Table Of ContentZOOSYSTEMATICA 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-
sediments from the Upper Pliocene of Wil-
shuk et al., 2010a,b; Kovalev et al., 2012).
lershausen (Niedersachsen, Germany).
The family Coccinellidae Latreille, 1807
Besides, among the inclusions from Bal-
is known in fossils only from some Cainozoic
tic amber Berendt (1845), Menge (1856),
resources, and no amber specimen has been
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-
Many specimens recovered among in-
linae Latreille, 1807). The subfamily Coc-
clusions from the Lowermost Eocene
cidulinae was also recorded in the Lower/
French amber are deposited in the Labora-
Middle Miocene of Shanwang Basin [Shan-
toire de Paléontologie, Muséum National
dong, China: Scymnus cf. kawamurai Ohta,
d’Histoire Naturelle, Paris (MNHN). For
1929 (by Zhang, 1989) and Hippodamia ol-
this study ordinary optic equipment was
bia J. Zhang, Sun et X. Zhang, 1994] and in
used, in particular the stereomicroscope
the Oligocene of Brunnstatt (Haut-Rhine,
Olympus SCX9 and inverted microscope
Elsas, France: Scymnus angulatus Förster,
Olympus CK 40 in the Paris museum, and
1891). The subfamily Chilocorinae was also
also the stereomicroscope Leica MZ 16.0
recorded in the Earliest Oligocene of Flo-
in the Zoological Institute, St Petersburg.
rissant (Colorado, USA: Chilocorus ulkei
All the holotypes and most paratypes of the
Scudder, 1900) and also from Brunnstatt
new species are deposited in the Paris muse-
[Chilocorus foersteri Ukrainsky, 2010 (=pol-
um, one paratype of Rhizobius antiquus sp.
itus Förster, 1891, non Mulsant, 1850) and
nov. (“PA 245”) is deposited in the Zoolo-
C. inflatus Förster, 1891].
gical Institute of the Russian Academy of
Finally, the subfamily Coccinellinae was
Sciences (St Petersburg).
mentioned for Florissant [Adalia subversa
The general classification of the fam-
Scudder, 1900 (also by Wickham, 1912); An-
ily Coccinellidae has several versions; the
atis resurgens Wickham, 1917; Coccinella flo-
ideas developed by Sasaji (1968) and Kovár
rissantensis Wickham, 1914; C. sodoma Wick-
(1996) were used in the paper. The recent
ham, 1913], the Oligocene of Brunnstatt
monograph on the genus Rhyzobius (To-
[Aphidecta marginata (Foerster, 1891); Ada-
maszewska, 2010) was rather an important
lia sp. (by Théobald, 1937)], the Latest Oli-
source of information on the species of this
gocene of Rott am Siebengebirge [Germany:
genus in addition to the recent specimens
Coccinella antiqua Heyden et Heyden, 1862;
from the collections of the Paris museum
C. bituminosa Heyden et Heyden, 1866; C.
and the Zoological Institute, St Petersburg.
fossilis C. Heyden et L. Heyden, 1866; C.
Type strata. Lowermost Eocene, circa –
krantzi C. Heyden et L. Heyden, 1866; C.
53 Myr, Sparnacian, level MP7 of the mam-
prisca Schlechtendal, 1894; Sospita haagi (C.
mal fauna of Dormaal.
Heyden et L. Heyden, 1866); Lasia primitiva
Type locality. Farm Le Quesnoy, Chev-
C. Heyden et L. Heyden, 1866], the Late
rière, region of Creil, Oise department
Miocene of Oeningen [Baden-Wurtemberg,
(north of France).
Germany: Coccinella amabilis Heer, 1865;
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 dorsoven tral 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
specimen of Trichoptera, 1 specimen of Ephem-
Remarks on probable bionomy. Like the
eroptera, 1 specimen of Hemiptera, 1 specimen
recent congeners, the two new species de-
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-
lotype because it is the most complete (although
(Figs 1–6, 9–13)
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
figuration of submeso- and submetacoxal
Balsam” medium and fixed between two square
lines. The pronotum of this new species,
coverslips. ‘PA 245’, probably female; nearly
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
onstrate such feature.
specimen is included in an amber bar (length 11
Etymology. The epithet of the new spe-
mm), semicircular in cross-section and with a
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 am-
men; dorsum with rather dense and dif-
ber parallelepiped (8.0 mm in length and about
fusely spread, long, very thin, moderately
4.0 mm in width of the widest facet); around the
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
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144
A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE 135
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.
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144
A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE 137
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)
distance between their roots), pronotal and
consisting of many layers, with two incomplete
elytral sides without clear cilia; underside
cracks going from the dorsum towards the flat
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
(a part of the right half of base of the pronotum,
tinct and deviating from posterior edge of
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.
Diagnosis. The new species is rather
half without a clear line along inner edge.
small and, in contrast to most of the recent
species, somewhat more oval; its epipleura
Subfamily SCYMNINAE Mulsant, 1846
are more gradually narrowing apically and
Tribe SCYMNINI Mulsant, 1846 more gradually outlined along inner edge
(extending along abdominal ventrites up
Genus Nephus Mulsant 1846
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
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144
A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE 139
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).
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144
140 A.G. KIREJTSHUK & A. NEL. OLDEST REPRESENTATIVES OF COCCINELLIDAE
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.
© 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 131–144
