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Annales de la Société entomologique de France (N.S.)
International Journal of Entomology
ISSN: 0037-9271 (Print) 2168-6351 (Online) Journal homepage: http://www.tandfonline.com/loi/tase20
New beetles of the suborder Polyphaga from
the Lowermost Eocene French amber (Insecta:
Coleoptera)
Alexander G. Kirejtshuk & André Nel
To cite this article: Alexander G. Kirejtshuk & André Nel (2008) New beetles of the suborder
Polyphaga from the Lowermost Eocene French amber (Insecta: Coleoptera), Annales de la
Société entomologique de France (N.S.), 44:4, 419-442, DOI: 10.1080/00379271.2008.10697578
To link to this article: http://dx.doi.org/10.1080/00379271.2008.10697578
Published online: 31 May 2013.
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Ann. soc. entomol. Fr. (n.s.), 2008, 44 (4) : 419-442
419
ARTICLE
New beetles of the suborder Polyphaga from the Lowermost
Eocene French amber (Insecta: Coleoptera)
Abstract. The paper deals with the descriptions of t hree new genera, one new subgenus and eleven new
species of the superfamilies Scirtoidea, Cleroidea, and Cucujoidea (suborder Polyphaga) originated
from the Lowermost Eocene amber, namely: the Scirtidae: Cyphon gallicus nov. sp. and Cyphon
lobanovi nov. sp.; the Melyridae: Malachiinae Colotes constantini nov. sp. and C. impexus nov. sp.;
the Nitidulidae: Cybocephalinae Pastillocenicus polyaki nov. gen., nov. sp., and P. gra nd ic lavis nov.
gen., nov. sp. and P. longifro ns nov. gen., nov. sp.; the Kateretidae: Hetherelus expressus nov. sp.
and Eoceniretes yantaricus nov. gen., nov. sp.; the Smicripidae: Smicrips europeus nov. sp.; and the
Anthicidae: Eurygeniinae Oisegenius antiquus nov. gen., nov. sp. The systematic positions of these
new taxa and hypotheses on their bionomy are discussed.
Résumé. Nouveaux Polyphaga de l’ambre éocène basal français (Insecta : Coleoptera). Tro is
nouveaux genres, un sous-genre et onze nouvelles espèces de Scirtoidea, Cleroidea et Cucujoidea
(sous-ordre Polyphaga) sont décrits de l’ambre éocène basal de France. Il s’agit des Scirtidae : Cyphon
gallicus nov. sp. et Cyphon loban ovi nov. sp. ; des Melyridae : Malachiinae : Colotes constantini nov. sp.
et C. impexus nov. sp. ; des Nitidulidae : Cybocephalinae: Pastillocenicus polyaki nov. gen., nov. sp.
et P. gra nd ic lavis nov. gen., nov. sp. et P. longifrons nov. gen., nov. sp. ; des Kateretidae : Hetherelus
expressus nov. sp. et Eoceniretes yantaricus nov. gen., nov. sp . ; du Smicripidae: Smicrips europeus
nov. sp. ; et de l’Anthicidae : Eurygeniinae : Oisegenius antiquus nov. gen., nov. sp. Les positions
systématiques de ces taxons nouveaux et des hypothèses sur leurs bionomies sont discutées.
Keywords: Scirtidae, Melyridae, Nitidulidae, Kateretidae, Smicripidae, Anthicidae, France.
A G. K (1) & A N (2)
(1) Zoological Institute of the Russian Academy of Sciences, Universitetskaya emb. 1, St. Petersburg, 199034, Russia
(2) CNRS UMR 5202, CP 50, Entomologie, Museum national d’Histoire naturelle, 45 rue Buff on, F-75005 Paris, France
E-mail: agk@zin.ru, ak3929@ak3929.spb.edu, alexander_kirejtshuk@
yahoo.com, anel@mnhn.fr
Accepté le 29 juillet 2008
France is known by many places, where amber of
diff erent age was deposited (Lacrois 1910; Schlüter
1978; etc.). Since 1996 a great number of amber inclu-
sions has been obtained in the outcrop with the Low-
ermost Eocene sediments in the Oise Department.
Recently these inclusions have been started to investi-
gate by specialists on diff erent groups of animals and
plants (Nel et al. 2004). e crucial diff erences of this
source from Baltic amber are connected with the age
and taxonomic attribution of resin producing plant.
e age depository of French amber in Oise falls on
the end of the ‘thermoera’, while Baltic amber more or
less coincides with the beginning ‘crioera’. e infrared
spectrum of the French amber is rather similar to that
of the recent Hymenaea copal (Nel et al. 2004), while
the resin for Baltic amber seemed to be produced by
coniferous plants. ese diff erences make possible to
explain an essential distinction in composition of these
amber entomofaunas. For now about 300 specimens
were determined as beetles among 20 thousands inclu-
sions deposited in MNHN. is is only a small portion
of the inclusions prepared for study and yet examined.
Among them there are recognized the following fami-
lies: Cupedidae, Micromalthidae, Carabidae, Staphy-
linidae, Scydmaenidae, Pselaphidae, Catopidae, Scirti-
dae, Eucinetidae, Elateridae, Eucnemidae, Bupresti-
dae, Dermestidae, Cleridae, Anobiidae, Malachiidae,
Nitidulidae, Phalacridae, Cryptophagidae, Kateretidae,
Smicripidae, Silvanidae, Coccinellidae, Endomychidae,
Corylophidae, Latridiidae, Ciidae, Scraptiidae, Me-
landryidae, Mordellidae, Rhipiphoridae, Mycteridae,
Anthicidae, Aderidae, Chrysomelidae, Apionidae, and
Curculionidae. e representatives of Archostemata,
Buprestidae and Rhipiphoridae have been described in
separate papers (Batelka et al. 2006; Bílý & Kirejtshuk
2007; Kirejtshuk et al. in press a). Some further species
are considered in this publication. Nevertheless, even
the forms here described give some evidence of a com-
plex origin of this fauna. Particularly some fossils show
how many unexpected fi ndings should be obtained in
this staff of amber, viz. some Cybocephalinae (earliest
record of the subfamily), parasitoids of coccids spread
in recent conditions mostly in areas with warm sub-
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420
A. G. K & A. N
tropical climate, and a member of the genus Smicrips
Leconte 1878 (Smicripidae) (fi rst fossil record of the
family), characteristic of recent fauna of Central and
South America.
At the same time the characteristic of both Baltic
and French amber is the comparative abundance of
representatives of Aderidae, probably connected with
certainty of the period of resin production in both ‘Baltic’
and ‘French’ forests. Probably this period in both cases
coincided with time of most pest attack. In modern
forests of the North Hemisphere this time is more or
less stretched through the spring and early summer,
although in some areas with more intertropical climate
this family is one most abundantly represented in the
Recent epoch (for example, islands of West Indies). e
emergence of adults of Aderidae usually coincides with
fl owering of many angiosperms. Perhaps, frequency
of ‘stellate hairs’ (of Quercus origin?) in Baltic amber
supports this conclusion. However, Aderidae are well
represented in the Lower Cretaceous Lebanese amber
(Kirejtshuk & Azar in press). In contrast to the Baltic
fauna, this representation of Aderidae in the ‘French’
forest is combined with a more important frequency of
Corylophidae, while that in Baltic amber agrees with a
great representation of Scirtidae, which are present by
only few specimens in the French Eocene amber.
Spahr (1981) and Ponomarenko & Kirejtshuk
(2008) published references to the coleopterous
inclusions in amber of diff erent origins.
Among other things some species of the subfamily
Alticinae and other leaf beetles (Chrysomelidae) have
been discovered in this French amber. is group is
more characteristic of grass habitats than arboreal ones
and it can be regarded as an indicator of grassland biotas,
although some Alticinae and Eumolpinae from this
French amber seemed to be associated with tree canopy.
However, some genera found in the Baltic amber are
not represented in the French amber. ey belong, for
example, to the Scirtidae and Nitidulidae, although
seven specimens of Cyphon Paykull 1799 and Scirtes
Illiger 1807 from the fi rst family (three of them are
here described), six specimens of Cybocephalinae from
the second have been found among the sorted material
(species of Cybocephalinae are here described).
Material and methods
e material under consideration is deposited in the Laboratoire
de Paléontologie, Muséum national d’Histoire naturelle, Paris,
although one paratype of Colotes constantini nov. sp. (PA 2764)
and one paratype of P. longifrons nov. gen., nov. sp. (PA 2451)
are deposited in the collection of the Zoological Institute of the
Russian Academy of Sciences (St. Petersburg).
e type stratum is Lowermost Eocene, in amber, circa - 53
Myr, Sparnacian, and level MP7 of the mammal fauna of
Dormaal. e type locality is Farm Le Quesnoy, Chevrière,
region of Creil, Oise Department (north of France).
Systematic palaeontology
Infraorder Elateriformia Crowson 1960
Superfamily Scirtoidea Fleming 1821
Family Scirtidae Fleming 1821
(= Sinodryopitidae Hong 2002)
Specimens of this family are quite common in the
Upper Eocene Baltic amber (Iablokoff -Khnzorian
1961; Klausnitzer 1976, 2004; Kubisz 2000) and
recorded from the Eocene Fushun amber (Hong 2002),
although it is known also from one limestone site of
sedimentary deposits with many fi ndings of this family
at the boundary between the Oligocene and Eocene
(Bembridge Marls: Kirejtshuk et al. in press b). Besides,
many representatives of this family are present in the
Upper Jurassic Karatau (still remaining undescribed
in the collection of the Palaeontological Institute of
the Russian Academy of Sciences, Moscow) and some
other outcrops of diff erent ages. Members of this family
were found in the Lower Cretaceous Burmese amber
(Grimaldi et al. 2002) and in the Lower Cretaceous
Lebanese amber (Kirejtshuk & Azar in press). e
genus Miocyphon Wickham, 1914 from the Florissant
shales (Lower Oligocene) can be scarcely considered
as related to Scirtidae (Kirejtshuk et al. in press b).
In contrast to Scirtidae, according to the original
description and drawings, its body is comparatively
large (8 mm long), its prothoracic segment is not rather
short, its head is not clearly declined, and its elytral
apices are subtruncate. e position of Sinodryopidae
is discussed in the latter publication.
All the recent Scirtidae, and especially Cyphon, with
known bionomy need water for larval development,
because their larvae fi lter algae and other small
particles from liquid substrates. Adults are usually
adhered to vegetation near water basins. erefore,
marsh beetles not infrequently use for such purpose
water-fi lled tree holes, leaf bases, wet soil, rotten logs,
and similar sources of water. Sometimes larvae of some
recent species develop in groundwater as deep as 10
meters. Adults of some species can occur in many dry
localities, including even arid places in deserts.
Genus Cyphon Paykull 1799
Type species. Cistela pallida Fabricius 1775 (by subsequent designation
by Westwood 1838), recent.
e specimens here considered as new members of
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Polyphaga from Eocene French amber
421
the genus Cyphon, in contrast to other scirtid groups with
oval and convex body, are characterized by the narrow
metafemora, gently outlined anterior and lateral edges
of pronotum, pronotal base only slightly narrower than
elytral base and not modifi ed antennae. is genus is
very abundant in both recent fauna and fossil records.
About 100 recent species spread throughout diff erent
zoogeographical regions; however, the most of them
are recorded in the Northern Hemisphere. Iablokoff -
Khnzoryan (1961) described three species of Cyphon
from the Baltic amber. Klausnitzer (2004) described
three other species from the Baltic amber with exposed
apical part of the aedeagus, which have various
structures of the lateral lobes of tegmen. Hong (2002)
fi gured a fossil from the Fushun amber (China) without
species epithet, which, however, is certainly a Scirtidae,
but the latter scarcely belongs to the genus Cyphon at
all because of its much strongly declined metacoxae.
Recently some species of this genus were recovered
from Bembridge Marls (boundary between Oligocene
and Eocene: Kirejtshuk et al. in press b). us, one of
the species described herein (Cyphon gallicus nov. sp.)
is the oldest representative of the genus. Another new
species (C. lobanovi nov. sp.) preliminarily put in this
genus supports a rather ancient origin for the diversity
of this family.
Cyphon gallicus nov. sp.
(Figs 1–8; 70)
Material. Holotype PA 3404, undefi ned sex [in a narrow
amber stick, the beetle with missing left metatibia and tarsus
as well as missing terminal antennomeres, together with a small
Nematocera in front of it and a surface mould of Nematocera
dorsally and behind the beetle. e consistence of the amber
substrate is not homogenous, with some layers of very soft
substrate and, therefore, surface of the amber is not very regular.
Small pieces of organic matter diff usely spread throughout
the amber piece and some concentration of them is observed
around mouthparts of the beetle].
Paratype PA 3404, undefi ned sex [in a triangular amber bar,
apparently the complete beetle visible only from above. e
consistence of the amber substrate is rather heterogenous, with
many layers of very soft substrate are below the beetle. Small
pieces of organic matter, small gas bubbles and cracks diff usely
spread throughout the amber piece with some concentration
along the underside].
Note. e paratype and holotype of this new species
are regarded as conspecifi c because they have the
comparable proportion of visible dorsal sclerites,
the same character of the dorsal puncturation and
sculpture and, particularly, the character of the dorsal
pubescence and similar antennae.
Etymology. Named after the Roman name Gallia of the
country of amber origin.
Diagnosis. In contrast to the recent species of the genus as well as
to Cyphon lobanovi nov. sp., and, perhaps, the species described
from Baltic amber (Iablokoff -Khnzoryan 1961; Klausnitzer
2004), this new species has the mesoventrite sharply divided
into the sloping anterior part and posterior part, the latter being
in the same plane as metaventrite and abdominal ventrites and
somewhat elevated before the sloped part. Besides, this new
species is characterized by comparatively rather wide elytra,
very wide epipleura, and not expressed submesocoxal line.
Other particular features are the nearly straight anterior edge
of the pronotum, characteristic antennae, and comparatively
widely separated procoxae. is genus is known also by fossils
from the lacustrine deposits from Bembridge Marls of the Isle
Wight (Kirejtshuk et al. in press b), which, in contrast to this
new species, have a markedly more slender body and narrower
head.
Description. Holotype: body 3.4 mm long, 2.0 mm wide,
1.5 mm high; oval, rather convex dorsally and ventrally;
subunicolorous dark bronze with somewhat lighter appendages;
with a strong bronze shine; dorsum with well conspicuous,
suberect and comparatively long yellow greyish hairs about 2-3
times as long as distance between their insertions; underside
somewhat sparser, very fi ne, subrecumbent and shorter hairs
somewhat longer than distance between their insertions; elytral
sides fringed with more conspicuous hairs.
Figures 1–8
Cyphon gallicus nov. sp. (Scirtidae): 1, body, dorsally; 2, proximal part of
antenna, dorsally; 3, maxillary palpus, ventrally; 4, apex of labial palpus,
ventrally; 5, prosternal process and median part of mesoventrite, ventrally;
6, meso- and metacoxae, metepisternum and mesofemur, ventrally; 7,
metafemur, ventrally; 8, protibia and tarsus, dorsally. Scales: A, to fi g. 1
representing bar of 1.0 mm; B, to fi gs 2-5 representing bar of 0.5 mm; C, to
fi gs 6, 7 representing bar of 0.5; D, to fi g. 8 representing bar of 0.5 mm.
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422
A. G. K & A. N
Head and pronotum with distinct, sparse and small punctures,
somewhat smaller than eye facets in diameter, interspaces
between them about three puncture diameters and smoothly
alutaceous; besides, pronotum with a irregular cellular net of
lines over the puncturation and microsculpture. Elytra with
much larger and denser punctures (somewhat larger than eye
facets in diameter); interspaces between them about as large
as a puncture diameter and smoothly alutaceous; besides,
with an irregular cellular net of lines over the puncturation
and microsculpture which is coarser than that on pronotum.
Underside with punctures almost as large as eye facets;
interspaces between them on thoracic sclerites and metacoxae
subequal to or slightly greater than a puncture diameter and
more or less smoothed; those on ventrites much less than a
puncture diameter and with very dense, fi ne and pronounced
microreticulation.
Head oval and well exposed dorsally, not clearly visible because
of position of the beetle in amber, apparently somewhat longer
than wide, considerably narrower than pronotum, with rather
large eyes not reaching prothoracic segment, base somewhat
convex and frons subfl attened; labrum apparently quadrangular
and transverse (about 1.5 as wide as long); mandibles
invisible because of surrounding organic matter, apparently
comparatively small; antennae with missing terminal segments
(right one only with six antennomeres and left one with fi ve),
proximal six antennomeres combined about as long as pro- and
mesotibiae, very shortly pubescent; scape moderately thick and
subcylindrical, about twice as long as wide, about 2.5 times
as long as oval antennomere 2 and nearly twice as long as
subconical antennomere 3, antennomeres 4-6 subcylindrical
and slightly shorter than scape; mentum not visible; ultimate
maxillary palpomere subconical, rather narrowing apically,
about 3.5 times as long as wide at its thickest place and about
twice as long as ultimate labial palpomere; labial palpi three-
segmented, ultimate labial palpomere only slightly narrowing
apically and about twice as long as thick at base.
Pronotum with distinctly bisinuate anterior edge (very convex
in its middle) and rather convex posterior edge, all angles
rounded, widest at posterior angles, slightly and gradually
narrowing anteriorly, posterior and lateral edges not bordered;
disk rather convex and sides rather sloping; scutellum moderately
large and looking almost like equilateral triangle; prosternum
moderately convex in anterior part and its process in narrowest
place extremely narrow, far projecting together with coxae and
strongly curved and somewhat widened before abrupt apex,
looking like an isolated and widened plate; procoxae distinctly
transverse and far projecting below; mesoventrite evenly sloping
anteriorly from narrow transverse stripe before mesocoxae
and somewhat elevating before the sloped part; mesocoxae
transversely oblique, very narrowly but distinctly separated;
metaventrite rather short and medially convex, with paracoxal
line before metacoxae not interrupted in the middle and rather
convex; submesocoxal line not expressed; metepisterna exposed
along the whole length, rectilinearly widening anteriorly;
ventrites comparable in length, although ventrite 1 slightly
shorter, hypopygidium very widely rounded at posterior edge;
epipleura very distinct, at base about four times as wide as distal
plate of prosternal process and twice as wide as scape; pygidium
very widely rounded to widely subtruncate at apex and slightly
exposed from under elytral apices.
Elytra only slightly longer than wide combined, longest at
suture, gently arcuate at sides; their apices apparently forming
a joint arc, slightly convex to subfl attened at disk and rather
steeply sloping (but not subvertically) at sides, and sutural lines
not clearly traced at distance from suture as great as antennomere
2 thick and expressed along the whole length.
Legs well developed and quite narrow; metacoxae contiguous
and moderately oblique, with very distinct femoral plates at
median part; trochanters rather of elongate type; tibiae slightly
compressed, narrow, about as wide as fl agellomeres, almost
comparable in length (although posterior tibia somewhat
longer) and subparallel-sided; their apices truncate and with
one clear spur, outer edges with setiferous border; femora of
usual shape, pro- and mesofemora about four times as wide as
tibiae and metafemur somewhat wider. Tarsi fi ve-segmented and
comparatively short, tarsomere 1 subconical (on anterior and
intermediate legs slightly longer, on posterior one apparently
about twice longer than following ones), tarsomeres 2-4 widely
bilobed, tarsomere 5 more or less subequal in length to pro-
and mesotarsomere 1; claws simple narrow and short.
Variation. Paratype: Body 2.9, mm long, 1.7 mm wide; more
slender than the holotype, rather convex dorsally and with
abdominal apex more exposed from under the elytral apices;
basal antennomeres looking like those of the holotype; the
Figures 9–16
Cyphon lobanovi nov. sp. (Scirtidae): 9, body, dorsally; 10, thorax, ventrally;
11, antenna, dorsally; 12, apex of maxillary palpus, ventrally; 13, mentum,
ligula and labial palpus, ventrally; 14, prosternal process with procoxa,
laterally; 15, abdominal apex, dorsally; 16, metatarsus, dorsally. Scales: A,
to fi g. 9 representing bar of 1.0 mm; B, to fi gs. 10, 11, 14–16 representing
bar of 0.5 mm.
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Polyphaga from Eocene French amber
423
antennomeres 7-10 more or less subcylindrical, about as large
and as shaped as previous ones and antennomere 11 slightly
longer and gently narrowing at subrounded apex.
Cyphon lobanovi nov. sp.
(Figs 9–16, 71)
Material. Holotype PA 5605, probably male [beetle with
missing left posterior leg included into an amber bar (about 8.0
mm long, 2.0 mm wide, and 2.0 mm high) containing many
gas vesicles, some cracks along beetle body, and an elongate
hole in amber on the right side from the beetle of the same size
with remains of chitin, a mould of insect mummifi ed together
with the beetle].
Etymology. is species is named after A. L. Lobanov, colleague
of the senior author and great enthusiast in creation of the web-
site <http://www.zin.ru/Animalia/Coleoptera>.
Diagnosis. is new species that we tentatively attribute to
the genus Cyphon, is distinct from the recent and Baltic amber
species (Iablokoff -Khnzoryan 1961; Klausnitzer 2004) in the
very sparse puncturation of the dorsum with rather smoothed
interspaces and much reduced pubescence. Besides, this new
species is rather slender in contrast to most members of the
genus. Its parametacoxal line is nearly transverse before the
anterior edge of coxae. See also above the diagnosis of the
previous species. is species has the male genital structures
quite diff erent from those in the species described from Baltic
amber. It is distinct from the species known from Bembridge
Marls (Kirejtshuk et al. in press b) in the much more slender
body and very long elytra.
Description. Body 2.1 mm long, 0.7 mm wide, 0.7 mm
high; elongate oval, strongly convex dorsally and ventrally;
subunicolorous dark bronze brown with lighter appendages; with
a strong bronze shine; dorsum with extremely fi ne, moderately
long, subrecumbent and scarcely visible hairs; underside yet
fi ner hairs; elytral sides fringed with more conspicuous hairs.
Head and pronotum with distinct, dense and very small
punctures, markedly smaller than eye facets in diameter,
interspaces between them more or less greater than a puncture
diameter and smoothly alutaceous; elytra with larger and
sparser punctures (about as large as eye facets in diameter or
somewhat larger); interspaces between them about as great
as a puncture diameter and smoothly alutaceous to smoothly
microreticulated; meso- and metaventrites as well as metacoxae
with extremely small and sparse and very smoothed punctures;
abdominal ventrites with shallow and almost distinct punctures,
slightly smaller than eye facets, separated markedly less than a
puncture diameter and with smoothed microreticulation.
Head oval and well exposed dorsally, apparently not longer
than wide, scarcely narrower than pronotum, with rather large
eyes not reaching prothoracic segment, base somewhat convex
and frons subfl attened; labrum invisible because of position
of the beetle in amber, although its truncate anterior edge is
visible between mandibular apices; mandibles well developed,
rather thin, with simple and projecting apices and far extending
beyond frons; antennae 11-segmented, comparatively long,
reaching the anterior edge of metacoxae, very shortly pubescent
and submoniliform, scape moderately thick and subcylindrical,
about 1.5 times as long as wide, about twice as long as each
of antennomeres 2 and 3 as well as somewhat longer than
ultimate antennomere; antennomere 2 shortest and almost
as thick as scape; antennomere 3 narrowest and slightly
longer than antennomere 2; antennomeres 4-9 about as long
as antenomere 2 and gradually becoming thicker and more
moniliform; antennomeres 10 and 11 longest in fl agellum and
subcylindrical; ultimate antennomere forming a narrowly blunt
apex; mentum subquadrangular, distinctly widened anteriorly;
ultimate maxillary palpomere conical, rather narrowing apically,
about 2.5 times as long as wide at its thickest part and about
twice as long as ultimate labial palpomere; labial palpi three-
segmented, ultimate labial palpomere narrowing apically and
about twice as long as thick at base.
Pronotum with nearly straight anterior edge and rather convex
posterior edge, all angles rounded, widest at posterior angles,
very slightly and gradually narrowing anteriorly, posterior
and lateral edges very distinctly bordered; disk rather convex
and sides rather sloping; scutellum moderately large, slightly
transverse and subtriangular; prosternum anteriorly looking like
an extremely narrow stripe and its process in narrowest place
markedly narrower than narrowest antennomere, far projecting
together with coxae and vertically abrupt, as its apex looking like
an isolated and somewhat widened plate; mesoventrite evenly
sloping from posterior edge anteriorly; metaventrite rather
short and medially convex, with straight parametacoxal line not
interrupted in the middle; submesocoxal line reaching anterior
angle of metaventrite and ending at inner anterior angle of
metepisternum; metepisterna exposed along the whole length,
arcuately widening anteriorly. Abdominal ventrites comparable
in length, although distal ones slightly shorter, hypopygidium
very widely rounded at posterior edge; epipleura very distinct
and comparatively narrow, at base about twice wider than
prosternal process and about as wide as scape.
Elytra about 1.5 time as long as wide combined, longest at
suture, subparallel-sided in proximal 2/3, distally gently and
arcuately narrowing; their apices forming a joint arc, moderately
convex at disk and steeply sloping (subvertically) at sides, and
sutural lines not expressed.
Pygidium and hypopygidium widely rounded at apex and
slightly exposed from under elytral apices; besides, a moderately
sclerotized and slightly narrowing anteriorly ‘strut’ protruding
beyond the apices of these sclerites.
Legs well developed and quite narrow; procoxae distinctly
transverse and far projecting below; mesocoxae transversely
oblique, very narrowly but distinctly separated; metacoxae
contiguous and rather oblique, with very distinct femoral
plates at median part; trochanters rather of elongate type; tibiae
slightly compressed, narrow, somewhat wider than fl agellomeres,
almost comparable in length and subparallel-sided; their apices
truncate and with one clear spur, outer edges with setiferous
border; femora of usual shape, pro- and mesofemora about
three times as wide as tibiae and metafemur somewhat wider;
tarsi fi ve-segmented and comparatively short, tarsomere 1
subconical (on anterior and intermediate legs slightly longer,
on posterior one about twice longer than following ones),
tarsomeres 2-4 widely bilobed, tarsomere 5 more or less shoter
than tarsomere 1; claws simple narrow and short.
Note. e ‘strut’ protruding beyond the apices of
the last abdominal segment shows some resemblance
with the penis trunk of the recent Cyphon furcillatus
Nyholm 1948.
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424
A. G. K & A. N
Infraorder Cucujiformia Lameere 1938
Superfamily Cleroidea Latreille 1802
Family Melyridae Leach 1815
is family is scarce in fossils, although it is
rather frequent in Baltic amber (Spahr 1981; Hieke
& Pietrzeniuk 1984). It is also found in the Lower
Oligocene Florissant (Wickham 1912, 1917 etc.) and
the Oligocene French Massif Central ( éobald 1937).
Recently this family was recovered from Bembridge
Marls at the boundary between the Oligocene and
Eocene (Kirejtshuk et al. in press b). us, our new
species are the oldest described melyrids. Besides, this
family was mentioned in the list of inclusions found in
the Burmese amber deposited in the Natural History
Museum in London (Rasnitzyn & Ross 2000), and
one probable representative was determined in the
Lebanese amber (Kirejtshuk & Azar in press). e
recent range of this family includes all zoogeographical
regions, although its maximum diversity corresponds
to warm areas, particularly in the Old Mediterranean,
including the Middle and Central Asia. Adults are
phytophagous, live on vegetation and usually on
fl owers, although sometimes they can also feed on
other insects. Larvae are primarily predaceous.
Subfamily Malachiinae Fleming 1821
is subfamily is known in fossils mostly by some
genera from the Upper Eocene Baltic amber: Anthocomus
Erichson 1840; Attalus Erichson 1840; Apalochrus
Erichson 1840; Collops Erichson 1840; Colotes Erichson
1840; Ebaeus Erichson 1840; Malachius Fabricius
1775; Macrocerus Motschulsky 1845 non Oken 1817
(= Malchinus auctorum non Kiesenwetter in Erichson
1863) (Berendt 1845; Motschulsky 1956; Klebs 1910;
Iablokoff -Khnzorian 1960; Larsson 1978; Kubisz 2001
etc.). e genus Malachius is recorded from the Lower
Oligocene Florissant Shales (Wickham 1916, 1917),
and the genus Troglops Erichson 1840 is recorded from
the Oligocene French Massif Central ( éobald 1937).
Finally, a formal genus apparently similar to Colotes
was recently established in the Bembridge Marls at the
boundary between Oligocene and Eocene (Kirejtshuk
et al. in press b).
Genus Colotes Erichson 1840
Type species. Malachius maculatus Laporte 1838, recent.
Remark. All specimens examined in the Oise amber are
or look like females and seem to belong to one group of
relatives, which is here tentatively put in Colotes because
of many characters more or less similar, including the
type of body coloration and integument sculpture.
Recent species (about 100 species) are characterized
by a sexual dimorphism in the maxillary palpi, but
with a rather great variability of this structure in both
sexes within the diff erent subgroups of this genus. e
antennae of the recent species are usually rather long
and tend to be simple, although antennomere 2 can be
as small as that in the fossil specimens here considered.
e tarsi of the recent species are comparatively longer
than those in the fossil specimens from the Oise amber.
is genus was listed from Baltic amber (Klebs 1910;
Larsson 1978 etc.), with one described species Colotes
sambicus Kubisz, 2001.
On the other hand, the fossil specimens under
consideration are rather similar to the recent species
of the genus Attalus Erichson, 1840, subgenus Abrinus
Mulsant, 1867, but the studied specimens diff er from
those of the latter subgenus in the more robust body
with larger eyes, ultimate maxillary palpomere not
Figures 17–23
Colotes constantini nov. sp. (Melyridae): 17, body, dorsally; 18, idem,
laterally; 19, head, anterodorsally; 20, antenna, dorsally, 21, abdominal
apex, ventrally; 22, metatarsus, dorsally; 23, idem, laterally. Scales: A, to
fi g. 17, 18 representing bar of 1.0 mm; B, to fi gs 19, 21 representing bar of
1.0 mm; C, to fi gs 20, 22, 23 representing bar of 1.0 mm.
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Polyphaga from Eocene French amber
425
narrowed apically, more regularly oval head, clearly
shorter antennae, and shorter tarsi. e recent fauna
of Attalus includes at least 400 species divided into
some subgenera, some of them have comparatively
short antennae similar to that in the new fossil species
and very short antennomere 2 (Wittmer 1988; etc.).
Nevertheless the new fossil species diff er from the
recent species of this genus in the maxillary palpi with
comparatively wide apex in the females, and in the very
large eyes. ey also diff er in the rather small body size,
comparatively short and subpectinate antennae. ey
could belong to a separate genus. But this problem
of taxonomic separation should be solved after a
futher study of male specimens from the Oise amber.
Besides, our fossil species are clearly distinct from
Colotes sambicus from the Baltic amber in the smaller
body size, shorter antennae and tarsi, less developed
puncturation, and alutaceous sculpture of dorsum.
e recent species of Colotes have a rather strong
sexual dimorphism in the structure of their maxillary
palpi. All fossil specimens here considered have no
clear trace of such a dimorphism. Both type specimens
of C. impexus nov. sp. are females (ovipositor). Only
one paratype of C. constantini nov. sp. is clearly a
female with a visible ovipositor, while the sex of other
specimens is undetermined, but they all show a stable
structure of the last maxillary palpomere. It can be also
suggested that these Eocene fossils could still have the
sexual dimorphism not so pronounced as in the recent
Colotes.
Both series are here regarded as separate species
because they show structures which are usually
characteristic of species separation.
Colotes constantini nov. sp.
(Figs 17–23, 72–74)
Material. Holotype PA 9530, female(?) [a complete beetle
with partly exposed hind wings included into an amber bar
containing many small cracks, pieces of organic matter and
very small gas vesicles, two Nematocera, one Hymenoptera,
and one ysanoptera. e consistence of the amber substrate
is not homogenous, with some layers after concentric rings].
Paratypes: PA 2764 3/6, female(?) [complete beetle, the
consistence of the amber substrate is more or less homogenous,
with two Dermestidae and one Hymenoptera]; PA 1781
10/15, female [complete beetle included into an elongate
amber bar with irregular longitudinal planes partly unpolished,
together with paratype of Oisegenius antiquus nov. gen., nov.
sp. (‘11/15’), one small Hymenoptera (‘11/16’), paratype of
C. constantini nov. sp., small gas vesicles and small pieces of
organic matter; the consistence of the amber substrate is more
or less homogenous]; 11959, female(?) [a complete beetle in a
more or less homogenous large amber piece].
Etymology. is new species that we tentatively attribute to the
genus Colotes, is named after R. Constantin, the friend of one of
authors and specialist on family Melyridae, who contributed to
the identifi cation of this family in amber inclusions.
Diagnosis. is new species diff ers from C. impexus nov. sp.
in the body coloration, particularly in the light transverse
stripes on elytra, narrower prothoracic segment and head,
shorter anterior part of frons, somewhat larger eyes, shape of
ultimate maxillary palpomere, not so compressed fl agellomeres,
shape of pronotum, small eversible vesicles in both pro- and
metathoraces, clearly oblique elytral apices, somewhat longer
legs, fi ner and sparser puncturation on elytra, and bigger tooth
of tarsal claw.
Description. Holotype: Body 2.4 mm long, with declined
prothoracic segment and head 1.9 mm long, 0.7 mm wide,
0.6 mm high; elongate oval, moderately convex dorsally and
ventrally, with an eversible vesicle at anterior pronotal angles
moderately small and an eversible vesicle at distal angle of
metathorax not extending along abdomen; head, metathorax,
basal abdominal ventrites, base and distal parts of elytra (but
not stripes along apices) and metatibiae rather dark to blackish,
abdominal ventrites and median part of metafemora also dark
(brownish); remainder lighter, almost straw reddish, including
appendages of head, prothorac segment, transverse stripes
through middle and apices of elytra and legs; with a some
bronze shine; dorsum with well conspicuous, recumbent and
short greyish hairs about two times as long as distance between
their insertions; underside with sparse, very fi ne, subrecumbent
and short hairs with distance between them less than length of
hairs.
Head, pronotum and elytral base with rather small, sparse
and shallow punctures, about 1/3 as large as eye facets in
diameter; interspaces between them about three times as great
as a puncture diameter and smoothly alutaceous; elytra apically
becoming impunctured to unclearly covered with small and
rather shallow punctures; underside and tergites uncovered by
elytra with small and sparse unclear punctures, much smaller
than eye facets in diameter; interspaces between them alutaceous
to smoothly microreticulated.
Head oval and apparently not wider than long, slightly convex
dorsally, somewhat narrower than pronotum, with rather
large and oval eyes far not reaching prothoracic segment and
about half as long as head; clypeus looking like a wide isolated
stripe; labrum apparently quadrangular and transverse, about
three times as wide as long; mandibles small and invisible
because of milky foggy around anterior part of head; antennae
subserrate and 11-segmented (both well preserved), scape
rather large and somewhat curved; antennomere 2 very small
and subcylindrical; antennomeres of fl agella subtriangular and
dorsoventrally compressed, about 2.5 as long as scape, although
ultimate antennomere about half as long as scape; mentum
invisible; penultimate maxillary palpomere narrow and twice
as long as wide at pex; ultimate maxillary palpomere slightly
widened apically and with obliquely truncate apex, rather
widening apically, about twice as long as wide at its thickest
place at apex and slightly longer than penultimate ones; labial
palpi invisible.
Pronotum suboval, with convex anterior and posterior edges
and arcuate lateral edges, all angles not expressed, gradually
narrowing anteriorly; disk rather convex and sides rather
steeply sloping; scutellum rather large, looking almost like
an elongate triangle and rounded at apex; prosternum not
visible; mesoventrite invisible clearly; metaventrite rather long
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426
A. G. K & A. N
and strongly convex in the middle, with paracoxal line before
metacoxae; metepisterna exposed along the whole length and
rather wide.
Elytra about 1 and 7/11 as long as wide combined, longest at
suture and widest before apices, their sides subdivergent distally
and apices suboblique, slightly convex to subfl attened at disk
and rather steeply sloping (subvertically) at sides and at base
turning ventrally
Abdominal ventrite 1 longest; rest ventrites comparable in length;
four distal abdominal tergites uncovered; distal abdominal
segments gradually narrowing apically; hypopygidium very
widely truncate to subemarginate at posterior edge; pygidium
very widely rounded at apex.
Legs very long and very narrow; procoxae moderately projecting
below; mesocoxae transversely oval, very narrowly separated;
metacoxae apparently narrowly separated and strongly oblique;
trochanters rather of elongate type; tibiae scarcely compressed,
narrow, slightly wider than fl agellomeres; pro- and mesotibiae
about as long as antenna, almost comparable in length, nearly
straight and subparallel-sided; metatibia nearly 1.5 times as long
as antenna and somewhat curved and narrowing at apex; their
apices truncate and without clear spurs, outer edges unbordered;
femora of usual shape, pro- and mesofemora about three times
as thick as tibiae and about as long as antenna; metafemur
slightly wider and about 1 and 1/3 as long as antenna; tarsi fi ve-
segmented and comparatively short, tarsomere 1 subconical,
tarsomeres 2-4 widely bilobed, tarsomere 5 more or less
subequal in length to tarsomeres 1-3; claws long and toothed
at the middle, simple, narrow and slightly curved, ungular
appendages rather long and almost reaching the apex of claw.
Variations. Paratype PA 2764: body 2.5 mm long, 0.7 mm
wide, 0.6 mm high, with declined prothoracic segment and
head 2.0 mm long; body elongate oval, moderately convex
dorsally and ventrally, with an eversible vesicle at anterior
pronotal angles moderately small and an eversible vesicle along
metathorax and slightly extending along abdomen; light reddish
to slightly brownish, although pterothorax, base and distal parts
of elytra (but not stripes along apices) darkened (brown); with
some bronze shining; dorsum with well conspicuous, not quite
recumbent and short greyish hairs about two times as long as
distance between their insertions; underside sparse, very fi ne,
subrecumbent and short hairs with distance between them
less than length of hairs. Abdomen with somewhat exposed
ovipositor. Paratype PA 1781: body 2.3 mm long, 0.6 mm wide,
with declined prothoracic segment and head 1.8 mm long;
body straw reddish with somewhat darkened elytral base and
distal parts of elytra; maxillary palpi of both paratypes identical
to those of holotype. Paratype PA 2764: body 2.5 mm long,
0.7 mm wide, 0.6 mm high, with declined prothoracic segment
and head 2.0 mm long; body elongate oval, moderately convex
dorsally and ventrally, with an eversible vesicle at anterior
pronotal angles moderately small and an eversible vesicle along
metathorax and slightly extending along abdomen; light reddish
to slightly brownish, although pterothorax, base and distal parts
of elytra (but not stripes along apices) darkened (brown); with
a some bronze shine; dorsum with well conspicuous, not quite
recumbent and short greyish hairs about two times as long as
distance between their insertions; underside sparse, very fi ne,
subrecumbent and short hairs with distance between them
less than length of hairs. Abdomen is with somewhat exposed
ovipositor.
Colotes impexus nov. sp.
(Figs 24–27, 75–77)
Material. Holotype PA 890, female [complete beetle obliquely
passing through a crevice included into an amber bar, with many
some small cracks, organic matter, and very small gas vesicles
spread throughout of amber].
Paratype PA 9094, female [beetle lying closely to amber surface
with left side of body opened].
Etymology. e epithet of this new species means ‘coarse’.
Diagnosis. See the diagnosis of the previous new species.
Description. Holotype: body 2.2 mm long, 0.8 mm wide, 0.6
mm high, with declined prothoracic segment and head 1.6
mm long; body elongate oval, moderately convex dorsally and
ventrally, with a very small eversible vesicle at anterior pronotal
angles and a large one along distal half of metathorax and basal
abdominal segments; metathorax, basal abdominal ventrites and
elytra rather dark to blackish, abdominal ventrites and median
part of metafemora also dark (brownish); remainder lighter,
almost straw reddish; very slightly shining with a clear bronze
luster on pronotum and head; body and appendages with well
conspicuous, recumbent and short greyish hairs somewhat
longer than distance between their insertions.
Head and pronotum with very small and shallow punctures,
interspaces between them smoothly alutaceous; elytra with
more or less distinct and shallow punctures about 1/2 as large
as eye facets in diameter; interspaces between them a puncture
diameter and smoothly alutaceous; integument of underside
Figures 24-27
Colotes impexus nov. sp. (Melyridae): 24, body, dorsally; 25, idem, laterally;
26, apical antennomeres, dorsally, 27, metatarsomere 5 with claws, laterally.
Scales: A, to fi gs 24, 25 representing bar of 0.5 mm; B, to fi gs 25, 27
representing bar of 0.25 mm.
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Polyphaga from Eocene French amber
427
apparently somewhat similar to that on head and pronotum,
although at least on ventrites punctures somewhat larger and
interspaces between them smoothly microreticulated rather
than alutaceous.
Head oval and apparently not wider than long, slightly convex
dorsally, somewhat wider than pronotum, with rather large and
oval eyes far not reaching prothoracic segment and about 2/5 as
long as head; clypeus as a wide isolated stripe; labrum not clearly
visible, but apparently well exposed, transverse and somewhat
convex anterior edge; mandibles rather long and projecting;
antennae 11-segmented, scape moderately large and moderately
curved; antennomere 2 rather short; fl agellar antennomeres
distinctly dorsoventrally compressed, somewhat less than 1/3
as long as scape, although ultimate antennomere nearly half
as long as scape; mentum not visible; penultimate maxillary
palpomere narrow and almost twice as long as wide at apex;
ultimate maxillary palpomere slightly widened apically and with
obliquely truncate apex, rather widening apically, slightly longer
wide at its thickest place at apex and markedly shorter than
penultimate ones; labial palpi not visible.
Pronotum suboval, with convex anterior and posterior edges and
gently arcuate lateral edges, all angles not expressed, disk rather
convex and sides rather steeply sloping; scutellum rather large,
looking almost like a slightly elongate triangle and rounded at
apex; prosternum and mesoventrite not visible; metaventrite
rather long and strongly convex in the middle, with paracoxal
line before metacoxae; metepisterna exposed along the whole
length and rather wide.
Elytra about 1 and 5/11 as long as wide combined, apparently
longest at suture and widest before apices, their sides subdivergent
distally and apices suboblique to subtransverse, slightly convex
to subfl attened at disk and rather steeply sloping (subvertically)
at sides and at base turning ventrally.
Abdominal ventrite 1 longest; ventrites 2-5 comparable in
length, hypopygidium markedly shorter than ventrite 1 and
much longer than ventrites 2-5, very widely rounded at posterior
edge; remaining four distal tergites uncovered; distal abdominal
segments gradually narrowing apically; pygidium somewhat
projecting and comparatively narrowly subtruncate at apex.
Legs moderately long and very narrow; procoxae moderately
projecting below; mesocoxae transversely oval, apparently very
narrowly separated; metacoxae apparently narrowly separated
and strongly oblique; trochanters rather of elongate type;
tibiae slightly compressed, narrow, 3/4 as wide as fl agellomeres;
pro- and mesotibiae somewhat shorter than antenna, almost
comparable in length, nearly straight and very slightly widened
apically; metatibia somewhat longer than antenna and somewhat
curved; their apices truncate and without clear spurs, outer
edges unbordered; femora of usual shape, pro- and mesofemora
about three times as thick as tibiae and markedly shorter than
antenna; metafemur slightly wider and about as long as antenna;
tarsi fi ve-segmented and comparatively short, tarsomere 1
subconical, tarsomeres 2-4 widely bilobed, tarsomere 5 more or
less subequal in length to tarsomeres 1-3; claws slightly toothed
at base, narrow and slightly curved, ungular appendages rather
long and almost reaching apex of claw.
Variation. Paratype: Body 2.0 mm long, 0.6 mm high, with
declined prothoracic segment and head 1.5 mm long; body
elongate oval, moderately convex dorsally and ventrally;
subunicolorous straw reddish with slightly darker elytra.
Superfamily Cucujoidea Latreille 1802
Family Nitidulidae Latreille 1802
e sap beetles is a rather numerous and diverse
family in the recent fauna, although only few fossils
have been described. Recent Nitidulidae have rather
diverse bionomies. ey spread in many diff erent
environments, but this family is not present in water
basins and in very wet substrates. It is also quite rare in
arid areas. Sap beetles very seldom inhabit in soil. Most
representatives are mycetophagous but many groups are
associated with fl owers. Few species are phyllophagous
or predaceous. Some groups live on colonies of scale
insects and whitefl ies (Cychramptodini Kirejtshuk &
Lawrence 1992 and Cybocephalinae). Highest diversity
of sap beetles is recorded in forest communities. e
Nitidulidae fi rst appear in the fossil record from the
Lower Cretaceous, with all earlier records erroneous
(Kirejtshuk & Ponomarenko 1990). e carpophilin
lineage of the family was recorded somewhat earlier
(Baissa, Middle Neocomian) than the nitidulin lineages
(Obeshtshayushtshiy Creek, Cenomanian). During
the Cenozoic this family is mostly known from Baltic
Amber (listed without description by Klebs 1910;
Larsson 1978; Spahr 1981; Hieke & Pietrzeniuk 1984)
and Dominican amber (Kirejtshuk & Poinar 2007),
where all subfamilies of Nitidulidae have been found,
except Amphicrossinae Kirejtshuk, 1986 and Cillaeinae
Kirejtshuk & Audisio, 1986 (in Kirejtshuk 1986). e
latter was recorded in the Lower Oligocene Florissant
shales (Wickham 1913) together with other sap beetles
from Carpophilinae Erichson 1842 and Nitidulinae.
Besides, Nitidulinae are known from the Middle
Miocene Wishnevaya Balka beds (North Caucasus,
Karaganian: Kirejtshuk & Ponomarenko 1990), the
Lower Oligocene Kleinkembs (Germany) ( éobald
1937), the Lower Miocene Radoboj (Croatia) (Heer
1847). e Cybocephalinae are recorded from the
Baltic amber (Hieke and Pietrzeniuk 1984), and from
the Middle Miocene Mojave Desert deposits (USA)
(Cybocephalus cf. californicus Horn 1879) (Palmer et al.
1957). us, the species here described are the oldest
fossil members of the subfamily Cybocephalinae.
Subfamily Cybocephalinae Jaquelin du Val 1858
Genus Pastillocenicus nov. gen.
Type species. Pastillocenicus grandiclavis nov. sp., other species: P. polyaki
nov. sp. and P. longifrons nov. sp.
Etymology. Named after the generic name ‘Pastillodes’ and
‘cenicus’, masculine gender.
Diagnosis. is genus diff ers from all other groups in
Cybocephalinae in the combination of the comparably short
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428
A. G. K & A. N
metaventrite, meso- and metacoxae more or less rather widely
separated intermesocoxal line convex, and particularly lack
of submetacoxal lines. e very important and characteristic
features of this genus are the nearly four-segmented and very
long antennal club (such long antennal club is unknown
among the recent representatives of the subfamily), and also the
protibia strongly widened along its inner edge. Finally, the eyes
contour of Pastillocenicus polyaki nov. gen., nov. sp. is more or
less even, only slightly interrupted by a fold of the frons along
their anterior edge. However, the posterior part of the eyes of
this new species is somewhat concealed into the prothoracic
segment.
e lack of the submetacoxal lines and very long
antennal club in the type species of this new genus are
its most distinct characters, allowing distinguishing
it among other representatives of the subfamily. e
type species of this new taxon diff ers from members
of Cybocephalus in the entire set of the characters
mentioned above, although metacoxae of the latter are
usually widely separated (but the distance between these
coxae are usually markedly less than the width of the
coxae). At the same time, the species of Endroediellus
Endrödy-Younga 1962, in addition to general
characters mentioned above, diff er from Pastillocenicus
polyaki in the very short and wide metafemora as
well as in the shorter frons and labrum, not exposed
from its ventral part. Pastillocenicus polyaki can be
distinguished from Hierronimus Endrödy-Younga
1968 and Pastillodes Endrödy-Younga 1968 in the eyes
occupying both dorsal and ventral head surface, not so
short metaventrite, and much more widely separated
metacoxae.
Species of the genus Pastillus Endrödy-Younga
1962, in contrast to Pastillocenicus polyaki, have a much
shorter metaventrite and equally narrowly separated
meso- and metacoxae (not widely separated at all).
Pastillocenicus polyaki can be distinguished from species
of Pycnocephalus Sharp 1891 in the smaller body size,
eyes occupying both dorsal and ventral head surface,
shorter metaventrite, much more widely separated
metacoxae, narrow protibia without crenellation along
outer edge, and longer and narrower metafemora and
metatibae. Pastillocenicus polyaki can be distinguished
from Taxicephomerus Kirejtshuk 1994 in the longer
frons, shorter and transverse mentum, protibia without
crenellation and projecting outer apical angle, lobed
tarsomeres 1-3, and more widely separated metacoxae
(in Taxicephomerus the tarsomeres are wider than
in Pastillocenicus polyaki but scarcely lobed and with
short setae, while its meso- and metacoxae are equally
separated, but distance between coxae in each pairs
much less than width of metacoxae). Pastillocenicus
polyaki can be distinguished from Horadion Endrödy-
Younga 1976 in the much longer and not transverse
head, 11-segmented antennae with four-segmented
antennal club, and much narrower protibia. e
large eyes of Pastillocenicus longifrons nov. sp. share
some resemblance with some species of the subgenus
eticephalus Kirejtshuk 1988, including species close
to C. (T.) decamerus Endrödy-Younga 1968, but the
subacute elytral apices of the new species from amber are
rather diff erent from those in species of eticephalus,
which are separately and widely rounded.
Note. All specimens examined show no sexual
dimorphism in the structure of their abdomen, rather
characteristic among species of the Recent and Baltic
amber faunas. Males of this new fossil genus could
have the anal sclerite not exposed from under apex of
pygidium.
Description. Body rather small, oval, dorsally very convex and
slightly convex ventrally; integument more or less smoothed and
fi nely punctured; dorsum very fi nely and slightly conspicuously
pubescent; underside with fi ne and dense hairs; posterior
edge of pygidium and hypopygidium along posterior edge of
penultimate ventrite with a regular row of more or less long
and stout setae.
Head rather wide, subtriangular, dorsally weakly convex and
with a very short and wide frons, rather large eyes more or less
extended on underside, although along anterior edge of eye a
narrow fold of continuing frons; labrum exposed from under
frons, unilobed; mandibles with bidentate apices and more or
less exposed from under frons; antennae 11-segmented and
with three-segmented elongate and comparatively compact
club; mentum subquadrangular and transverse; ultimate
maxillary palpomere subcylindrical to slightly narrowing
apically; ultimate labial palpomere comparatively wide and
short; antennal grooves expressed; gular sutures subparallel-
sided.
Pronotum large, subsemicircular and widest at base, rather
and evenly convex, with steeply sloping sides, its anterior edge
Figures 28–33
Pastillocenicus longifrons nov. gen., nov. sp. (Nitidulidae): 28, body,
dorsally; 29, body, laterally; 30, head, anteroventrally; 31, apex of antenna,
dorsally; 32, metacoxa and precoxal depression, ventrally; 33, posterior leg,
ventrally. Scales: A, to fi gs 28, 29 representing bar of 0.5 mm; B, to fi g. 30
representing bar of 0.5 mm; C, fi gs 32, 33 representing bar of 0.5 mm.
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Polyphaga from Eocene French amber
429
slightly emarginated or excised; scutellum subtriangular and
rather large; prosternum very short and with narrow process,
curved along coxae and not extending the level of posterior edge
of procoxae; distance between mesocoxae somewhat greater
and that between metacoxae yet more greater than that between
procoxae; mesoventrite excavate and medially subcarinate at
bottom; metaventrite rather short and with posterior edge
between coxae apparently shallowly emarginate; abdominal
ventrite 1 longer than hypopygidium and each ventrites 2-
4 shorter than both ventrite 1 and hypopygidium; epipleura
moderately narrow and sloping downwards laterally.
Elytra somewhat shorter than their combined width, rather
convex at disk, evenly and gently sloping laterally, with their
sides more or less arcuate, longest at suture. Pygidium widely
rounded at apex.
Tibiae rather widened along inner edge and apparently
comparable in width at apex, wider antennal club; outer edge
of meso- and metatibiae with a row of long setae; femora of
usual shape and with gently convex anterior and posterior
edges; metafemur wider and more enlarged apically; tarsi
comparatively narrow, tarsomeres 1-3 narrowly lobed, claw
moderately long and narrow.
Pastillocenicus longifrons nov. sp.
(Figs 28–33, 78–79)
Material. Holotype PA 5132, female(?) [complete beetle with
strongly defl ected head and almost clear integument included
in the small elongate amber piece].
Paratype PA 2451, female(?) [complete beetle with clear
integument, somewhat defl ected head (broken at frons)
and slightly exposed posterior wing in the very small amber
piece in Canada balsam on the microscope slide and covered
by the round cover glass. e consistence of amber is more
or less homogenous, the piece has a crack passing through
the beetle and some pieces of organic matter and many very
small gas bubbles.]. Paratype PA 2545, female(?) [complete
beetle somewhat damaged at suture of in anterior half of left
elytron, at both eyes and in the middle of metaventrite, and
also with defl ected head and integument partly with very thin
milky bloom along body sides in the very small amber piece
in Canada balsam on the microscope slide and covered by the
standard cover glass. e consistence of amber is heterogenous,
the boundaries of diff erent layers passing in perpendicular
directions along both body sides of the beetle and some
heterogeny goes along body sides in the plane of the beetle].
Additional specimen PA 1032, female(?) [complete beetle with
somewhat defl ected head and integument fi nely covered with
very thin milky bloom, although elytra and legs have nearly
clear surface, in a small and elongate amber piece, with some
very small vesicles spread through this piece and one depressed
vesicle ventrally and behind from abdomen of the beetle. Apex
of right wing is exposed from under elytral apex and declined
on ventral body side].
Note. e specimen PA 1032 looks like more slender
and more convex than other examined and recognized
as conspecifi c (holotype and paratypes of P longifrons
nov. sp.) and with some other diff erences (see below),
but the characters that can be observed in it are mostly
the same as in the females. Unfortunately, puncturation
and sculpture of the integument of the male are unclear
and cannot be used for comparison with the available
females. erefore this fossil is here regarded as an
additional specimen, although futher specimens from
this amber will probably clarify a distinctness of this
specimen from those of the type series of P. longifrons
nov. sp.
Etymology. Epithet formed from the Latin ‘longus’ (long) and
‘frons’ (frons, forehead).
Diagnosis. Pastillocenicus longifrons nov. sp. diff ers from the
other species of the genus in the more slender and subovoid
body with longer abdomen; more pubescent and not so shining
dorsum; head with much longer frons and shorter labrum
exposed from under frons; larger eyes; pronotum longer, more
arcuate at sides and bi-emarginate but not concave at base;
somewhat more distinct posterior angles of pronotum; longer
elytra; less widely separated metacoxae; comparatively longer
metacoxae, and somewhat narrower femora and tibiae.
Description. Holotype: Body 1.0 mm long (with defl ected
head), 0.5 mm wide, 0.3 mm high; elongate oval, dark chestnut
brown to blackish with brown appendages; rather convex
dorsally and slightly convex ventrally; dorsum with a slight
shine and underside nearly mat; dorsum with rather sparse and
well conspicuous hairs, about as long as distance between their
insertions; underside with similar but denser hairs; a regular
row of rather long and more stout setae along posterior edge
of pygidium (about twice as long as those on the remainder of
integument).
Head with some trace of weak, sparse and very fi ne irregular
rugosity, nearly alutaceous, and without punctuation;
pronotum with very sparse and small punctures bearing roots
of hairs, interspaces between them apparently smooth or
very fi nely alutaceous; elytra with punctures similar to those
on pronotum, but in addition with some smaller punctures
between them and interspaces fi nely alutaceous; pygidium
with sparse and very small punctures, interspaces between
them rather smoothed; underside seemingly fi nely and densely
sculptured (not smoothed), ventrite 1 with milky cover on the
median part; sides of metaventrite with smoothed and slightly
depressed lateral area at each side before coxae allowing femur
to move; this area distinctly bordered by a curved line, returning
to anterior edge of coxa and composing about a third of total
length of metaventrite.
Head weakly convex, with a frons rather projecting anteriorly,
almost twice as long as distance between very large eyes
(composed of moderately large facets); eyes seemingly extended
on underside; labrum slightly exposed from under frons,
unilobed and with a rather convex anterior edge; mandibles not
exposed from under frons; antennae completely not visible, but
the apex of their club can be seen from right side (apparently
antennomere 10 slightly wider than antennomere 11, the latter
somewhat longer than wide and transversely abrupt at apex).
Pronotum somewhat more than 2/5 as long as elytra, rather
and evenly convex, with steeply sloping sides, its anterior
edge bisinuate, lateral edges very slightly rounded, posterior
edge with a shallow emargination at each side of scutellum,
both anterior and posterior angles with somewhat rounded
apices (but so widely as in the recent members of the genus);
scutellum not visible dorsally due to a crack in the amber piece,
but at least, as it is seen laterally, it has a moderate length;
Downloaded by [46.231.117.154] at 09:35 29 January 2016
430
A. G. K & A. N
distance between mesocoxae apparently somewhat greater and
that between metacoxae markedly greater than that between
procoxae (metacoxae apparently somewhat more widely
separated that width of femora, but somewhat less than width of
metacoxa); distance between metacoxae markedly smaller than
width of metacoxa; metacoxa about 2.5 times as wide as long;
metaventrite somewhat shorter than ventrite 1, its posterior
edge between coxae apparently shallowly emarginate; ventrite
1 somewhat longer than hypopygidium and somewhat shorter
than ventrites 2-4 combined; hypopygidium widely rounded to
subtruncate at apex; epipleura moderately narrow and steeply
sloping downwards laterally.
Elytra about 1.1 times as long as wide combined, rather convex
at disk, evenly and gently sloping laterally, their sides slightly
arcuate and apices nearly forming a joint arc, although at suture
a shallow but distinct sutural angle expressed. Pygidium widely
rounded at apex.
Legs well developed; tibia comparable in width at apex, which
is slightly greater than that of antennal club (haetotaxy of all
tibiae and outline of external edge of protibia invisible); femora
about twice as wide as tibiae and about 1.5 times as wide as
metacoxa long, with both anterior and posterior edges slightly
convex; metafemur about three times as long as wide; tarsi of
moderate length and rather narrow, tarsomeres 1-3 apparently
narrowly lobed.
Variations. Paratype PA 2451: Body 0.9 mm long (with
defl ected head), 0.6 mm wide; very similar to the holotype,
although the setae along the posterior edges of pygidium and
hypopygidium are somewhat sparser and shorter as well as its
metafemora are somewhat wider (about twice as wide as wide)
in comparison with the holotype. Paratype PA 2545: Body 0.8
mm long (with defl ected head), 0.6 mm wide; similar to other
specimens of the type series, although the general shape of body
looks like more narrowing posteriorly (more subovoid than
oval); elytra somewhat shorter and abdomen more extending
posteriorly; besides, its metafemora are as wide as in the
paratype PA 2451.
Additional specimen PA 1032: Body 0.9 mm long (with
defl ected head), 0.5 mm wide, 0.3 mm high; elongate oval, dark
brow to blackish with somewhat lighter appengages; strongly
convex dorsally and sufl attened ventrally; dorsum and underside
apparently with a slight shine; dorsum with very sparse and very
fi ne hairs; underside with fi ne and comparatively dense hairs;
along posterior edge of pygidium, hypopygidium and previous
ventrite as well as on anal sclerite there are somewhat longer
and fi ne hairs, although sides of pygidium (but not their apices)
and hypopygidium with sparse thicker setae (not so thick as
those in the holotype and other paratypes).
Head and pronotum with some comparatively relief, dense and
very fi ne irregular rugosity (alutaceous) and without distinct
puncturation (may be because of milky bloom); elytra with
very sparse and slightly visible punctures bearing root hairs,
interspaces between them about as fi nely alutaceous as head
and pronotum: pygidium without visible punctures, but
with rather relief sculpture; underside seemingly fi nely and
densely sculptured (not smoothed); sides of metaventrite with
smoothed and slightly depressed lateral area at each side before
coxa allowing femur to move; this area distinctly bordered by a
curved line returning to anterior edge of coxa and composing
about a third of total length of metaventrite.
Antennae 11-segmented and rather long (almost 4/5 as long
as head wide), antennal club about 1/4 of total antennal club
and with comparable length of antennomeres composing it;
ultimate maxillary palpomere much narrower than subultimate,
about four times as long as thick and slightly narrowing
apically; scutellum very wide and widely rounded at apex, more
than three times as wide as long; abdomen slightly extended
beyond elytral apices; metacoxa about 3.5 times as wide as long;
pygidium subtruncate at apex; hypopygidium subtruncate to
shallowly emarginate at apex; short apex of anal sclerite slightly
exposed; metafemur about twice as long as wide; metatibia
with long hairs along its outer edge; tarsomeres 1-3 apparently
narrowly lobed, tarsal claws moderately short and thin.
Pastillocenicus polyaki nov. sp.
(Figs 34–38, 81)
Material. Holotype PA 8980, female [almost complete beetle
with strongly defl ected head and clear integument included in a
rather small amber piece. One crack transverse to body plane of
the beetle is approached ventrally to its defl ected head. Dorsum
of the beetle is lying close to the amber surface, an irregular hole
is opened at the place of its median basal part of pronotum,
scutellum and median basal parts of elytra; besides, a milky gas
vesicle is located at apex of mandibles and between all coxae].
Etymology. e epithet of this new species is devoted to
Vladimir A. Polyak, friend of the senior author from the
childhood, when we are spent a lot of time to in mountains and
forests, observing life of wild animals, including insects.
Diagnosis. is new species diff ers from all congeners in the
more oviform body with rather shining dorsum and particularly,
in contrast to other new species, very shining head. Besides,
its labrum seems to be longer than that in other species. is
new species diff ers also from Pastillocenicus grandiclavis nov.
sp. in the markedly shorter elytra, wider anterior edge of the
shorter frons, which is far extended along anterior edge of eye
as a rather thin fold, peculiarities of pubescence, puncturation
and sculpture of integument, confi guration of line isolating
a smoothed place before metacoxae, metafemur less widened
apically, pro- and mesofemora with more convex anterior and
posterior edges, and subrectilinear inner edge of all tibiae. P.
polyaki nov. sp. diff ers from P. longifrons nov. sp. in a shorter
Figures 34–38
Pastillocenicus polyaki nov. gen., nov. sp. (Nitidulidae): 34, body, dorsally;
35, idem, laterally; 36, head, anteroventrally; 37, metacoxa and precoxal
depression, ventrally; 38, metafemur and tibia, ventral. Scales: A, to fi gs
34, 35 representing bar of 0.5 mm; B, to fi gs 36-38 representing bar of
0.25 mm.
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Polyphaga from Eocene French amber
431
abdomen; less developed pubescence; head with much shorter
frons; smaller eyes; shorter pronotum, less arcuate at sides and
concave at base; somewhat more distinct posterior angles of
pronotum; shorter elytra; more widely separated metacoxae;
and somewhat narrower femora and tibiae.
Description. Body 0.7 mm long (with defl ected head), 0.6 mm
wide, 0.3 mm high; oval, blackish with dark brown appendages;
rather convex dorsally and slightly convex ventrally; dorsum
rather shining with a bronze lustre and underside with a slight
shine; dorsum nearly glabrous, although in distal half of elytra
with very sparse and rather reddish hairs, about 2/3 as long as
distance between their insertions and in apical part of head very
small and slightly visible hairs; underside with fi ne and dense
hairs; along posterior edge of pygidium and hypopygidium
along posterior edge of penultimate ventrite there is a regular
row of rather long and stout hairs (about twice longer than
those on elytra).
Head with some trace of weak, sparse and very fi ne irregular
transverse rugosity (nearly alutaceous) and with extremely small
and very sparse punctures between eyes; pronotum completely
smooth and with extremely small punctures; elytra with similar
punctures, which in the distal half having a hair, interspaces
between them fi nely and smoothly alutaceous; pygidium not
visible; underside seemingly fi nely and densely sculptured (not
smoothed); sides of metaventrite with smoothed and slightly
depressed lateral area at each side before coxa allowing femur to
move, isolated by curved line turning to anterior edge of coxae
and nearly as long as half of total length of metaventrite.
Head weakly convex and with a very short and wide frons, not
more as long as distance between very large eyes (composed of
moderately large facets); eyes seemingly extended on underside,
although along anterior edge of eye a long and narrow fold
continuing frons disposed, which is nearly meeting with
anterior angle of pronotum; labrum far exposed from under
frons (about as long as frons before eyes), unilobed and with a
rather convex anterior edge; mandibles moderately long, with
bidentate apices and far exposed from under frons; antennae
invisible.
Pronotum almost 1/3 as long as elytra, rather and evenly
convex, with steeply sloping sides, its anterior edge bisinuate,
lateral edges subrectilinear, posterior edge regularly emarginate,
both anterior and posterior angles with somewhat rounded
apices; scutellum mostly missing, only left part of base visible;
distance between mesocoxae apparently much greater and
that between metacoxae yet more greater than that between
procoxae (that between metacoxae apparently twice as great as
femur wide); metaventrite somewhat shorter than ventrite 1, its
posterior edge between coxae apparently shallowly emarginate;
metacoxae about 3.5 times as wide as long and distance between
them about as great as width of metacoxae; ventrite 1 about as
long as hypopygidium and somewhat longer than ventrites 2-
4 combined; submetacoxal line not expressed; hypopygidium
widely rounded at apex; epipleura moderately narrow and
steeply sloping downwards laterally.
Elytra about 5/6 as long as wide combined, rather convex at
disk, evenly and gently sloping laterally, their sides slightly
arcuate and apices nearly forming a joint arc, although at suture
a shallow but distinct sutural angle expressed; pygidium widely
rounded at apex.
Legs well developed; tibiae apparently comparable in width at
apex, somewhat less than a half as wide as femora (chaetotaxy
and outline of external edge of protibia invisible, but metatibia
subtrapezium-shaped and with long hair along its outer edge);
femora more than twice as wide as tibiae, with both anterior
and posterior edges slightly convex; metafemur about twice
as long as wide; tarsi of moderate length and rather narrow,
metatarsomeres 1-3 narrowly lobed, claw moderately long and
narrow.
Pastillocenicus grandiclavis nov. sp.
(Figs 39–46, 80)
Material. Holotype PA 602, female [complete beetle with
not defl ected head and almost clear integument included in a
subquadrangular amber piece].
Etymology. Named after the Latin ‘grandis’ (large, big, great)
and ‘clavus’ (nail, peg, club).
Diagnosis. is new species seems to be more subquadrate than
other congeners with wider head and nearly rectilinear elytral
sides. Its head is characterized by the rather coarce sculpture,
but not expressed puncturation. See also the diagnoses of two
Figures 39–46
Pastillocenicus grandiclavis nov. gen., nov. sp. (Nitidulidae): 39, body,
dorsally; 40, body, laterally; 41, head, anterodorsally; 42, head and
prosternum, ventrally; 43, pterothorax and abdomen, ventrally; 44,
antenna, dorsally; 45, protibia and tarsus, dorsally; 46, metafemur, ventral.
Scales: A, to fi gs 39-41, 43 representing bar of 0.5 mm; B, to fi g. 44
representing bar of 0.25 mm; C, to fi g. 42 representing bar of 0.5 mm; D,
to fi gs 45, 46 representing bar of 0.5 mm.
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432
A. G. K & A. N
other new species of the genus.
Description. Body 0.9 mm long (with not defl ected head),
0.7 mm wide, 0.4 mm high; oval, blackish with dark brown
appendages; rather convex dorsally and slightly to moderately
convex ventrally; dorsum rather shining (but without bronze
luster) and underside with a slight shine; dorsum with very
sparse and slightly conspicuous hairs (uniformly spread
throughout dorsal sclerites), 1/2-3/5 as long as distance
between their insertions; underside with fi ne, denser and more
conspicuous hairs becoming to posterior edges of pygidium
and hypopygidium as well as posterior edge of penultimate
ventrite longer and markedly more conspicuous (up to twice
longer than those on dorsum).
Head with very fi ne lines of somewhat transverse cellularity,
and with very small and very sparse punctures; pronotum
completely smooth and with extremely small punctures; elytra
with similar punctures, interspaced even smaller punctures and
smoothly alutaceous; pygidium invisible; underside very fi nely
and very sparsely punctured, fi nely and densely sculptured (not
smoothed); sides of metaventrite with smoothed and slightly
depressed lateral area at each side before coxa allowing femur to
move, isolated by rectilinear line nearly as long as half of total
length of metaventrite.
Head weakly convex, with a moderately short and not wide
frons, distinctly shorter than distance between very large
eyes (composed of moderately large facets); eyes extended on
underside in posterior part (where they looking like a triangle),
along anterior edge of eye a rather short and narrow fold
continuing frons disposed; labrum far exposed from under
frons (markedly shorter than length of frons before eyes),
unilobed and with a rather convex anterior edge; mandibles
moderately long and moderately exposed from under frons;
antennae 11-segmented and somewhat more than 2/3 as
long as head wide, their three-segmented and elongate
club consisting of about 2/7 of total antennal length and
antennomere 8 enlarged apically and looking like involved
in the club; scape and antennomere 2 subequal in length and
together nearly as long as club, antennomeres 3-5 about 1/2 as
long as preceding ones and subconical, antennomeres 6 and
7 shortest and oval, antennomeres of club subequal in width
and ultimate somewhat longer than preceding one; mentum
subquadrangular, about twice wider than long and emarginate
anterior edge; penultumate maxillary palpomere more than
twice thicker than following one; ultimate maxillary palpomere
subcylindrical to slightly narrowing apically and about fi ve
times as long as thick; ultimate labial palpomere rather wide
and slightly narrowing apically; antennal grooves well expressed
up to the level of posterior edge of mentum; gular sutures
subparallel-sided.
Pronotum almost 1/3 as long as elytra, rather and evenly
convex, with steeply sloping sides, its anterior edge very slightly
bisinuate, lateral edges slightly arcuate, posterior edge regularly
emarginate, both anterior and posterior angles with somewhat
rounded apices; scutellum very transverse, subtriangular and
about six times as wide as long; prosternum with process about
as long as mentum and subparallel-sided, strongly curved along
coxae and not extending to level of posterior edge of procoxae;
distance between mesocoxae apparently much greater and
that between metacoxae yet more greater than that between
procoxae (that between metacoxae apparently twice as great
as femora wide); distance between metacoxae bout as great
as width of metacoxa; mesoventrite strongly excavate and
medially subcarinate at bottom; metaventrite about as long as
ventrite 1, its posterior edge between coxae apparently shallowly
emarginate; metacoxae about three times as wide as long and
distance between them about as great as width of metacoxae;
ventrite 1 slightly longer than hypopygidium and somewhat
longer than ventrites 2-4 combined; submetacoxal line not
expressed; hypopygidium widely rounded at apex; epipleura
moderately narrow and steeply sloping downwards laterally.
Elytra very slightly longer than wide combined, rather convex
at disk, evenly and gently sloping laterally, their sides slightly
arcuate and apices nearly forming a joint arc, although at suture
a shallow but distinct sutural angle expressed; pygidium widely
rounded at apex.
Legs well developed; tibiae rather widened along inner edge
and apparently comparable in width at apex, about 1.5 times as
wide as antennal club and nearly nearly twice as wide as pro- and
mesofemora; outer edge of meso- and metatibiae with a row of
long setae; pro- and mesofemora about twice as wide as tibiae,
with both anterior and posterior edges very slightly convex;
metafemur wider (about 2.5 times as long as wide) and more
enlarged apically; tarsi of moderate length and rather narrow,
tarsomeres 1-3 narrowly lobed (particularly in intermediate and
posterior legs), claw moderately long and narrow.
Family Kateretidae Erichson 1843
is family is known from fossils only after the
description of Amartus petrefactus Wickham 1912
from the Lower Oligocene Florissant shales and one
new genus from the Lebanese amber (Kirejtshuk
& Azar 2008), although undescribed genera and
species are present in Baltic amber as well. e recent
members of this group are associated with angiosperm
Figures 47–52
?Heterhelus expressus nov. sp. (Kateretidae): 47, body with outline dark
spot on each elytron, dorsally; 48, head, anterodorsally; 49, antennal club,
dorsally; 50, scape and antennomere 2, dorsally; 51, labial palpus, ventrally;
52, metafenur and tibia, dorsally. Scales: A, to fi g. 47 representing bar of
1.0 mm; B, to fi g. 48 representing bar of 1.0 mm; C, to fi g. 52 representing
bar of 0.5 mm.
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Polyphaga from Eocene French amber
433
fl owers. e specimens here described as new members
of this family are usually quite distinct from the sap
beetles (Nitidulidae) by the elongate, not strongly
dorsoventraly compressed and looser antennal clubs,
however, all Cybocephalinae have also elongate club,
but compact and strongly dorsoventraly compressed.
Genus Hetherelus Jacquelin du Val 1858
Type species. Cercus sambuci Erichson 1843 [= Heterhelus scutellaris (Heer
1841)], recent.
Remark. is genus was never recorded from fossils
before this study. Its recent species are associated with
infl orescences of Sambucus (Caprifoliaceae). is new
species looks rather diff erent from the recent species
(see diagnosis below), however its separation into a
diff erent genus or subgenus cannot be proposed at this
stage.
Hetherelus expressus nov. sp.
(Figs 47–52, 82)
Material. Holotype PA 1937, undefi ned sex
[incomplete beetle with one Diptera: Ceratopogonidae,
legs of a large insect].
Etymology. e epithet of this new species means ‘expressive’.
Diagnosis. is fossil belongs to Kateretidae because of the
quite characteristic shape of all sclerites, in particular the
transverse procoxal cavities, antennal club and legs. Its shape
of pronotum, elytra and antennal clubs show that this species
should be regarded as a member of the Palaearctic Heterhelus
Jacque du Val 1858, although its body is outlined as in Sibirhelus
corpulentus (Reitter 1900). In contrast to the East Palaearctic
Sibirhelus corpulentus, Heterhelus expressus nov. sp., like other
species of Heterhelus, has not a strongly convex pronotum, its
disk being slightly convex, although its sides at lateral edges are
quite steeply sloping, a quite distinct antennal club, and much
sparser and fi ner dorsal puncturation. e tarsal claws of the
new species are somewhat bulbous, but not toothed, as those
in Sibirhelus Kirejtshuk 1984. Nevertheless, this new species is
very distinct from the recent Heterhelus in the general outline
of its body, as that in Sibirhelus, and quite distinct basal border
of pronotum. Besides, this new species has a ratio between
prosternum and metaventrite, basal border of pronotum, and
distance between metacoxae, all characteristic of Heterhelus.
It is impossible to defi ne the sex of the holotype of this new
species, because the ultimate abdominal segment in this genus
has no clear sexual dimorphism. However, its very narrow
anterior tarsi make possible to suppose that the holotype could
be a female rather than a male.
Description. Body 3.6 mm long, 1.5 mm wide, 1.2 mm high;
moderately convex ventrally and dorsally; subunicolorous
reddish brown with a round dark spot at inner apical angle
of each elytron; dorsum with rather fi ne hairs, about as long
as distance between their insertions (hairs on pygidium more
conspicuous).
Head with distinct punctures, markedly smaller than eye
facets in diameter, intersparces between them about as great
as a puncture diameter and somewhat alutaceous; pronotum
and pygidium with punctures similar to those on head, but
somewhat smaller and sparser and interspaces between them
alutaceous; elytra with markedly smaller and sparser punctures,
interspaces between them 2-3 puncture diameters; puncturation
and sculpture of underside invisible because of milky condition
of amber.
Head somewhat shorter than distance between eyes (consisting
of rather large facets), subfl attened, its anterior edge distinctly
emarginate and acute tops of lateral angles; labrum unlobed and
moderately exposed from under frons; mandibles well developed
and moderately projecting anteriorly; antennae somewhat
shorter that head wide, scape moderate large, antennomere 2
about 3/4 times as long as scape and more or less longer than
the rest antennomeres of fl agellum, club composing of about
2/7 of total antennal length with ultimate segment largest and
subacute at apex; mentum apparently of usual shape and about
2.5 times as wide as long; last maxillary palpomere narrow and
rather long; last labial palpomere strongly widened apically and
obliquely truncate at apex.
Pronotum transverse, slightly convex at disc and rather steeply
sloping at lateral edges, distinctly bordered along sides and
base as well as with widely rounded all angles, its anterior edge
somewhat convex, sides regularly arcuate and posterior edge
almost straight, but with a shallow sinuation at each posterior
angle; scutellum subsemicircular; prosternum moderately short
with a quite narrow process; metaventrite about twice as long
as prosternum; outlines of ventrites not visible.
Elytra somewhat wider than pronotum, gently convex at disk
and rather steeply sloping at sides, slightly longer than wide
combined, with subparallel sides, their apices transversely
truncate; pygidium nearly completely exposed from under
elytral apices, with very widely rounded to truncate apex.
Right anterior and median legs complete and well developed;
pro- and mesofemora as well as pro- and mesotibiae compared
shape; tibiae about twice as wide as antennal club and femora
nearly twice wider than tibiae; protibia apparently with bordered
outer edge; protarsus about 1/3 as wide as tibiae; tarsal claws
rather long and slightly bulbous at base.
Genus Eoceniretes nov. gen.
Type species. Eoceniretes yantaricus nov. gen., nov. sp.
Etymology. Named after Eocene and ‘retes’, usual suffi x for
generic names in the family Kateretidae; masculine gender.
Diagnosis. Most characters of this fossil are similar to those in
the recent species of Brachypterus Kugelan, 1794, which range
in the North Hemisphere, mostly in the Holarctic regions. But
the shape of penultimate abdominal segment resembles that
of species of Brachypterolus Grouvelle, 1912. For both genera
the strongly toothed tarsal claws are characteristic, although
development of the tooth within Brachypterus is rather variable.
At the same time the unique shape of scutellum of the new
species makes possible to erect for it a new genus. From these
mentioned recent genera Eoceniretes yantaricus nov. gen., nov.
sp. diff ers also in the not strongly convex body, comparatively
shorter elytra, remaining completely uncovered three last
abdominal segments, and narrowly explanate pronotal sides.
e new genus shows some similarity with the member of the
family recently described from the Lower Cretaceous Lebanese
amber (Kirejtshuk & Azar 2008), although it diff ers from
the latter in the larger and not so narrow body, antennal club
not subparallel-sided and with subacute apex, wider mentum
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434
A. G. K & A. N
with subquadrangular shape, arcuate outline of eye edge on
underside, almost regularly oval pronotum with distinctly
bordered and narrowly explanate sides, characteristic scutellum,
much shorter elytra, ends of the two penultimate abdominal
segments markedly wider than base of following segments,
widely separated metacoxae, and strongly toothed tarsal claws.
e outstanding feature of this genus is also the absence of
visible tibial spur.
Eoceniretes yantaricus nov. sp.
(Figs 53–57, 83–84)
Material. Holotype PA 9167, undefi ned sex [complete beetle
with strong milky cover around its abdomen included in an
elongate amber piece].
Etymology. e epithet of the new species is formed from
Slavic adjective with meaning amber.
Description. Body 2.2 mm long (with rectifi ed abdomen
apparently 2.5 mm long), 0.9 mm wide, 0.5 mm high; body
elongate oval, moderately convex dorsally and ventrally; dark
brown with lighter appendages (nearly reddish); dorsum with
very conspicuous silvery grey hairs, somewhat longer than
distance between their insertions; underside with similar
pubescence, but not so conspicuous.
Head and pronotum with rather deep punctures, about as
large as eye facets in diameter, interspaces between them
slightly narrower than a puncture diameter and without clear
microreticulation; elytra with similar, but partly elongate
punctures, interspaces between them markedly narrower
than those on head and pronotum; prosternum with
pucturation becoming microtuberculation and rather coarse
microsculpture; metaventrite and ventrites 1-3 with punctures
almost as large as those on head and pronotum, but much
shallower and interspaces between them with rather contrasting
microreticulation.
Head about as long as distance between eyes (consisting
of rather large facets), subfl attened, anterior edge of frons
shallowly emarginate and with narrowly rounded lateral angles;
labrum well exposed from under frons; mandibles moderately
developed and well exposed from dorsally; antennae somewhat
shorter than head broad; oval scape somewhat longer than
subcylindrical antennomere 2, subconical antennomere 3
rather narrow and nearly as long as scape; antennomeres 4-7
suboval, comparable in shape and size, less than half as long as
antennomere 4; antennomere 8 similar to previous, but slightly
larger; club somewhat wider than scape and composing much
less than 1/3 of total antennal length, antennomeres 9 and
10 comparable in shape and size and each of them somewhat
shorter than antennomere 11 (the latter subacute at apex); eyes
on underside somewhat enlarging and somewhat wider than
dorsal side, with arcuate outline, minimal distance between
them almost twice greater than width of mentum; mentum
covered by milky cover and apparently of usual shape, i.e.
subquadrangular and not wider than length of antennal club,
its emarginate anterior edge disposed much anteriorly than the
level of anterior edge if eyes; ultimate labial palpomere suboval,
somewhat narrowed apically and almost twice longer than
wide; antennal grooves very slightly impressed.
Pronotum transversely oval, moderately and evenly convex and
with rather rounded all angles; its sides arcuate and narrowly
explanate, anterior and poisterior edges weakly convex;
scutellum subtriangular and with rather broader base, as a result
of this with very peculiar outline; prosternum gently convex
with a narrow process subparallel-sided, somewhat narrower
than antennal club and narrowly rounded at apex; distance
between mesocoxae markedly narrower and that between
metacoxae almost four times as great as that between procoxae;
mesoventrite very short. Metaventrite about 1 and 1/4 as long
as prosternum and medially nearly subfl attened, its posterior
edge between coxae nearly straight; ventrites 1-3 considerably
shorter than ventrite 4 and hypopygidium, ventrite 4 at apex
much wider than base hypopygidium and the latter somewhat
longer than penultimate ventrite and widely rounded at apex;
two penultimate tergites completely and apex of previous tergite
remaining uncovered by elytra, laterosternites comparatively
wide.
Elytra nearly as long as wide combined, subparallel-sided,
moderately convex at disk and steeply sloping at sides, longest
at outer apical angles and their apices subrectilinearly inclined
to suture.
Legs well developed; protibia with a simple and unbordered
outer edge; tibiae similar in shape, comparable in width and
at subtruncate apex about 1 and 2/3 as wide as antennal club
and without distinct spur at tarsal insertion; profemur about
Figures 53–57
Eoceniretes yantaricus nov. gen., nov. sp. (Kateretidae): 53, body with
rectifi ed abdomen, dorsally; 54, mentum and eyes, ventrally; 55, apex of
labial palpus, ventrally; 56, antenna, dorsally; 57, metafemur and metatibia,
ventral. Scales: A, to fi g. 52 representing bar of 0.5 mm; B, to fi gs 54, 56
representing bar of 0.5 mm; C, to fi g. 57 representing bar of 0.5 mm.
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Polyphaga from Eocene French amber
435
1 and 1/3, mesofemur about 1.5 times and metafemur nearly
twice as wide as corresponding tibiae, of usual outline and
with somewhat convex anterior and posterior edges; protarsus
somewhat wider than antennal club, meso- and metatarsi about
as wide as antennal club, their claws moderately developed and
strongly toothed at base.
Remarks. e sex of this specimen cannot be defi ned
due to a rather thick milky cover around its abdomen,
which can have sexual character in the shape of
pygidium.
Family Smicripidae Horn 1879
is monogeneric family has never been recorded
from fossils before. e new fossil species here described
is rather similar to the recent species (see the below
diagnosis). All the recent species of the genus occur
in Central America and are registered from decaying
fl owers (from the genus Pseudobombax - Bombacaceae),
leaf litter and under bark. e presence in the Eocene
of Western Europe of this American family supports
its possible great antiquity, at least in the Cretaceous. A
similar distribution occurs for the Paleogene dragonfl y
family Palaeomacromiidae Petrulevicius, Nel & Muzon,
1999, known from Argentina and Italy (Petrulevicius
& Nel 2007).
Genus Smicrips Le Conte 1878
Type species. Smicrips palmicola Le Conte 1878, recent.
Smicrips europeus nov. sp.
(Figs 58–60, 85–87)
Material. Holotype PA 4167, male [complete beetle included
into a fl at amber plate].
Etymology. e epithet of this new species refers to the
continent of origin of the type specimen.
Diagnosis. e fossil diff ers from all recent species in its
somewhat broader body with the head wider than pronotum,
much wider anterior part of pronotum and much wider and
subfl attened head with the very wide frons, much narrower
and more subparallel-sided tibiae; markedly sparser dorsal
puncturation and less conspicuous fi ner pubescence as well as
smoothed integuments between punctures. Besides, it is also
distinct from:
- Smicrips distans (Sharp, 1900) (examined type series in the
Natural History Museum in London) in the fl attened dorsum,
much longer antennomere 2, much narrower and subparallel-
sided tibiae;
- Smicrips mexicanus (Sharp, 1900) (examined type series in the
Natural History Museum in London) in the widely rounded
posterior angles of pronotum;
- Smicrips chontalenus (Sharp, 1900) (examined type series in
the Natural History Museum in London) in widely rounded
posterior angles of pronotum.
As this new species is very similar to the recent members of the
genus, some of its characters are omitted in the description.
Description. Body 1.6 mm long, 0.5 mm wide, 0.2 mm high;
elongate, subfl attened dorsally and ventrally; subunicolorous
brownish; slightly shining; dorsum and ventrite with well
conspicuous, recumbent and short yellowish hairs about two
times as long as distance between their insertions; thoracic
sterna and underside of head with fi ner, shorter and less
conspicuous hairs.
Head and pronotum with more or less distinct, shallow and
small punctures, somewhat smaller than eye facets in diameter,
interspaces between them markedly greater than a puncture
diameter and more or less alutaceous, but at pronotal sides
punctures becoming denser. Elytra with similar puncturation
and sculpture, although punctures less distinct and sculpture
somewhat more relief; uncovered tergites, ventrites 2-4 and
hypopygidium with somewhat smaller and denser punctures
than on head and pronotum, with very dense and fi ne
microreticulation; prosternum as punctured as head and middle
of pronotum, but with very dense and fi ne microreticulation;
metaventrite and ventrite 1 with obsolete puncturation and
microsculpture similar to that on prosternum.
Head subtriangular and prognathous, somewhat shorter than
Figures 58–62
Species of Smicrips (Smicripidae): 58-60, S. europeus nov. sp.: 58, body,
dorsally; 59, idem, ventrally; 60, antenna, dorsally; Smicrips sp. (Natural
History Museum in London: ‘W. Indies’, ‘Soubise (Windward side),
Grenada, W.I., H.H. Smith’); 61, mentum and labial palpi, ventral;
Smicrips sp. (Natural History Museum in London: ‘W. Indies, St. Lucia,
balionnean, 29.IV.84, H. Alam’); 62, mentum and labial palpi, ventral.
Scales: A, to fi gs 58, 59, 61, 62 representing bar of 1.0 mm; B, to fi gs. 60
representing bar of 0.25 mm.
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436
A. G. K & A. N
long, slightly wider than pronotum, with rather large eyes not
reaching prothoracic segment, subfl attened; labrum apparently
well exposed but boundary between it and frons invisible
because a thick layer of amber; mandibles slightly exposed from
under frons and labrum; antennae 11-segmented with three
segmented loose club composing about 2/7 of total antennal
length and consisting of antennomeres comparable in width and
with ultimate one longest; scape about 2/3 as long as antennal
club and somewhat narrower than the latter; antennomere
2 somewhat shorter and somewhat narrower than scape;
antennomeres 3-5 comparable in length, somewhat narrower
and shorter than antennomere 2; antennomeres 6-8 comparable
in length and shortest in antennae; mentum rather large, about
3/4 as long as wide, with subrectilinearly narrowing sides and
strongly excised anterior edge, its anterior angles pointed and
projecting anteriorly; ultimate maxillary palpomere subconical,
slightly narrowing apically, about three times as long as wide at
its thickest place; labial palpi moderately small and its ultimate
segment somewhat longer than wide at apex.
Pronotum with subtruncate anterior and posterior edges, all
angles rounded, widest in anterior 1/2, slightly and gradually
narrowing anteriorly from base, posterior and lateral edges not
bordered; its disk rather fl at and sides rather sloping; scutellum
moderately large and subtriangular; prosternum slightly convex
and its process in narrowest place extremely narrow, moderately
projecting till the level of posterior edge of coxae, nearly
subparallel-sided and subtruncate at apex; procoxae distinctly
transverse and with exposed trochantin; mesoventrite slightly
convex and with the same plane as metaventrite; distance beween
mesocoxae nearly three times and that between metacoxae
nearly four times as great as that between procoxae; metaventrite
slightly longer than prosternum with process, slightly convex,
without both median line and paracoxal line before metacoxae;
submesocoxal lines following closely to posterior edge of cavities
and along anterior half of metepisterna; submesocoxal line not
expressed; metacoxae not very wide; ventrites 1 somewhat
longer than ventrites 2-4 combined and slightly shorter than
wide, hypopygidium convex in the middle of its posterior edge
and sinuate at each sides before apex; epipleura well expressed;
pygidium apparently as long as wide and widely subtruncate at
apex; apex of anal sclerite distinctly exposed from under apex
of pygidium.
Elytra about 1 and 2/5 as long as wide combined, truncate at
apices, subparallel-sided; with a small sutural angle, subfl attened
at disk and rather steeply sloping at sides and somewhat declined
on ventral side, adsutural lines absent, leaving uncovered
pygidium and previous tergite.
Legs well developed and quite narrow; trochanters rather of
elongate type; tibiae slightly compressed, narrow, about as wide
as scape, almost comparable in length and slightly widening
apically; their apices somewhat oblique to subtruncate and
with two clear spurs, outer edge of tibia simple; femora of usual
shape, pro- and mesofemora about two times and metafemur
about four times as wide as tibiae; tarsi fi ve-segmented and
comparatively short, tarsomeres 1-3 narrowly lobed (in anterior
legs 2/3 as wide as tibiae, in intermediate and posterior ones
somewhat narrower); tarsomere 4 very small and tarsomere
distinctly longer than previous tarsomeres combined; claws
simple, narrow and short.
Superfamily Tenebrionoidea Latreille 1802
Family Anthicidae Latreille 1819
Fossils of this family are known mostly from Baltic
amber (Klebs 1910; Abdullah 1964; Larsson 1978;
Spahr 1981; Hieke & Pietrzeniuk 1984, etc.), where the
subfamilies Macratriinae Le Conte 1862; Steropinae
Jacquelin du Val 1863; Tomoderinae Ranadona 1961,
and Anthicinae (including Notoxini Stephens 1829)
have been recorded. e oldest record of this family is
in the Lower Cretaceous Lebanese amber (Kirejtshuk
& Azar 2008) and Burmese amber (Cockerell 1917:
Eurygeniinae Le Conte 1862). e further Cenozoic
records originated from Palaeocene (?Eocene) Sunchal
(Cockerell 1926), boundary Eocene-Oligocene in
Bembridge Marls (Kirejtshuk et al. in press), Lower
Oligocene Florissant shales (Wickham 1910) and Aix-
en-Provence (France) (Oustalet 1874) as well as from
the Miocene Izarra beds (Spain) (Arillo & Ortuno
1997).
Recent representatives of the family spread in
all zoogeographical regions. Larvae and adults of
most species are mycetophagous, although few are
predaceous, and live in decaying matter of plant origin,
frequently in leaf litter and other kinds of debris. Some
of them are characteristic of the edges of lakes and
other bodies of water.
Subfamily Eurigeniinae LeConte 1862 (?)
e new fossil seems to belong to this subfamily,
although it is rather diff erent from its other members
(see diagnosis below). is group is known in fossils
only after the description of ‘Eurygenius’ wickhami
Cockerell 1917 from the Lower Cretaceous Burmese
amber, which could be indeed a member of the
subfamily Macratriinae Leconte, 1862 (Kirejtshuk &
Azar 2008).
Genus Oisegenius nov. gen.
Type species. Oisegenius antiquus nov. gen., nov. sp.
Etymology. e name of this new genus is formed from type
locality (Oise Department in north of France) and ‘genius’, usual
suffi x for generic names in the family Anthicidae; masculine
gender.
Diagnosis. is fossil reminds the other representatives
of the subfamily, but in general it is somewhat smaller in
size and more robust. e type species of this new genus is
characterized by the procoxal cavities seeming to be opened or
very narrowly closed, contiguous metacoxal cavities, neck less
than half as wide as head at eyes, not elongate antennomere 3,
not emarginate eyes, lack of distinct spur, and rugose sculpture
of the neck. ese peculiarities make possible to approach this
fossil to Eurygeniinae (well expressed ‘frontoclypeal’ suture,
microtuberculate pronotum and elytra, and extremely narrow
distances between both mesocoxae and metacoxae), although
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Polyphaga from Eocene French amber
437
it has some clear peculiarities that allow distinguishing it from
the other groups of the subfamily. In fi rst turn these concern
the eyes that are not emarginate anteriorly in contrast to the
known representatives of the subfamily. However even among
the recent representatives this character varies to a great level,
and some forms have a very feeble emargination. e fl ange of
collar of the new fossil is rather narrow, but more or less distinct
(especially clearly visible in the paratype). e lack of distinct
tibial spurs makes it distinct from all members of Eurygeniinae
and Macratriinae having such spurs.
e subfamily Eurygeniidae is distinct due to the wide neck
and emarginate eyes. In our new taxon this latter character
is not expressed at all (see above and the description below).
Besides, it is diffi cult to fi nd a clear placement for this fossil in
any of the known tribes of the subfamily. It is most probable
that the new fossil should be regarded as a member of the tribe
Eurygeniini because of the pronotum bearing a distinct apical
fl ange of collar, although its procoxal cavities are not visible
enough to determine whether they are open externally or not.
Besides, the type species of the new genus has very coarse and
diff use scupture of dorsum, rather similar to that in species of
Eurygenius Ferté-Senecrète, 1849.
‘Eurygenius’ wickhami Cockerell 1917 seems to be also a
member of the subfamily Macratriinae, but not Eurygeniinae.
At least its long and modifi ed antennae and shape of maxillary
palpi are similar to those in the species of Macratriinae from the
Lebanese amber (Kirejtshuk & Azar 2008) rather than to those
of Eurygeniinae. e species here considered diff ers from that
from Burmese amber according to the description of the latter
in the smaller and much more robust body, shorter head with
smaller eyes and smaller mandibles, shorter antennae, shorter
maxillary palpi with the rather widened ultimate palpomere,
not ‘subcircular’ pronotum, shorter legs, and lack of tibial
spurs, subuniform sculpture, and puncturation of elytra.
Oisegenius antiquus nov. sp.
(Figs 63–69, 88)
Material. Holotype PA 1154, male [almost complete
beetle with missing ultimate right antennomere
and slightly covered with milky fogging, together
with one medium-sized Coleoptera: Anobiidae, two
Hymenoptera: Chalcidoidea and one Psocoptera:
Lachesillidae, included in an elongate amber piece. A
print of another coleopterous specimen (probably the
same anthicid species) is at about 10 mm from the type].
Paratype PA 1781 11/15, female [anterior part of the
beetle showing the head with appendages, prothoracic
segment with anterior legs and base of pterothorax
with base of elytra included in a quadrangular side
of an elongate amber bar, together with the paratype
of Colotes constantini n. sp (10/15) and one small
Hymenoptera (11/16)].
Description. Holotype. Body 4.6 mm long, 1.7 mm wide, 1.3
mm high; elongate, rather convex both dorsally and ventrally;
brownish grey with some silver-bronze lustre; dorsum with
very dense and fi ne, suberect, moderately conspicuous, silvery
grey hairs, less dense on head and pronotum and more dense
on elytra, where they are about fi ve times as long as distance
Figures 63–69
Oisegenius antiquus nov. gen., nov. sp. (Anthicidae): 63, body, laterodorsally;
64, idem, lateroventrally; 65, pronotum with base of head and base of elytra,
dorsally; 66, apex of antenna, dorsally; 67, ultimate maxillary palpomere,
ventrally; 68, mesotarsus, dorsally; 69, metatarsus, dorsally. Scales: A, to
fi gs 63, 64 representing bar of 1.0 mm; B, to fi g. 65 representing bar of 1.0
mm; C, to fi gs. 66, 68, 69 representing bar of 1.0 mm.
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438
A. G. K & A. N
Figures 70–74
70, Cyphon gallicus nov. sp. (Scirtidae), holotype PA 3404, body, lateroventral, length of inclusion 3.4 mm. 71, C. lobanovi nov. sp. (Scirtidae), holotype PA
5605, body, laterodorsal, length of inclusion 2.1 mm. 72, 73, 74, Colotes constantini nov. sp. (Melyridae), holotype PA 9530, length of inclusion 2.4 mm: 72,
body, dorsal; 73, idem, lateral; 74, idem, ventral.
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Polyphaga from Eocene French amber
439
Figures 75–82
75, 76, 77, Colotes impexus nov. sp. (Melyridae), holotype PA 890, length of inclusion 2.2 mm: 75, body, lateroventral; 76, idem, dorsal; 77, idem,
posterodorsal. 78, 79, Pastillocenicus longifrons nov. gen., nov. sp. (Nitidulidae): 78, paratype PA 1032, body, lateroventral, length of inclusion 0.9 mm; 79,
holotype PA 5132, body, ventral, length of inclusion 1.0 mm. 80, P. grandiclavis nov. gen., nov. sp. (Nitidulidae), holotype PA 602, body, ventral, length of
inclusion 0.9 mm. 81, P. polyaki nov. gen., nov. sp. (Nitidulidae), holotype PA 8980, body, dorsal length of inclusion 0.9 mm. 82, Hetherelus expressus nov.
sp. (Kateretidae), holotype PA 1937, body, dorsal, length of inclusion 3.6 mm.
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440
A. G. K & A. N
Figures 83–88
83, 84, Eoceniretes yantaricus nov. gen., nov. sp. (Kateretidae), holotype PA-9167, length of inclusion 2.2 mm: 83, body, dorsal; 84, idem, lateral. 85, 86, 87,
Smicrips europeus nov. sp. (Smicripidae), holotype PA 4167, length of inclusion 1.6 mm: 85, body, lateroventral; 86, idem, lateral; 87, antenna. 88, Oisegenius
antiquus nov. gen., nov. sp. (Anthicidae), holotype PA 1154, body, lateroventral, length of inclusion 4.6 mm.
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Polyphaga from Eocene French amber
441
between their insertions; underside with much shorter and
subrecumbent hairs, scarcely longer than distance between
their insertions.
Head integument not visible because of milky cover and
its position, but it seemingly very densely microtuberculate
(about as pronotum) and sculpture of neck rather rugose than
smoothed; pronotum also with milky cover, although dense
microtuberculate integument clearly visible, very dense tubercles
about as large as eye facets in diameter; elytra with similar
sculpture, but tubercles 2-3 times as large as those on head and
pronotum, interspaces between them somewhat narrower than
a tubercle diameter, integument smoothly microreticulated;
prosternum with rather shallow and quite distinct punctures
about as large as tubercles on head and pronotum, interspaces
between them somewhat narrower than a puncture diameter,
very densely and fi nely microreticulated; metaventrite about
as punctured and sculptured as prosternum, but punctures
somewhat denser, shallower and less distinct; ventrites with
indistinct, denser and smaller punctures than those on thoracic
sterna, interspaces between them with rather contrasting dense
microreticulation; although hypopygidum very densely and
very fi nely microtuberculate.
Head markedly longer than distance between rather large eyes
(consisting of comparatively fi ne facets) with not emarginate
anterior edge, gently swollen frons transversely abrupt at
anterior edge; antennal insertions rather close to eye edge and
frons above them slightly elevated; temples unraised; neck rather
short (about as long as each of antennomeres 3-5 and about
3/5 as wide as as head at eyes), with distinct subbasal line and
almost 3/5 as wide as head at eyes; ‘frontoclypeal’ suture well
expressed; labrum and mandibles clearly not visible; antennae
subfi liform and almost twice as long as head wide; scape
somewhat wider than fl agellomeres (except ultimate one) and
about four times as long as thick; antennomere 2 less than half as
long as scape; antennomeres 3-8 subcylindrical: antennomeres
3-5 somewhat longer than antennomeres 6-8; antennomeres
9 and 10 enlarged apically and dorsoventrally compressed;
ultimate antennomere thickening in proximal half and in distal
half narrowing to acuminate apex; eyes arcuately enlarging on
underside; mentum and palpomeres mostly invisible, although
ultimate maxillary palpomere quite large and securiform, and
ultimate labial palpomeres rather small, elongate and somewhat
narrowing apically; gular sutures invisible.
Pronotum slightly longer than wide (to nearly as long as wide),
moderately and evenly convex at disk and steeply sloping at
sides, widest before anterior edge; its sides slightly subsinuate at
base and evenly widened anteriorly, without distinct carine; its
apex with a very narrow fl ange of collar; scutellum subtriangular
and widely rounded at apex; prosternum medially subfl attened,
its length before procoxae somewhat shorter than the latter;
procoxae subtriangular with longest outer edge and rather
projecting (apparently open posteriorly); distance between
mesocoxae and that between metacoxae extremely narrow;
mesoventrite not visible; metaventrite medially subfl attened
and about twice as long as procoxae; premetacoxal lines well
expressed; metacoxal cavities contiguous; abdominal ventrite
1 longest and about as long as ventrites 2 and 3 combined;
hypopygidium somewhat shorter than ventrite 1 somewhat
excavate before angularly excised posterior edge and with a
swollen places bearing somewhat longer hairs at each side
of subapical depression; pygidium very widely rounded to
subtruncate at apex.
Elytra complete and about 1.5 times as long as wide combined,
moderately convex at disk and steeply (subvertically) sloping
at sides, longest at very small sutural angle; widest in anterior
forth and very gently narrowing to separately rounded apices.
Legs well developed and rather long; all trochanters distinctly
of elongate type; tibiae similar in shape and size, very slightly
compressed and evenly covered with rather long setae, slightly
narrower than ultimate maxillary palpomere, obliquely
subtruncate at apex; without both outer borders and distinct
spur (or the latter very short and thin and therefore not visible
among the apical setae); femora rather enlarged apically and
widest in distal third, profemur about twice, mesofemur
somewhat more than twice and metafemur apparently about
three times as wide as corresponding tibiae; tarsi characteristic
of family, viz. protarsomere 1 widened apically (nearly as wide
as tibiae) and about twice longer than wide; protarsomere 2
somewhat shorter and narrower; metatarsomeres 1 and 2 rather
narrow with the former at least twice longer than the latter;
metatarsomere 3 about twice wider than previous ones; claws
distinctly dentate at base; protarsus somewhat wider than
antennal club, meso- and metatarsi about as wide as antennal
club, their tarsi moderately big and strongly toothed at base.
Variation. Paratype: Looking like the holotype, although its
sculpture of dorsum is clearer because of vicinity of the beetle
to the amber surface. Elytron with coarse oval punctures
reminding of the cells in the elytra of species of Cupes Fabricius,
1801.
Acknowledgements. We thank the company Lafarge-Granulat
for the help with the sampling of the fossil and the family
Langlois-Meurinne for the authorization of working in their
property. We also thank G. Hodebert (MNHN) for the
realization of the drawings, and also G. De Ploëg (MNHN)
and Dr. D. Azar (MNHN) for the careful preparation of
the material. e authors have possibility to cooperate with
many other colleagues, who willingly gave the authors ideas
and opinions. In some cases the diagnoses of new forms were
impossible without contribution of our friends and colleagues.
e authors have a pleasant duty to involve thanks to Dr. W.
Schawaller (Staatliches Museum für Naturkunde in Stuttgart)
and Dr. A.G. Ponomarenko (Palaeontological Institute of the
Russian Academy of Sciences, Moscow), D. Telnov (Latvian
Entomological Society, Riga), M.V. Barclay (Natural History
Museum in London), Dr. T. Deuve (MNHN), and R.
Constantin (Saint-Lo, France). e senior author has a pleasant
duty to express his recognition to the Museum national
d’Histoire naturelle provided him a possibility to work in this
museum in 2006, 2007 and 2008. is study was also partly
supported by the Programme of the Presidium of the Russian
Academy of Sciences ‘Origin and Evolution of Biosphere’ and
Russian Foundation of Basic Research (grants 070400540a and
07-04-92105-GFENа).
References
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Amber of Eastern Prussia and Gum Copal of Zanzibar. Transactions of
the Royal Entomological Society of London 116: 329-346.
Arillo A., Ortuno V. 1997. First records of the families Anthicidae
and Chrysomelidae from the Oligocene of Izarra (Alava, Spain).
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