. Order COLEOPTERA hesione Heer, 1847; C. perses Heer, 1847; C. ponomarenkoi Ukrainsky, 2010 (=spectabilis Family COCCINELLIDAE Latreille, 1807 Heer, 1865, non Faldermann, 1835)], and Subfamily COCCIDULINAE Mulsant, 1846 the Pliocene of Willershausen [Halyzia sp., Harmonia sp. and genus incertus (Gersdorf, Tribe COCCIDULINI Mulsant, 1846 1969)]. Thus, the species described in this Genus Rhyzobius Stephens, 1829 paper represent the oldest members of the family known at present. A more detailed Type species: Nitidula litura Fabricius, review of this coleopterous family in the fos- 1787. sil record can be taken from Ponomarenko & Notes. The specimens considered here Kirejtshuk (2012). were assigned to the genus Rhyzobius be- © 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE 133 cause of (a) large and coarsely faceted eyes, teristic shape of antennal club, configura- (b) relatively long antenna with flattened, tion of submeso- and submetacoxal lines. three-segmented club, (c) deeply split tarsal The new species are somewhat similar to claws and other features. All the specimens Rhyzobius confinis Lea, 1902, particularly are rather similar. Three specimens, which by the somewhat brownish abdomen, char- are visible in the dorsove ntral plane, have acters of puncturation and sculpture of in- differences in some characters that could be tegument, configuration of submeso- and considered as diagnostic ones for two sepa- submetacoxal lines and shape of antennal rate species. The remaining four specimens club. However, in contrast to the latter, are visible in pieces of amber from the later- the new species have the rather smaller al view and cannot be completely compared body size, pronotum with nearly bisinuate with the first three specimens. Neverthe- base and only slightly narrower than ely- less, these four specimens are placed in one tral ones, almost distinct posterior angles of the two new species recognized among and gradually narrowing anteriorly, shorter the specimens with the underside exposed hairs on the dorsum and antennal club more because they have characteristic pronotal distinctly oblique at apex. Nevertheless, sides. The recent species of this genus are a somewhat similar pronotum is known in spread in all continents except Antarctica, some other recent species. Rhyzobius cya- and the greatest diversity of the genus is neus Blackburn, 1889 (MNHN: “Eber. N. known from Australia. Gallyd. S.”) is, in contrast to both new spe- These new species differ from most re- cies, characterized by much larger and more cent congeners (about 60 recent species are slender body, bluish dorsum, yellow abdo- recorded in the genus after the recent revi- men, longer pubescence with erect hairs, sion by Tomaszewska, 2010) in the shape different configuration of submeso- and of pronotum which is comparatively wide submetacoxal lines, different antennal club at base, its posterior angles with an almost and ultimate maxillary palpomere. Rhyzobi- clear top, sides nearly gradually narrowing us pulchellus (Montrouzier, 1861) (MNHN: anteriorly and forming an even continu- “New Caledonia, Mueo, 19.V.1928, J.D.A. ous line with sides of elytra [in most recent Cockerell, on Citrus”) differs from the new members of the genus the pronotal base is species in the lighter and more slender markedly narrower than