The Oldest Representatives of the Family Coccinellidae(Coleoptera

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 dorsoventral 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 the elytral base; body, unbordered pronotal base, longer pu- usually pronotal sides at base are more or bescence with erect hairs, smoothed inter- less straight or sometimes the pronotum spaces between markedly larger and sparser is subquadrangular; posterior and anterior dorsal punctures, configuration of submeso- pronotal angles are more or less rounded]. and submetacoxal lines, shape of antennal The new species have uniform and com- club and ultimate maxillary palpomere, paratively short pubescence on the dorsum, much shorter ultimate tarsomere. Rhyzobi- although many recent pubescent species of us trimeni Casey, 1899 (MNHN, “Leap” and the genus demonstrate two types of hairs: “Cape Town, Dr Martin”) differs from the shorter and semirecumbent hairs are inter- new species in the somewhat larger body, mixed with longer and erect ones. In gener- yellow spots at anterior pronotal angles and al, the combination of the diagnostic char- elytral base, longer pubescence with erect acters of the new species, except those of the hairs, configuration of submeso- and sub- shape of pronotum and pubescence consid- metacoxal lines, shape of antennal club and ered above, is quite unique and includes the ultimate maxillary palpomere, much short- unicolorous dark and elongate oval body of er ultimate tarsomere. Finally, Rhyzobius medium size, dense and comparatively fine javeli Mulsant, 1899 (MNHN: “Leap” and dorsal puncturation, raised microsculpture “Cape Town, Dr Martin”) differs from the on interspaces between punctures, charac- new species in somewhat larger, markedly

© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144 134 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE more slender and much lighter body, un- Additional specimen. ‘PA 5162’, sex unknown, bordered pronotal base, longer tarsal lobes, badly preserved specimen with right half of its longer pubescence with erect hairs. body destroyed is included together with 1 Remarks on probable bionomy. Like the specimen of Trichoptera, 1 specimen of Ephem- recent congeners, the two new species de- eroptera, 1 specimen of Hemiptera, 1 specimen of Fulgoroidea and some separate organs or scribed here could be predaceous or could sclerites of different insects in mostly unpolished have some associations with colonies of elongate amber piece with mixture of layers, scale insects that appeared before the Cai- many small gas bubbles and different grains of nozoic. organic matter. The beetle is obscured with an ephemeropteran wing and not clearly visible. Rhyzobius antiquus sp. nov. The specimen ‘PA 11862’ is chosen as the ho- (Figs 1–6, 9–13) lotype because it is the most complete (although the piece of amber including it and two other in- Holotype. ‘PA 11862’, male with slightly ex- sects also contains many small gas bubbles mak- posed genital capsule; complete specimen is in- ing it difficult to observe the whole beetle but cluded in a small irregular elongate amber paral- allowing one to study most structures separate- lelepiped (8.0 mm in length and about 3.0 mm ly). The mandibles, palpi, procoxae and procoxal in width of one facet) with many small pieces of process of all the specimens examined are not organic matter and small gas bubbles diffusely visible because of the “milky” cover. However, spread throughout it; besides, there are a very the labial and maxillary palpi of the paratype small specimen of Nematocera and a very small ‘PA 245’ can be more or less clearly observed, larva of Thysanoptera near the posterior half of whereas only the maxillary palpi are more or less the beetle underside; most sclerites of the beetle visible in other specimens. are covered with “milky” cover. Diagnosis. This new species differs from Paratypes. ‘PA 972’, female; broken specimen the other congener from the same resource with somewhat exposed apex of the left wing but in the more oval body, shape of pronotum, with most part of dorsal sclerites missing is in- narrower prosternal process, greater rela- cluded in a very small irregular elongate amber tive distance between mesocoxae and con- parallelepiped, which is embedded in “Canada Balsam” medium and fixed between two square figuration of submeso- and submetacoxal coverslips. ‘PA 245’, probably female; nearly lines. The pronotum of this new species, complete specimen with somewhat exposed in contrast to that in R. gratiosus sp. nov., apices of both hind wings is included together is comparatively narrow at base and with with a small immature dipteran larva below the more arcuate sides. The holotype of this left part of the beetle underside in a very small new species has a clear line along the inner irregular elongate amber parallelepiped with edge of epipleura, while the paratype have many small gas bubbles, which is embedded in well visible epipleura (‘PA 245’) and the ho- “Canada Balsam” and fixed between two rect- lotype of R. gratiosus sp. nov. do not dem- angular coverslips. ‘PA 5388’, female; complete specimen is included in an amber bar (length 11 onstrate such feature. mm), semicircular in cross-section and with a Etymology. The epithet of the new spe- flat plane of 5 mm in width, also including many cies is the Latin adjective meaning “an- layers and many small pieces of organic matter, cient” or “archaic”. very small cracks; the beetle is visible laterally Description . Holotype, male. Length 2.8, and most of its sclerites are covered with “milky” width 2.0, height 1.2 mm. Oval, strongly cover of different thickness. ‘PA 4840’, sex un- convex dorsally and moderately ventrally; known; specimen with broken distal part of the unicolorous dark brown to blackish with body, which is out of a rather clear piece of am- brown to brownish appendages and abdo- ber, is included in a small irregular elongate amber parallelepiped (8.0 mm in length and about men; dorsum with rather dense and dif- 4.0 mm in width of the widest facet); around the fusely spread, long, very thin, moderately beetle there are some very small cracks, particu- conspicuous brownish, subrecumbent hairs larly at head and prothorax. (3–5 times as long as the distance between

Figs 1–6. Rhyzobius antiquus sp. nov. 1, 2, body of holotype (‘PA 11862’); 3, antenna of paratype (‘PA 972’); 4, labial palpus of paratype (‘PA 245’); 5, mesofemur of paratype (‘PA 972’); 6, hind leg of paratype (‘PA 972’). Ventral (1, 3, 4) and dorsal (2) view. Scale bars: 1.0 mm; A – to figs 1, 2; B – to figs 3–6. their roots), pronotal and elytral sides surface of some places of elytra and thorac- without clear cilia; underside with some- ic sclerites with very fine and sparse punc- what similar pubescence consisting of less tures, interspaces between them finely and conspicuous hairs (which are somewhat smoothly alutaceous. sparser on thoracic sclerites). Sculpture Head transverse and somewhat declined and puncturation of integument mostly not (subhypognathous) and scarcely visible visible because of “milky” cover, although dorsally, much narrower than pronotum,

© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144 136 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE with very large and coarsely faceted eyes pygidium comparable in length and some- bearing clear interfacetal setae. Mandibles what longer than each of ventrites 3 and 4, very small and scarcely exposed from un- posterior edge of hypopygidium transverse der frons. Pronotum slightly narrower than to shallowly emarginate. Epipleura gently combined elytral base, about twice as wide outlined, at anterior third their plane with as long, widest at base and gradually arcu- a curve visible laterally and in anterior half ately narrowing anteriorly towards widely with a line along inner edge. rounded anterior angles, moderately and Legs well developed, moderately narrow gently vaulted; its anterior edge slightly and long, diffusely covered with compara- emarginate; its posterior edge indistinctly tively short setae. Tibiae moderately com- bordered, strongly convex at the middle pressed, comparable in width and shape, and slightly emarginate at sides; posterior narrowing at apex by a comparatively wide angles with distinct top. Scutellum looking isolated stripe reaching place of insertion of like a subequilateral triangle. Elytra some- tarsus. Femora of usual shape and slightly what shorter than wide combined, longest compressed, about 2.5 times as wide as pro- at suture and nearly regularly arcuate along tibiae. Tibial spur not raised. Tarsi tetram- sides, rather steeply sloping laterally (with erous and moderately long, about 2/3 as lateral edges visible dorsally) and with ex- long as tibiae, tarsomeres 1 and 2 with very tremely narrowly explanate edges, adsutur- wide lobes and wider than tibiae; ultimate al lines not visible. Pygidium with widely tarsomeres much longer than tarsomeres rounded to subtruncate apex. Anal sclerite 1 and 2 combined; claws strongly dentate, well exposed from under pygidial apex. about 1/4 as long as ultimate tarsomere, Most part of underside not clearly vis- apex of dens nearly reaching apex of claw. ible because of optical aberration in differ- Paratypes. ‘PA 972’: length 3.0 mm. ent layers of amber and rather thick “milky Head slightly convex and with frons slight- cover”. Procoxae moderately large, trans- ly extending beyond the anterior edge of verse and moderately narrowly separated. eyes and clear left maxillary palpus. La- Prosternal median part whether is isolated brum far projecting anteriorly, slightly less from lateral parts and process subparallel- than twice as wide as long and truncate at sided or slightly widened at apex, where it anterior edge. Antennae narrow and about is about one-third as wide as the distance as long as head width, 3 segmented club between metacoxae. Mesocoxae apparently (with widest ultimate segment) compris- subtransversely oval and widely separated ing nearly 2/7 of total length, flagellomeres (somewhat less widely separated than meta- between pedicel and club subcylindrical coxae). Metaventrite subflattened along and comparable in length, rather elongate. the middle, posterior edge between coxae Hypopygidium widely rounded at apex. ‘PA straight. Submesocoxal line distinct and 245’: length 2.7, width 1.8 mm. Specimen deviating from posterior edge of mesocoxal nearly subunicolorous blackish. Distances cavity at its median part, then arcuately between pro-, meso- and metacoxae as 1:3:4; joining to the inner edge of metepisterna labial comparatively short and wide, with before the middle of the latter. Metepister- ultimate palpomere obliquely transverse at na moderately narrow and subparallel-sid- apex. ‘PA 388’: length 2.6, height 1.1 mm. ed. Metacoxae transversely oval, somewhat Dorsum with more conspicuous and some- less wide than the distance between them. what longer pubescence. ‘PA 4840’: length Abdominal ventrite 1 longest, submetacox- 2.6, width 1.6, height 1.0 mm; dorsal pubes- al lines arcuately deviating from posterior cence much less conspicuous than that in edge of the cavity and returning to the lat- other specimens of the type series; subme- ter at outer edge – this line reaching distal socoxal line almost not returning anteriorly fifth of the ventrite; ventrite 2 and hypo- at the edge of metepisternum.

Additional specimen. ‘PA 5162’, length Etymology. The epithet of this new spe- nearly 3.0 mm; the general outline of body, cies is formed from the Latin “gratia” (grace puncturation and sculpture of the dorsum, or graceful) and “-osus” (means abundan- and particularly posterior angles of pro- ce of). notum are rather similar to those of other Description. Holotype (taking into con- specimens of R. antiquus sp. nov., however, sideration a great similarity of this new spe- the dorsal hairs are clearly longer and seem cies to the previous one, most characters to have some difference in length as in many shared by both species are omitted in the recent species. description below). Length 2.6, width 1.7, height about 1.2 mm. Elongate oval, strong- Rhyzobius gratiosus sp. nov. ly convex dorsally and moderately ventral- (Figs 7–8, 14–15) ly; unicolorous dark brown to blackish with slightly lighter tarsi; dorsum with rather Holotype. ‘PA 1290’, probable female (ab- dense and diffusely spread, long, very thin, dominal apex not visible clearly); complete moderately conspicuous brownish, sub- specimen is included in a flat piece of amber of recumbent hairs (3–5 times as long as the irregular triangular shape (17, 11 and 14 mm) consisting of many layers, with two incomplete distance between their roots), pronotal and cracks going from the dorsum towards the flat elytral sides without clear cilia; underside plane of the amber piece; most sclerites of the with somewhat similar pubescence con- beetle are covered with “milky” cover. sisting of less conspicuous hairs. Sculpture Diagnosis. See the diagnosis of the previ- and puncturation of integument mostly not ous new species. visible because of “milky” cover, although

Figs 7–8. Rhyzobius gratiosus sp. nov., holotype (‘PA 1290’). 7, body, dorsal view; 8, metatorax with abdomen, ventral view. Scale bar: 1.0 mm.

© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144 138 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE surface of some places of elytra and thoracic sence of both projection of anterior edge sclerites with very fine and sparse punc- and longitudinal ridges of its prosternum. tures, interspaces between them finely and The characters accessible in the specimen smoothly alutaceous. examined do not make it possible to find a Pronotum at base about as wide as com- strict subgeneric attribution for it. bined elytral base, about twice as wide as Remarks on probable bionomy. Like the long, widest at base and gradually, nearly recent congeners, the new species described rectilinearly narrowing anteriorly to widely here could be predaceous or could have rounded anterior angles, moderately and some associations with colonies of parane- gently vaulted; its anterior edge slightly opteran groups (including aphids and scale emarginate; its posterior edge indistinctly insects) which could also exist in the early bordered, strongly convex at the middle and Eocene (while pseudococcids seem to have slightly emarginate at sides; posterior angles appeared later). with distinct top. Scutellum subcardiform. Elytra about as long as wide combined. Py- Nephus (subgenus incertus) gidium widely rounded at apex. Prosternal subcircularis sp. nov. median part not isolated from lateral parts (Figs 16–20) and process slightly widened to apex, where it is at least half as wide as the distance be- Holotype. ‘PA 1047’, male with somewhat ex- tween metacoxae. Submesocoxal line dis- posed genital capsule; almost complete specimen tinct and deviating from posterior edge of (a part of the right half of base of the pronotum, and base of the right elytron are cut) is included mesocoxal cavity at its median part, then in a small irregular elongate amber piece (8.0 mm arcuately joining to the inner edge of met- in length and about 5.0 mm in width of the wid- episterna behind the middle of the latter. est facet) with some small pieces of organic mat- Abdominal ventrite 1 longest, submetacoxal ter and small cracks diffusely spread through- lines arcuately deviating from posterior out; some sclerites of the beetle is covered with edge of the cavity and returning to the lat- “milky” cover. ter at outer edge – this line reaching distal The antennae, mouthparts and procoxae of fourth of the ventrite. Epipleura in anterior the examined specimen are not clearly visible. half without a clear line along inner edge. Diagnosis. The new species is rather small and, in contrast to most of the recent Subfamily SCYMNINAE Mulsant, 1846 species, somewhat more oval; its epipleura are more gradually narrowing apically and Tribe SCYMNINI Mulsant, 1846 more gradually outlined along inner edge Genus Nephus Mulsant 1846 (extending along abdominal ventrites up to ventrite 4); tarsi of the new species are Type species: Sphaeridium quadrimacu- somewhat narrower and its underside is less latum Herbst, 1783. convex. Notes. The species described here has Etymology. The epithet of the new spe- submeso- and submetacoxal lines similar to cies is the complex adjective of “sub” (near- the species of the genus Nephus rather than ly) and “circularis” (circular). other genera with small members. This ge- Description. Holotype, male. Length 1.2, nus includes about two hundred recent spe- width 1.0, probable height 0.4 mm. Short cies and is divided into various subgenera. oval, strongly convex dorsally and moder- Like other congeners, the new species is ately ventrally; unicolorous dark brown to characterized by the incomplete submeta- blackish with brown appendages; dorsum coxal line (not reaching the lateral edge of with comparatively sparse and diffusely ventrite 1), pseudotrimerous tarsi (with a spread, not long, very thin, moderately very small intercalary tarsomere) and ab- conspicuous brownish, subrecumbent hairs

Figs 9–12. Rhyzobius antiquus sp. nov. 9, 10, body of holotype (‘PA 11862’), length of body – 2.8 mm; 11, body of paratype (‘PA 972’), length of body – 3.0 mm ; 12, head of paratype (‘PA 972’). Dorsal (9), ventral (10) and lateroventral view (11, 12).

Figs 13–16. Rhyzobius spp. 13, Rhyzobius antiquus sp. nov., body of paratype (‘PA 245’), length of body – 2.7 mm; 14, 15, Rhyzobius gratiosus sp. nov., body of holotype (‘PA 1290’), length of body – 2.6 mm; 16, Nephus subcircularis sp. nov., body of holotype (‘PA 1047’), length of body – 1.2 mm. Dorsal (14) and ventral (13, 15, 16) view.

Figs 17–20. Nephus subcircularis sp. nov., holotype (‘PA 1047’). 17, body; 18, antennal club; 19, protarsus; 20, tarsal claws. Dorsal (19) and ventral (17, 18) view. Scale bars: A – to fig. 17, 0.3 mm; B – to figs 18, 19, 0.7 mm

(somewhat shorter than the distance be- but there are very fine and very sparse dif- tween their roots), pronotal and elytral fuse punctures and rather smoothed (alu- sides without clear cilia; underside with taceous) broad interspaces between them; somewhat similar pubescence consisting abdominal ventrites with somewhat denser of less conspicuous and somewhat shorter and coarser punctures (with interspaces hairs (which are somewhat sparser on tho- somewhat greater than a puncture diame- racic sclerites). Sculpture and punctura- ter); metaventrite with yet larger punctures tion of part of dorsal integument mostly and interspaces about as great as a puncture not clearly visible because of very fine and diameter or very slightly greater and nearly dense cracks intermixed with “milky” cover, completely smooth.

Head transverse and somewhat declined while returning to the posterior edge of cav- (subhypognathous) and somewhat extend- ity and not joining with lateral edge of ven- ing anteriorly from anterior edge of prono- trite – this line reaching distal fourth of the tum, moderately narrower than pronotum, ventrite; ventrite 2 and hypopygidium com- with moderately large and not coarsely fac- parable in length and somewhat longer than eted eyes. Mandibles very small and scarce- each of ventrites 3 and 4, posterior edge of ly exposed from under frons. Antennal club hypopygidium emarginate. Epipleura about three-segmented, only slightly narrower as wide as prosternal process, gently out- than maxillary palpomere and with larg- lined along outer and inner edges gradually est penultimate antennomere. Pronotum narrowing apically and extending along ab- markedly narrower than combined elytral dominal ventrites to ventrite 4; apparently base, nearly 2.5 times as wide as long, wid- only with a slight curve visible laterally. est along posterior half and from the middle Legs well developed, moderately narrow gradually narrowing anteriorly to widely and long, diffusely covered with compara- rounded anterior angles, moderately and tively short setae. Tibiae moderately com- gently vaulted; its anterior edge subtrun- pressed, comparable in width and shape, cate to subemarginate; its posterior edge narrowing at apex by a comparatively wide moderately strongly and regularly convex; isolated stripe reaching place of insertion of posterior angles rounded. Scutellum miss- tarsus. Femora of usual shape and slightly ing. Elytra apparently somewhat longer compressed, about 2.5 times as wide as pro- than wide combined, longest at suture and tibiae. Tibial spurs not raised. Tarsi mod- nearly regularly arcuate along sides, rather erately long, about 3/5 as long as tibiae, convex and gradually steeply sloping; later- tarsomeres 1 and 2 with long and not wide al edges very narrowly explanate; adsutural lobes (narrower than tibiae); ultimate tar- lines not visible. Pygidium with widely someres much longer than tarsomeres 1 and rounded apex. Anal sclerite well exposed. 2 combined; claws strongly dentate, about Considerable part of underside not 1/4 as long as ultimate tarsomere, apex of clearly visible because of optical aberra- dens reaching the middle of the claw length. tion in different layers of amber and partly because of “milky cover”. Maxillary palpi DISCUSSION apparently moderately short and wide, ul- The new species described here belong timate palpomere about as long as thick to genera whose recent species have a preda- and obliquely truncate at apex. Prosternum ceous mode of life, although it might scarce- somewhat medially convex, but not with ly be the initial one for the family. Different longitudinal ridges along the middle and its phylogenetic reconstructions (Sasaji, 1968; anterior edge not projecting anteriorly. Pro- Yu, 1994; Kovár, 1996, etc.) assume that coxae moderately large, transverse, rather the tribes Coccidulini and Scymnini are narrowly separated. Mesocoxae apparently not very close to the ancestor of the family. subtransversely oval and widely separated Therefore, it may be supposed that a con- (nearly as widely as metacoxae). Metaven- siderable diversification of the family with trite slightly convex in the middle, posterior probably mycetophagous archaic members edge between coxae straight. Submesocoxal took place before the time of the amber de- line more or less distinct, arcuately return- position in Oise. ing anteriorly at metepisterna. Metepi- sterna moderately narrow and subparallel- sided. Metacoxae transversely oval, about ACKNOWLEDGEMENTS as wide as the distance between them. Ab- The authors thank the company Lafarge- dominal ventrite 1 longest, submetacoxal Granulat for help with the sampling of the fos- lines not complete and becoming obsolete sil and the family Langlois-Meurinne for the

© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE 143 authorization of working in their property. They Hieke F. & Pietrzeniuk E. 1984. Die Bernstein- are also grateful to G. De Ploeg (MNHN) and Käfer des Museums für Naturkunde, Berlin D. Azar (Lebanese University) for the care- (Insecta: Coleoptera). Mitteilungen der Zoo- ful preparation of the material. The paper was logische Museum, 60(2): 297–326. prepared in the Muséum National d’Histoire Kirejtshuk A.G. & Nel A. 2008. Some new fos- Naturelle in Paris in the framework of the sils of the suborder Polyphaga (Insecta, Co- Programme “Research in Paris” (programme leoptera) from Lowermost Eocene French d’accueil des chercheurs étrangers de Mairie amber. Annales de la Société Entomologique de Paris). The work of the first author has been de France, (N.S.), 44: 419–442. supported for several years by the Muséum Na- Kirejtshuk A.G. & Nel A. 2009. New genera tional d’Histoire Naturelle according to the Pro- and species of Cucujiformia (Coleoptera, gramme of visiting professors, Programme of the Polyphaga) from lowermost Eocene French Presidium of the Russian Academy of Sciences amber. Denisia, 26: 103–118. “Problems of Origin of Life and Formation of Kirejtshuk A.G., Nel A., Collomb F.-M. 2009. Biosphere,” and his work with the collection of New Archostemata (Insecta, Coleoptera) the Zoological Institute was supported by the from the French Paleocene and Early Eo- Russian Ministry of Education and Science and cene, with a note on the composition of the a grant from the Russian Foundation for Basic suborder. Annales de la Société Entomolo- Research (12-04-00663-а). The authors greatly gique de France, (N.S.), 45: 216–227. appreciate the assistance of Stanislaw Adam Kirejtshuk A.G., Nabozhenko M.V. & Nel A. Ślipiński (Division of Entomology, CSIRO, 2010. New genus and species of the tribe Canberra) and Wioletta Tomaszewska (Museum Opatrini (Coleoptera, Tenebrionidae, Tene- and Institute of Zoology of the Polish Academy brioninae) from the Lowermost Eocene am- of Sciences, Warszawa) during elaboration of ber of Paris basin. Proceedings of the Zoologi- the diagnoses for the new species described here. cal Institute RAS, 314(2): 191–196. The preparation of drawings was made with as- Kirejtshuk A.G., Hava A J. & Nel A. 2010. New sistance of the first author’s daughter, Polina genus and species of subfamily Trinodinae A. Kirejtshuk. The authors appreciate very use- (Coleoptera, Polyphaga, Dermestidae) from ful comments and recommendations of anony- Lowermost Eocene French amber. Zoosyste- mous reviewers. matica Rossica, 19(1): 54–69. Klebs R. 1910. Uber Bernsteineinschlusse in all- REFERENCES gemeinen und die Coleopteren meiner Bern- steinsammlung. Schriften der Physikalischen- Batelka J., Collomb F.-M, Nel A. 2006. Macro- Okonomische Gesellschaft zu Königsberg, 51: siagon deuvei n. sp. (Coleoptera: Ripiphori- 217–242. dae) from the French Eocene amber. Annales Kovalev A.V., Kirejtshuk A.G. & Nel A. 2012. de la Société Entomologique de France, (N.S.), New species of the genus Trixagus Kugelann, 42: 75–78. 1794 (Coleoptera: Throscidae) from the Berendt G.C. 1845. Die im Bernstein befindli- Lowermost Eocene amber of Oise (France). chen Organischen Reste der Vorwelt gesam- Proceedings of the Zoological Institute RAS, melt in Verbindung mit mehreren bearbeitet. 316(1): 83–87. Erster Band. Abtheilung I. Der Bernstein und Kovár I. 1996. Phylogeny. In: Hodek I. & Hon- die in ihm befindlichen Pflanzenreste der Vor- ek A. (Eds.) Ecology of Coccinellidae: 19–31. welt. Danzig [= Gdansk], iv + 125 p. Dordrecht: Kluwer Academic Publishers. Bílý S. & Kirejschuk A. G. 2007. Philanthaxo- Kubisz D. 2000. Fossil beetles (Coleoptera) ides gallicus gen. n., sp. n. from the Lowermost from Baltic amber in the collection of the Eocene French amber (Coleoptera: Bupresti- Museum of Natural History of ISEA in Kra- dae). Folia Heyrovskyana, A, 14: 181–186. kow. Polish Journal of Entomology, 69(2): Gersdorf E. 1969. Käfer (Coleoptera) aus dem 225–230. Jungtertiär Norddeutschlands. Geologisches Larsson S.G. 1978. Baltic amber – A palaeobio- Jarbuch, 87: 295–331. logical study. Entomonograph, 1: 1–192. Grimaldi D.A. & Engel M.S. 2005. Evolution of Menge F.A. 1856. Lebenszeichen vorweltlicher the insects. Cambridge University Press. xv + im Bernstein eingeschlossener Thiere (Hete- 755 p. roptera, Coleoptera, Lepidoptera, Hymeno-

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