First Mesozoic representative of the subfamily Liparochrinae (Coleoptera: Hybosoridae) from the Lower Cretaceous Lebanese amber

2011·0.28 MB·

by  A. G. Kirejtshuk

ZOOSYSTEMATICA ROSSICA, 20(1): 62–70 28 JULY 2011
First Mesozoic representative of the subfamily Liparochrinae
(Coleoptera: Hybosoridae) from the Lower Cretaceous Lebanese amber
Первый мезозойский представитель подсемейства Liparochrinae
(Coleoptera: Hybosoridae) из нижнемелового ливанского янтаря
A.G. KIREJTSHUK, D. AZAR & O. MONTREUIL
А.Г. КИРЕЙЧУК, Д. АЗАР, О. МОНТРЕЙ
A.G. Kirejtshuk, Zoological Institute of Russian Academy of Sciences, 1 Universitetskaja Emb., St. Petersburg,
199034, Russia; CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 57 Rue Cuvier,
F-75231, Paris, France. E-mail: [email protected], [email protected]
D. Azar, Lebanese University, Faculty of Sciences II, Natural Sciences Department, Fanar-Matn, P.O. box
26110217, Lebanon; CNRS 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon,
F-75005, Paris, France. E-mail: [email protected]
O. Montreuil, CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon,
F-75005, Paris, France. E-mail: [email protected]
One specimen among coleopterous inclusions recently recovered in Lebanese amber is de-
scribed as Libanochrus calvus gen. et sp. nov. and assigned to the subfamily Liparochrinae
of the Hybosoridae. This specimen is incomplete but a large part of its head with appenda-
ges, prothoracic segment with anterior legs, remains of the median part of the pterothoracic
underside and the lateral base of the of the right elytron make possible the conclusion on the
subfamily attribution and diagnose it among the rest of fossil and recent taxa of this family. At
present it is the oldest representative of the subfamily.
Один экземпляр среди инклюзов жуков, недавно найденный в ливанском янтаре, описан
как Libanochrus calvus gen. et sp. nov. и отнесен к подсемейству Liparochrinae из семей-
ства Hybosoridae. Этот экземпляр неполный, но большая часть его головы с придатками,
переднегрудной сегмент с передними ногами, остатки медиальной части низа птерото-
ракса и основание бока правого надкрылья позволяют cделать вывод о его подсемей-
ственной принадлежности и определить его среди представителей семейства. В настоя-
щее время он является древнейшим представителем подсемейства.
Key words: Lower Cretaceous, fossil, Lebanese amber, Coleoptera, Hybosoridae, Leparochri-
nae, new genus, new species
Ключевые слова: нижний мел, ископаемые, ливанский янтарь, Coleoptera, Hybosoridae,
Leparochrinae, новый род, новый вид
INTRODUCTION Jurassic by members of the subfamily Hy-
bosorinae (Bajan-Teg, Uver-Khangay aj-
The family Hybosoridae Erichson, 1847
mag, Mongolia ‒ Jurahybosorus mongolicus
is a worldwide distributed family within
Nikolajev, 2005 and Karatau, Kazakhstan ‒
Scarabaeoidea, being most diverse in the
Protohybosorus karatavicus Nikolajev, 2010;
area s with tropical climate and currently
P. grandissimus Nikolajev, 2010 and P. me-
comprising more than 220 species belong-
ing to 35 genera within five subfamilies sasiaticus Nikolajev, 2010). This subfamily
(Ocampo, 2006). It is represented in the is known also from the Lower Cretaceous
fossil record since the Middle or Upper of Baissa (Buryatskaya Autonomous Repu-
© 2011 Zoological Institute, Russian Academy of Scienсes
64 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER
blic, Transbaikalia, Russia; Middle Neoco- MATERIAL AND METHODS
mian: Cretohybosorus buryaticus Nikolajev,
Till present day the oldest amber with
1999 and C. striatulus Nikolajev, 1999; and
many arthropod inclusions is from Lebanon
also Leptosorus zherikhini Nikolajev, 2006),
(Azar, 1997). The Lebanese amber ranges in
from the Lower Cretaceous of Chifeng
age from the Late Jurassic to Cenomanian.
(Inner Mongolia, China: Leptophorus for-
The fossiliferous outcrops are all approxi-
tus (Ren, Zhu & Lu, 1995) (Geotrupoides)),
mately of the same age and are mainly late
from the Oligocene of Kučlin [Kutschlin]
Barremian to lowermost Aptian (Azar et
(Czech Republic: Phaeochrous tertiarum
(Deichmüller, 1881) (Bolboceras)), from al., 2003a). The precise dating of the Leba-
nese material was possible after the disco-
the Lower Miocene Dominican amber
very of stratigraphical marker fossil pollen
(Dominican Republic: Procoilodes adrastus
trapped in the amber (Dejax pers. com.). In
Ocampo, 2002 and Tyrannasorus rex Rat-
the majority of amber outcrops in Lebanon,
cliffe & Ocampo, 2001) and from the Upper
Miocene of Oeningen (Baden-Württem- the amber is found in its primary deposits.
The material studied herein comes from
berg, Germany: Hybosorus lividus Heer,
the Lower Cretaceous outcrops of Bouarij
1862). Besides, from Baissa Cretanaides
(Caza Zahleh, Beqaa, Central Lebanon)
trogopterus Nikolajev, 1996 and Protanaides
(for details see Azar et al., 2010). The ma-
sibiricus Nikolajev, 2010 from the subfa-
terial has been prepared (cut and polished),
mily Anaidinae Nikolajev, 1996 and Mima-
then included in Canada balsam medium
phodius pusillus Nikolajev, 2007 from the
between two glass cover slips as described
subfamily Mimaphodiinae Nikolajev, 2007
by Azar et al. (2003b). Observations were
are known. Recently Mesoceratocanthus
made with usual optic equipment; in partic-
tuberculifrons Nikolajev, Wang, Liu & Zi-
ular, the stereomicroscope Olympus SCX9
iang, 2010 from the Yixian formation of the
and inverted microscope Olympus CK 40
Upper Jurassic – Lower Cretaceus was de-
scribed in the subfamily Ceratocanthinae. in Muséum National d’Histoire Naturelle,
Finally, Coprologus gracilis Heer, 1847 with Paris, and also the stereomicroscope micro-
unclear subfamilian attribution was de- scope Leica MZ 16.0 in Zoological Insti-
scribed from Oeningen. In spite of the fact tute, St. Petersburg, were used.
that some mentioned indications could be The material under consideration is
regarded as somewhat doubtful, they give a temporally deposited in the Laboratory of
preliminary imagination on the distribution entomology, Muséum National d’Histoire
of this family through time. Naturelle, Paris, until a national natural his-
This paper constitutes the sixth contri- tory museum in Lebanon is established.
bution to the knowledge on fauna of Co-
leoptera from Lebanese amber (Kuschel & RESULTS
Poinar, 1993; Lefebvre et al., 2005; Kirejt-
shuk & Azar, 2008; Kirejtshuk et al., 2009a, Order COLEOPTERA
2009b), which is devoted to the families,
Suborder POLYPHAGA
oldest representation of which in fossils is
Superfamily SCARABAEOIDEA
frequently put namely in Lebanese inclu-
sions. The specimen described here is also Family HYBOSORIDAE Erichson, 1847
the oldest representative of the hybosorid Subfamily LIPAROCHRINAE
subfamily Liparochrinae Ocampo, 2006. A Ocampo, 2006
more detailed overlook on representation of Libanochrus gen. nov.
the family Hybosoridae in the fossil record
can be taken from the catalogue by Pon- Type species Libanochrus calvus gen. et
omarenko & Kirejtshuk (2011). sp. nov.
© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70
A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 65
Diagnosis. Body of comparatively small notum projecting posteriorly and armature
size. Head and pronotum comparatively of protibia (intermixed by more prominent
subflattened and without any horn or ca- and small teeth).
rina. Head feebly transverse, its outline In contrast to most other members of
evenly curved, including the short but wide the Hybosoridae, this genus is mostly char-
eye canthus and clypeus; clypeo-frontal su- acterised by the comparatively short head
ture not visible; labrum transverse, short and mandibles not exposed dorsally. The
and rather wide, flat, prominent, exposed members of the subfamily Anadiinae dif-
from above beyond apex of clypeus, located fer from the new species in their markedly
in an evenly curved excision of the latter, longer head with frons not dilated later-
its anterior edge forming a gentle curved ally, projecting and not very wide procoxae,
line, its side forming a discontinuity with coarse sculpture of integument etc.; and
the curve of clypeal side; eyes surrounded Cretanaides Nikolajev, 1996 from Anaidinae
by fronto-clypeal folds; mandibles not ex- cannot be compared with the new species,
posed dorsally. Antenna short and with first because it was described after one elytron.
club segment suboval cupular-shaped. Pro- The shape of mesocoxae and very smooth
notum widest at the distinct posterior an- integument of this specimen are similar to
gles, with deeply excised anterior edge and those of some Ceratocanthinae rather than
widely explanate sides; lateral margins cili- to those of Hybosorinae and Liparochri-
ate. All pairs of coxae narrowly separated nae, although other characters, including
or conjoined; mesocoxae feebly transverse the outline of the anterior part of its head
and not very wide. Integument of dorsum and an exposed labrum are very similar to
smooth and unpunctured. Protibia triden- those in some extant representatives of the
tate on outer edge, with distinct crenula- last two subfamilies. Libanochrus gen. nov.
tions from the base and between the main is particularly distinguished from the mem-
teeth, and bearing long setae between each bers of Ceratocanthinae by the exposed
pair of teeth in the crenulation. Femora and labrum from under clypeus, much wider
tibiae with brushes of long hair at apices. procoxae, shape of anterior legs and arma-
Comparison. The genus Libanochrus ture of protibiae. In particular, the Meso-
gen. nov. showing explanate sides of pro- zoic Mesoceratocanthus Nikolajev, Wang,
notum, first antennal club segment cupu- Liu & Ziiang, 2010 from this subfamily, in
lar-shaped, position of coxa, clypeo-frontal contrast to the new genus, has elongate and
suture not visible, shape of protibia (with larger body (12 mm) with subquadrangu-
setae between crenulations) coincides with lar prothorax, oval labrum, exposed and far
the extant members of the family Hybosori- projecting mandibles, and also narrow pro-
dae. A particular evidence of the subfamily tibiae without prominent teeth.
attribution of the new genus is the position In fossils, the new genus differs from
of transverse labrum, exposed from above, Mimaphodius Nikolajev, 2007 (Mimapho-
at the same level as the dorsal surface of diinae) in the not very convex body, ex-
head, fitting an excavation of the clyp- posed labrum, lack of clypeo-frontal suture,
eus (as observed in Liparochrus Erichson, outline of head and pronotum, rather even
1848). Libanochrus gen. nov. differs from dorsal surface of head and widely explanate
other genera in the anterior outline of its pronotal sides. It is distinct also from the
rather short head (evenly curved, not sinu- Mesozoic Cretohybosorus Nikolajev, 1999,
ate at front base of the eye canthus), mark- Leptosorus Nikolajev, 2006 and Protohypo-
edly narrower mesocoxae and type of the ar- bosorus Nikolajev, 2010 in the not exposed
mature of protibia, and also from Antiochrus mandibles, laterally extended head sides
Sharp, 1873, in the not so convex body, lack making the head transverse, widely ex-
of hairs on dorsum, posterior angles of pro- planate pronotal sides, posterior angles of
© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70
66 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER
pronotum slightly projecting backwards bles with the outer anterior edge somewhat
(posterior edge of pronotum subsinuate shortly elevated and their elevated edge is
at sides) [although the pronotum of Cre- closely clasped to the anterior edge of the
tohybosorus could be somewhat similar]; labrum. However most species of the lat-
and also Protanaides sibiricus in the ante- ter genus have the mandibles far projecting
rior part of head strongly extended later- anteriorly and pronotal sides only subex-
ally and surrounding the eyes, narrower planate or narrowly explanate. Besides, the
head, pronotum more arcuately narrowing new species has a short labrum similar to
anteriorly and rounded anterior angles and members of Seleucosorus Kuijten, 1983 (the
also different armature of protibia. Since later have the mandibles also short, but dis-
Jurahybosorus Nikolajev, 2005 (Hybosori- tinctly projecting anteriorly). Finally, the
nae) was described after the metathoracic specimen examined is comparatively small,
ventral sclerites and posterior legs only, it however many groups of the generic rank
cannot be compared with the new species. include the members with small body size.
Libanochrus gen. nov. also differs from the Libanochrus calvus gen. et sp. nov. shows
Caenozoic Procoilodes Ocampo 2002 from the most similarity to some species of Phae-
Hybosorinae (probably with somewhat ochrous and Kuijtenous Paulian, 1981, but in
similar pronotum and small body size) in addition to the mandibles which are not ex-
the flatly depressed to excavate (and not posed from under labrum, a comparatively
tuberculate) frons, not pointed labrum and short head and gently outlined head sides
smoothed surface of head and pronotum; (not subsinuate at the base of eye canthus),
and from Tyrannasorus Ratcliffe & Ocampo the new species has no trace of border along
2001 in the completely different structure edges of its pronotum.
of the transverse head (concealed man- Etymology. The name of the new genus
dibles, not projecting labrum, laterally ex- is formed from Greek ‘Libanos’ (Lebanon)
tended anterior part of the head). Finally, and a usual root of generic names in Hy-
the new genus is very distinct from Copro- bosoridae descended from the Greek ‘chros’
logus gracil is Heer, 1847 in the strongly di- (skin, surface of body; skin colour). Gender
lated sides of the head, pronotum more nar- masculine.
rowing anteriorly and concealed mandibles.
All genera of Hybosorinae with later- Libanochrus calvus sp. nov.
ally expanded clypeus have more or less ex- (Figs 1–7)
pressed sinuation at the base eye canthus,
while Libanochrus calvus gen. et sp. nov. is Material examined. Holotype. ‘TAR 39’, sex
unknown; flat piece of amber, the inclusion bad-
characterised by the even outline of the lat-
ly preserved but with characters that allow the
eral edges of its head. Besides, the head of
description and placement of the fossil, such as
all recent Hybosorinae is markedly longer
the head and most of its appendages, most of the
than that in new species and the dorsum of
prothoracic segment with anterior legs and most
the recent Hybosorinae is clearly punctured of the ventral median surface of pterothorax,
(not infrequently rather coarsely), but the left meso- and metafemora, a trace of the dorsal
head and pronotum of the new species are surface of right meso- and metafemora is located
smooth and nearly unpunctured. Hypsolo- in the epoxy between round microscope glasses.
deres Fairmaire, 1893 has an appearance Pieces of yellowish to very dark organic matter
and very small gas vesicles surround the inclu-
dorsally somewhat similar to Libanochrus
sion. The remains of the body seemed to be some-
gen. nov., because its rather long labrum
what pressed before entombing in the fresh resin.
and mandibles are turned ventrally. Some
Most parts of integument are masked by very
species of Phaeochrous Laporte, 1840
small pieces of organic matter and gas vesicles.
demons trate comparatively short exposed The specimen examined seemed to be rather soft
labrum (as in the new species) and mandi- when it became stuck to the resin but still alive
© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70
A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 67
Fig. 1. Libanochrus calvus gen. et sp. nov., ho-
lotype. Head and pronotum, dorsal view. Head
width 2.6 mm.
Fig. 2. Libanochrus calvus gen. et sp. nov., holo-
type. Head and thorax, ventral view. Head width
(being evidence of the traces of moving in the
2.6 mm.
amber), then it was destroyed and decayed before
fossilization (as a result its remains including ap-
pendages became additionally depressed). Head transverse, somewhat depressed
Description of the holotype. Length of medially and widely explanate along frons,
head 1.4, width of head 2.6, width of avail- much narrower than pronotum; median part
able part of pronotum 3.4 mm (complete of frons apparently partly isolated by fine
body length of this specimen before its de- and oblique swellings at each side. Eyes not
struction could be about 5.0 mm). Head clearly outlined (because the layer of small
and pronotum without hairs; pronotal and pieces of organic matter and very small gas
elytral sides, and legs with moderately long bubbles covers them) apparently finely fac-
and fine hairs (particularly dense on entire eted eyes (although facets scarcely visible
surface of femora and they are arranged in and eyes seeming translucent) and with lat-
a long row along the anterior edge and two eral edge surrounded by fold of frons. Ulti-
shorter ones located along posterior edge of mate maxillary palpomere narrowest and as
meso- and metafemora; with brushes of long long as two previous ones combined. Apices
setae at apices of all femora and very dis- of maxillary lobes (narrow galea and rather
tinct brush of very long setae along anterior wide lacinia) exposed from under labrum
edge of profemur; besides longitudinal rows and bearing extremely dense setae. First
of setae are visible on protibiae). Dorsum of antennal club segment somewhat dorsoven-
head and pronotum as well as prohypomera trally compressed. Pronotum widest at base
apparently without puncturation. Along and compressed at right posterior angle,
lateral edges of frons there are dense trans- however it was apparently not very con-
verse strips looking like strengthening and vex, its anterior edge strongly trapezium-
lightening of sclerotization. likely excised, its sides narrowly explanate
© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70
68 A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER
Antennal club of the specimen examined
represented by the basal wall of the first seg-
ment, but remains seem to be moved away.
Etymology. The epithet of the new spe-
cies refers to the mostly smooth and hairless
integument of dorsum of the specimen.
DISCUSSION
The composition and placement of the
family Hybosoridae are in great discussion
during the last decade. The most gene-
ralised viewpoint, which is close to the
traditional one, was formulated by Browne
and Scholtz (1999) and Howden and Gill
Fig. 3. Libanochrus calvus gen. et sp. nov., ho-
(2000) according to which Hybosoridae,
lotype. Head with exposed maxillary lobes (nar-
row galea and rather wide lacinia) and palpi, Ceratocanthidae, Ochodaeidae Mulsant &
dorsal view. Head width 2.6 mm. Rey, 1871 and Geotrupidae Latreille, 1802
can be treated as most relatives putting
the latter as closer to their common ances-
and with undulate edges, posterior angles tor. Krell (2006) after a review of the fossil
with distinct top, posterior edge apparently records also regards the Hybosoridae and
widely and evenly convex and with a slight Ceratocanthidae as sister groups with the
sinuation at posterior angles. All coxae ap- closest link also to Ochodaeidae and joining
parently (sub)contiguous; procoxae appar- all three families with Geotrupidae. Ocam-
ently suboval to subquadrangular, meso- po (2006) and Ocampo and Hawks (2006)
coxae suboval and metacoxae transverse. on molecular and morphological compari-
Protibia comparatively short (slightly son put Hybosoridae (including Ceratocan-
longer than head) and rather compressed, thidae) as a sister group of Ochodaeidae,
its outer edge with clear not quite regu- both of which are linked with Glaphyridae
lar and oblique crenulations from the base MacLeay, 1819. Nikolajev (2007) having
and between three more prominent teeth analysed some fossil families considers as
disposed with subequal distances between sister groups the Hybosoridae (including
them, the proximal tooth rather small. Spur Belohinidae Paulian, 1959) and Trogidae
of anterior legs very long (about as long MacLeay, 1819 and approaching both to
as protarsomeres 1 and 2 combined) and Geotrupidae and some subfamilies of Scara-
slightly curved. Profemora gently curved baeidae Latreille, 1802. The new species
along posterior edge and strongly subangu- demonstrates that the subfamily Liparo-
larly curved along the anterior one, about chrinae has a Mesozoic root in addition to
twice as wide as probibia. Left meso- and the Middle Jurassic – Cretaceous Anaidi-
metafemora are visible only from posterior nae Hybosorinae and Ceratocanthidae
side and they look like strongly compressed. (Krell, 2007; Ponomarenko & Kirejtshuk,
Protarsus rather thin and apparently about 2011; Nikolajev et al., 2010). On the other
three quarters as long as protibia; protarso- hand, the comparatively great representa-
mere 1 apparently longer than each of pro- tion of this family in fossils from the Asian
tarsomeres 2–4 and protarsomere 5 longest outcrops could be evidence that this family
(about as long as protarsomeres 1–4 com- was rather diverse on this continent during
bined; one claw available apparently about the late Mesozoic, compared with diversity
two fifths as long as protarsomere 5. of Geotrupidae, Ochodaeidae and Glaphy-
© 2011 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 20(1): 62–70
A.G. KIREJTSHUK ET AL. MESOZOIC LIPAROCHRINAE FROM LOWER CRETACEOUS LEBANESE AMBER 69
ridae beginning from the Cretaceous (Krell, Browne, J. & Scholtz, C.H. 1999. A phylogeny
2007; Kirejtshuk et al., 2010; Ponomarenko of the families of Scarabaeoidea. Systematic
& Kirejtshuk, 2011). Entomology, 24(1): 51–84.
Howden, H.F. & Gill, B.D. 2000. Tribes of New
World Ceratocanthinae, with keys to genera
ACKNOWLEDGEMENTS
and descriptions of new species (Coleoptera:
Scarabaeoidea). Sociobilogy, 35: 281–329.
The authors greatly appreciate the gener-
Kirejtshuk, A.G. & Azar, D. 2008. New taxa
ous assistance of A. Nel during the study. The
of beetles (Insecta, Coleoptera) from Leba-
two first co-authors have a pleasant duty to ex-
nese amber with evolutionary and systematic
press their recognition to the Muséum National
comments. Alavesia, 2: 15–46.
d’Histoire Naturelle for providing them a pos-
Kirejtshuk, A.G., Azar, D. Beaver, R.A., Man-
sibility to work in this museum as visiting pro-
delstam, M.Yu. & Nel, A. 2009a. The most
fessors. The study of the first co-author was also
ancient bark beetle known: a new tribe, ge-
supported by the Programme of the Presidium
nus and species from Lebanese amber (Cole-
of the Russian Academy of Sciences ‘Origin of
optera, Curculionidae, Scolytinae). System-
Biosphere and Evolution of Geo-Biological Sys-
atic Entomology, 34(1): 101–112.
tems’ and by a grant of the Russian Foundation
Kirejtshuk A.G., Azar D., Tafforeau P., Bois-
of Basic Research (09-04-00789). This paper is
tel R. & Fernandez V. 2009b. New beetles
a contribution to a team project ‘Biodiversity:
of Polyphaga (Coleoptera, Polyphaga) from
Origin, Structure, Evolution, and Geology’ al-
Lebanese amber. Denisia, 26 (Zugleich Kata-
lotted to the second co-author and his team by
loge der OberösterreichischenLandesmu-
the Lebanese University. An essential help was
seen, Neue Serie. 86): 119–130.
obtained from G.V. Nikolajev (Al-Farabi Kazakh
Kirejtshuk, A.G., Ponomarenko, A.G., Pro-
University, Almaty) and Wang Bo (Nanjing In-
kin, A.A., Chang, H., Nikolajev, G.V.,
stitute of Geology and Palaeontology of Chinese
Ren, D. 2010. Current knowledge on Me-
Academy of Sciences) provided with their re-
sozoic Coleoptera from Daohugou and Lia-
cent publications. The authors greatly appreci-
oning (North East China). Acta Geologica
ate valuable comments on an earlier version of
Sinica, 84 (4): 783–792.
the manuscript received from F.T. Krell (Denver
Krell, F.-T. 2006. Fossil Record and Evolution of
Museum of Nature and Science) and S. Hisamat-
Scarabaeoidea (Coleoptera: Polyphaga). Co-
su (Ehime University, Matsuyama).
leopterists Society Monograph, 5 (Supplement
to The Coleopterists Bulletin, 60): 120–143.
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