A peer-reviewed open-access journal ZooKeys 130: 229-237 (2011) Ree er tore Fee #ZooKeys www.zookeys.o rg Launched to accelerate biodiversity research 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 T urn:lsid:zoobank.org:author:7 CBE9A76-16C4-4D6D-9B4D-B2BECOO0ADDF * urn:lsid:zoobank.org:author: 18SAOB9F9-537A-46EF-B745-3942F6A5AB58 Corresponding author: Bo Wang (savantwang@gmail.com) Academic editor: D. Shcherbakov | Received 23 March 2011 | Accepted 28 April 2011 | Published 24 September 2011 urn:lsid:zoobank.org:pub:AISF2DA6-1DE3-4869-BOAD-FC23D85A9EE4 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 Abstract 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. Keywords Protorabinae, Coleoptera, Lower Cretaceous, Inner Mongolia, China Introduction 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, 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. 230 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 lagerstatte 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 (Corel DRAW 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 pronotum. Systematic Paleontology Family Carabidae Latreille, 1802 Subfamily Protorabinae Ponomarenko, 1977 Genus Cretorabus Ponomarenko, 1977 http://species-id.net/wiki/Cretorabus Type species. Cretorabus capitatus Ponomarenko, 1977; by original designation. Type horizon and locality. Zaza Formation, Lower Cretaceous; Baissa, Buryatiya, Russia. 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) ole e ee eseeseeeeeeeeeeeee 3 — Bodydlarge ts 3: Mim). succes conten nee cdiice aesseesalsenaesbatealauiaca Aa tuts, 2 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 3 Body small (length 3.7 mm); elytra with punctate fUrrowS........ eee eee cs Mad ok Wy C. sulcatus Ponomarenko, Coram & Jarzembowski, 2005 - Bodyimediimesizedssely tra ss OOUln tas vscecseten cane ese etseruae debe etershagac cuneate thas 4 4 Prosternal process narrow, 0.4 times as wide as procoxae, slightly longer than procoxdermictacexde 1: Citaesaswide As LON es Fo a.5inainnlenerderoseiaesemetrensselaens _eaeemrneior wees ecemmvmm ie boven Freie ir WEES. Ste RE A. C. capitatus Ponomarenko, 1977 = Prosternal process almost as wide as procoxae, much longer than procoxae; MICACOMAS Ze liMVestas WIE: ASNONG Maen ensue net noun a a antlnnellaphaaeat 5 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 POStEHOr- Marcin NOteMaAreiiaten 22 .s.e.2. Sa. ie saps adensaasaes C. rasnitsyni sp. n. 232 Bo Wang & Haichun Zhang/ ZooKeys 130: 229-237 (2011) Cretorabus rasnitsyni Wang & Zhang, sp. n. urn:lsid:zoobank.org:act:4416C789-0065-4CCC-9279-5F1419A1D481 http://species-id.net/wiki/Cretorabus_rasnitsyni Figs 1-5 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 subtruncate. 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 o ft. ‘ r f os 4 ea Figures 1-2. Cretorabus rasnitsyni sp. n., photomicrograph of holotype NIGP152464 | 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 Russian palaeoentomologist. 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- 234 Bo Wang & Haichun Zhang/ ZooKeys 130: 229-237 (2011) Figures 3-4. Cretorabus rasnitsyni sp. n., drawings of holotype NIGP152464 3 without legs 4 with legs. ginate. Furthermore, it is different from C. capitatus in having the comparatively wider prosternal process and metacoxae. Discussion Ponomarenko and Kirejtshuk (2011) have revised and summarized the fossil record of Protorabinae, but the Chinese data require further revisions. Two monotypic genera, Sinocarabus Hong, 1982 and Obesofemoria Hong, 1982 were described from the Lower Cretaceous of Yumen of Gansu by Hong (1982), and also recorded A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous... 235 maxillary palp mandible scape mentum . stipes submentum cardo gula 0.5 mm Figure 5. Cretorabus rasnitsyni sp. n., drawing of mouthparts of holotype NIGP 152464. in the list of fossil beetles by Ponomarenko and Kirejtshuk (2011). Sinocarabus leptoceroides Hong, 1982 was erected based on a specimen with the part and coun- terpart (Hong 1982). Judging from the original photograph (Hong 1982, plate 26, Figs 3, 4), this specimen seems to belong to Carabidae because of its body shape and strong mandibles. In the Hong’s original description, he stated that the “meta- coxae round” and “metacoxae distant from each other”. Therefore, this specimen is not a carabid beetle. Moreover, Tan et al. (2004) also pointed out that the original drawing of this species showed some clear characters of Polyphaga. Thus, the sys- tematic position of this specimen remains unclear. Obesofemoria Hong, 1982 was described based on a poorly-preserved specimen in dorsal aspect. This specimen does not show any diagnostic characters of Adephaga; thus more clear evidence is needed to resolve its taxonomic issue. Three specimens from the Lower Cretaceous Lushangfen Formation of Lushangfen of Beijing were transferred to Protorabinae by Ponomarenko and Kirejtshuk (2011): Penecupes rapax Ren, 1995, Aethocara- bus levigata Ren, 1995, and Nebrorabus tumoculus (Ren, 1995). As a result, only four species, including the species described herein, within Protorabinae have been reported from the Mesozoic strata of China (Table 1). The diversity of Chinese Protorabinae based on the present data probably has been underestimated. More investigations on this topic should be done to understand better the true diversity of Protorabinae. 236 Bo Wang & Haichun Zhang/ ZooKeys 130: 229-237 (2011) Table 1. Fossil record of Protorabinae. Taxon Lithorabus incertus Ponomarenko, 1977 Occurrence Lower Jurassic of Issyk-Kul, Kirghizia Protorabus planus Ponomarenko, 1977 Upper Jurassic of Karatau, Kazakhstan P. magnus Ponomarenko, 1977 P. nigrimonticola Ponomarenko, 1977 P. kobdoensis Ponomarenko, 1986 Upper Jurassic of Karatau, Kazakhstan Upper Jurassic of Karatau, Kazakhstan Lower Cretaceous of Myangad, Mongolia P crassus Ponomarenko, 1989 Lower Cretaceous of Shiviya, Transbaikalia P. tsaganensis Ponomarenko, 1989 Ovrabites ovalis Ponomarenko, 1977 O. jurassicus Ponomarenko, 1977 O. incertus Ponomarenko, 1993 Lower Cretaceous of Bon-Tsagan, Mongolia Upper Jurassic of Karatau, Kazakhstan Upper Jurassic of Karatau, Kazakhstan Lower Cretaceous of Khetana, Russian Far East Cordorabus notatus Ponomarenko, 1977 C. antennatus Ponomarenko, 1977 C. minimus Ponomarenko, 1977 C. vittatus Ponomarenko, 1980 Upper Jurassic of Karatau, Kazakhstan Upper Jurassic of Karatau, Kazakhstan Upper Jurassic of Karatau, Kazakhstan Lower Cretaceous of Manlay, Mongolia C. striatus Ponomarenko, 1986 Lower Cretaceous of Gurban-Ereney, Mongolia Cretorabus capitatus Ponomarenko, 1977 C. latus Ponomarenko, 1977 C. orientalis Ponomarenko, 1989 Lower Cretaceous of Baissa, Transbaikalia Lower Cretaceous of Baissa, Transbaikalia Lower Cretaceous of Khutel Khara, Mongolia C. ovalis Ponomarenko, 1989 Lower Cretaceous of Bon-Tsagan, Mongolia C. sulcatus Ponomarenko, Coram & Jarzembowski, 2005 C. rasnitsyni sp. n. Nebrorabus baculum Ponomarenko, 1989 Lower Cretaceous of Dorset, England Lower Cretaceous of Inner Mongolia, China Lower Cretaceous of Chernyshevsk, Transbaikalia N. capitatus Ponomarenko, 1989 Lower Cretaceous of Baley, Transbaikalia N. nebrioides Ponomarenko, 1989 N. 2elongatus (Ponomarenko, 1986) N. tumoculus (Ren, 1995) Lower Cretaceous of Bon-Tsagan, Mongolia Lower Cretaceous of Myangad, Mongolia Lower Cretaceous of Lushangfen, China Aethocarabus levigata Ren, 1995 Lower Cretaceous of Lushangfen, China Penecupes rapax Ren, 1995 Acknowledgements Lower Cretaceous of Lushangfen, China BW thanks to the staff members of the palaeoentomological laboratory of the Palae- ontological Institute (Russian Academy of Sciences) for their help during his visit in Moscow (2010). Many thanks go to Alexander Ponomarenko and Alexander Kirejt- shuk for reviewing the manuscript and providing constructive suggestions, Lin Jih- Pai for checking English language of the manuscript, and Dmitry Shcherbakov for his helpful editorial comments. This research was supported by the National Natural Science Foundation of China (Grant Nos. 40872015, 41002006, J0930006), Major Basic Research Program of China (Grant No. 2006CB806400). A new ground beetle (Carabidae, Protorabinae) from the Lower Cretaceous... 237 References Chang SC, Zhang HC, Renne PR, Fang Y (2009) High-precision “Ar/*’Ar age for the Jehol Biota. Palaeogeography, Palaeoclimatology, Palaeoecology 280: 94-104. doi: 10.1016/j. palaeo.2009.06.021 Gromov VP, Dmitriev VYu, Zherikhin VV, Lebedev EL, Ponomarenko AG, Rasnitsyn AP, Sukacheva ID (1993) Cretaceous insect faunas from Uljya River basin (West Okhotsk region). Trudy Paleontologicheskogo Instituta Rossiyskoi Akademii Nauk 252: 5-60. 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