A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous of Inner Mongolia, China.
ABSTRACT Cretorabus rasnitsynisp. n., belonging to the extinct subfamily Protorabinae of Carabidae, was described based on a well-preserved specimen from the Lower Cretaceous Yixian Formation of Yangshuwanzi, Inner Mongolia. The diagnostic characters forCretorabus are revised, and the key to species of the genus was presented. The fossil record of Protorabinae was summarized. Sinocarabus Hong, 1982 and Obesofemoria Hong, 1982 cannot be attributed to Protorabinae.
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A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous...
A new ground beetle (Carabidae, Protorabinae) from
the Lower Cretaceous of Inner Mongolia, China
Bo Wang†, Haichun Zhang‡
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chi-
nese Academy of Sciences, 39 East Beijing Rd., Nanjing 210008, China
Corresponding author: Bo Wang (email@example.com)
Academic editor: D. Shcherbakov | Received 23 March 2011 | Accepted 28 April 2011 | Published 24 September 2011
Citation: Wang B, Zhang H (2011) A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous of Inner
Mongolia, China. In: Shcherbakov DE, Engel MS, Sharkey MJ (Eds) Advances in the Systematics of Fossil and Modern
Insects: Honouring Alexandr Rasnitsyn. ZooKeys 130: 229–237. doi: 10.3897/zookeys.130.1300
Cretorabus rasnitsyni sp. n., belonging to the extinct subfamily Protorabinae of Carabidae, was described
based on a well-preserved specimen from the Lower Cretaceous Yixian Formation of Yangshuwanzi, Inner
Mongolia. The diagnostic characters for Cretorabus are revised, and the key to species of the genus was
presented. The fossil record of Protorabinae was summarized. Sinocarabus Hong, 1982 and Obesofemoria
Hong, 1982 cannot be attributed to Protorabinae.
Protorabinae, Coleoptera, Lower Cretaceous, Inner Mongolia, China
Protorabinae is an extinct subfamily of Carabidae, and differs remarkably from all
other carabid beetles in that the metepisterna are extending to the mesocoxal cavi-
ties (Ponomarenko 1977). The earliest Protorabinae, Lithorabus incertus Ponomarenko,
ZooKeys 130: 229–237 (2011)
Copyright Bo Wang, Haichun Zhang. This is an open access article distributed under the terms of the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Bo Wang & Haichun Zhang / ZooKeys 130: 229–237 (2011)
1977, was described from the Lower Jurassic of Issyk-Kul, Kirghizia (Ponomarenko
1977). Up to date, nearly 30 species within 8 genera are known from the Jurassic of
Central Asia and the Lower Cretaceous of Mongolia, Russia, China and UK (Gromov
et al. 1993; Ponomarenko 1977, 1980, 1986, 1989; Ponomarenko et al. 2005; Ren
1995). However, some Chinese Protorabinae require more detailed descriptions and
revisions. The famous Early Cretaceous Jehol Biota yields abundant, well-preserved
insects, including beetles. However, no record of Carabidae was reported from this
lagerstätte until now. Herein, a new species of Protorabinae is described based on a
well-preserved beetle from the Yixian Formation of Inner Mongolia, and the current
fossil record of Protorabinae is summarized herein.
Material and methods
The specimen (NIGP152464) was from the Lower Cretaceous Yixian Formation of
Yangshuwanzi Village, Bisiyingzi Township, Ningcheng County, Chifeng City, Inner
Mongolia. The fossils from Yangshuwanzi are commonly preserved in yellow silty mud-
stone. The horizon in this locality is equivalent to either the Jianshangou or Dawang-
zhangzi bed (about 124–122 Ma) of the Yixian Formation (Chang et al. 2009). The
coleopteran assemblages from this locality were dominated by a variety of scarabs.
The specimen was examined dry and under alcohol, using a Nikon SMZ1000
stereomicroscope and drawings were made with the aid of a camera lucida. The pho-
tographs were prepared using a digital camera (DXM1200) connected to the above
stereomicroscope, and the line drawings were readjusted on photographs using im-
age-editing software (CorelDRAW X4 and Adobe Photoshop CS). In drawings, the
broken lines denote the hidden and presumably missing body parts. Body length
was measured along the midline from the anterior margin of frons to apex of elytra,
and width was measured across the broadest part of elytra. The length of pronotum
was measured along the midline; the width was measured across the broadest part of
Family Carabidae Latreille, 1802
Subfamily Protorabinae Ponomarenko, 1977
Genus Cretorabus Ponomarenko, 1977
Type species. Cretorabus capitatus Ponomarenko, 1977; by original designation.
Type horizon and locality. Zaza Formation, Lower Cretaceous; Baissa, Buryatiya,
A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous... 231
Diagnosis. Body wide, small or medium-sized. Head large, strongly transverse.
Pronotum transverse, widest in anterior or middle portion, constricted behind mid-
dle. Mesoventrite longer than mesocoxae. Metepisterna posteriorly tapering. Meta-
coxal plates tapering strongly in lateral half, extending as a narrow tongue up to lateral
margins of metacoxae. Abdomen short, with rounded apex; last ventrite long, its ante-
rior margin two-thirds narrower than base of abdomen. Legs short, femora slightly ex-
tending beyond body sides. Elytra smooth or with numerous rows of large punctures.
Remarks. The genus is different from other genera in the metacoxal plates ta-
pering strongly in lateral half, extending as a narrow tongue up to the lateral mar-
gins of metacoxae. Furthermore, it differs from Cordorabus Ponomarenko, 1977
in having the mesoventrite longer than mesocoxae, and abdomen short, with apex
rounded; from Ovrabites Ponomarenko, 1977 in possessing the pronotum con-
stricted behind middle, abdomen short, with apex rounded; from Protorabus Pon-
omarenko, 1977 by the last ventrite much narrower than base of abdomen, and
elytra with grooves; from Lithorabus Ponomarenko, 1977 in having the metepis-
terna posteriorly tapering; from Nebrorabus Ponomarenko, 1989 in possessing the
wider body and shorter legs.
Species included. Six species: C. capitatus Ponomarenko, 1977 and C. latus Pon-
omarenko, 1977 from the Lower Cretaceous of Baissa; C. orientalis Ponomarenko,
1989 from the Lower Cretaceous of Khutel Khara of Mongolia; C. ovalis Ponomaren-
ko, 1989 from the Lower Cretaceous of Bon-Tsagan of Mongolia; C. sulcatus Pon-
omarenko, Coram & Jarzembowski, 2005 from the Lower Cretaceous Purbeck Lime-
stone Group of England; and C. rasnitsyni sp. n. from the Lower Cretaceous Yixian
Formation of Inner Mongolia of China.
Key to species of Cretorabus
Body small or medium-sized (length < 10 mm) ..........................................3
Body large ( > 13 mm) ................................................................................2
Metacoxae 1.8 times as wide as long ..............C. latus Ponomarenko, 1977
Metacoxae 2.5 times as wide as long ............C. ovalis Ponomarenko, 1989
Body small (length 3.7 mm); elytra with punctate furrows ............................
........................C. sulcatus Ponomarenko, Coram & Jarzembowski, 2005
Body medium-sized; elytra smooth .............................................................4
Prosternal process narrow, 0.4 times as wide as procoxae, slightly longer than
procoxae; metacoxae 1.5 times as wide as long ..............................................
...............................................................C. capitatus Ponomarenko, 1977
Prosternal process almost as wide as procoxae, much longer than procoxae;
metacoxae 2.5 times as wide as long ............................................................5
Metaventrite twice as wide as long; metacoxal plates with emargination in
lateral part of posterior margin ................C. orientalis Ponomarenko, 1989
Metaventrite 3 times as wide as long; metacoxal plates with lateral part of
posterior margin not emarginate .....................................C. rasnitsyni sp. n.
Bo Wang & Haichun Zhang / ZooKeys 130: 229–237 (2011)
Cretorabus rasnitsyni Wang & Zhang, sp. n.
Holotype. NIGP152464, male, a well-preserved beetle in ventral aspect. Yixian For-
mation, Lower Cretaceous; a fossil locality (41°25'N, 118°57'E) near Yangshuwanzi
Village, Bisiyingzi Township, Ningcheng County, Chifeng City, Inner Mongolia,
China. Deposited in the Nanjing Institute of Geology and Palaeontology (NIGP),
Chinese Academy of Sciences.
Diagnosis. Body medium-sized. Pronotum 1.8 times as wide as long. Prosternal
process almost as wide as procoxae, much longer than procoxae. Metaventrite 3 times
as wide as long. Metacoxae 1.8 times as wide as long. Metacoxal plates with lateral part
of posterior margin not emarginate.
Description. Head and pronotum strongly transverse. Length of head (including
mandibles) slightly shorter than occiput width. Head capsule narrowing anteriorly
from base. Eyes shorter than temples. Gular plate 1.5 times as long as wide, wid-
ened anteriorly. Antennae inserted at anterior margin of eyes; scape dilated; pedicel-
lum slightly shorter than scape; first flagellomere slightly longer than scape. Mandi-
bles large, slightly incurved, asymmetrical, with retinaculum small. Maxillary palps
4-segmented, conspicuously longer than mandibles; palpomere 4 with apical margin
Pronotum 1.8 times as wide as long, widest nearly in the middle; pronotal angles
pointed, concealing the base of head dorsally. Propleura narrow. Prosternum before
procoxae 1.4 times longer than procoxae. Prosternal process wide, 1.4 times as long
as procoxae. Mesoventrite transverse, with subtriangular depression for reception
of prosternal process. Mesepisternum almost rectangular, transverse. Mesepimeron
short, extending to mesocoxal cavities, slightly widened laterally. Metaventrite short,
3 times as wide as long; its anterior margin half as long as the posterior. Metepis-
terna subtriangular, gradually widened anteriorly, its length 1.5 times width at ante-
rior margin. Metacoxae oblique, slightly projecting over abdomen, 3 times as wide
as long. Metacoxal plates slightly longer than coxae. Abdomen a little longer than
meso- and metathorax combined, widened from base to apex of second visible ster-
nite, then narrowing. Parameres rather long, with penis slightly projecting.
Profemur and protibia widened apically, almost equal in length. Tarsal segments
widened apically; tarsomeres 1 almost as long as tarsomeres 2; tarsomeres 2 slightly
longer than tarsomeres 3. Mesofemur and mesotibia almost equal in length. Mesotibia
widened apically, 1.2 times as long as protibia. Metatrochanters one-third as long as
metafemora. Metatibiae 1.1 times as long as metafemora, slightly widened apically; its
outer side possibly with pits.
Measurements. Body length 7.8 mm, width 4.0 mm. Head length (including
mandibles) 1.9 mm, occiput width 2.6 mm; mandible length 0.8 mm. Pronotum:
A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous... 233
Figures 1–2. Cretorabus rasnitsyni sp. n., photomicrograph of holotype NIGP152464 1 dry 2 under
alcohol. Scale bar represents 1 mm.
length 1.8 mm, maximal width 3.2 mm. Abdomen: length 3.3 mm. Fore legs: femur
length 1.5 mm; tibia length 1.5 mm. Middle legs: femur length 1.7 mm; tibia length
1.7 mm. Hind legs: femur length 2.0 mm; tibia length 2.3 mm.
Etymology. Specific epithet is devoted to Alexander P. Rasnitsyn, an outstanding
Remarks. This specimens can be undoubtedly attributed to Cretorabus by the fol-
lowing characters: the pronotum constricted behind middle; mesoventrite longer than
mesocoxae; metacoxal plates tapering strongly in lateral half, extending as a narrow
tongue up to the lateral margins of metacoxae; and abdomen short, with rounded
apex. The species mostly resembles C. orientalis in having the prosternal process much
longer than procoxae, and metacoxae 2.5 times as wide as long, but differs from the
latter in possessing the metacoxal plates with the lateral part of the margin not emar-