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A Remarkable New Genus of the Snakefly Family Mesoraphidiidae

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A Remarkable New Genus of the Snakefly Family Mesoraphidiidae Cretaceous Research 89 (2018) 119e125 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes Short communication A remarkable new genus of the snakefly family Mesoraphidiidae (Insecta: Raphidioptera) from the Lower Cretaceous of China, with description of a new species ** * Ya-nan Lyu a, Dong Ren b, , Xingyue Liu a, a Department of Entomology, China Agricultural University, Beijing 100193, China b College of Life Sciences, Capital Normal University, Beijing 100048, China article info abstract Article history: A fossil snakefly species, Mesoraphidia obliquivenatica (Ren, 1994), from the Lower Cretaceous (upper Received 20 October 2017 Barremian) of the Yixian Formation in Liaoning Province, China was discovered to possess an extremely Received in revised form prolonged occiput, which is a remarkable feature previously unknown in snakeflies. Based primarily on 21 January 2018 this feature, a new genus of the family Mesoraphidiidae, namely Stenoraphidia gen. nov., is erected to Accepted in revised form 26 February 2018 contain this species. In addition, a second and new species of Stenoraphidia gen. nov., i.e. Stenoraphidia Available online 27 February 2018 longioccipitalis sp. nov., is described from the same deposit. A summary of the morphological diversifi- cation of head shapes in snakeflies is given. Keywords: © Mesoraphidiinae 2018 Elsevier Ltd. All rights reserved. Taxonomy Head shape Yixian Formation Mesozoic 1. Introduction and MP, as well as fewer crossveins among these veins. However, many morphological characters, which are significant for dis- The extinct raphidiopteran family Mesoraphidiidae is the most tinguishing extant snakefly families, greatly vary within Meso- species-rich snakefly group from the Mesozoic era. It has a long raphidiidae, e.g. the presence/absence of ocelli (Liu et al., 2016) and evolutionary history ranging from the Middle Jurassic to the Late the pterostigmal crossvein (Martynov, 1925). This phenomenon Cretaceous (Engel, 2002; Jepson and Jarzembowski, 2008; Lü et al., prevents clear familial assignment of a number of genera to Mes- 2015; Makarkin and Khramov, 2015). Mesoraphidiidae is particu- oraphidiidae. In addition, some congeneric species described from larly diverse in the Early-Mid Cretaceous, with various morpho- the same deposit may represent different genera due to highly logically remarkable species. Nevertheless, the monophyly of this divergent non-wing characters; conversely other taxa are probably family remains unconfirmed due to a lack of conclusive support and synonyms since they lack solid diagnostic characters and show only convincing autapomorphies, despite previous phylogenetic studies subtle venational differences (Ponomarenko, 1988, 1993; Ren, 1994, (Willmann, 1994; Bechly and Wolf-Schwenninger, 2011; Liu et al., 1997). To date, the majority of mesoraphidiid species, as with other 2014). fossil snakefly families similarly preserved as compression fossils, Until now, Mesoraphidiidae consisted of two subfamilies, i.e. are principally diagnosed by wing characters. Exploration of other Alloraphidiinae and Mesoraphidiinae, presently including 27 diagnostic characters, other than wing venation, is important to genera and 69 species (Oswald, 2017). In general, compared with further clarify the paleodiversity of the Mesozoic Raphidioptera. the other major snakefly group, i.e. the Mesozoic Baissopteridae, In this study, we report on a group of Mesoraphidiidae from the the wings of Mesoraphidiidae contain far fewer branches of RP, MA Lower Cretaceous of Yixian Formation of China that contains more than 20 previously unidentified fossils with an exceptionally elongate occiput. This group represents a new genus, namely Stenoraphidia gen. nov. and comprises two species of the subfamily * Corresponding author. ** Corresponding author. Mesoraphidiinae: S. obliquivenatica (Ren, 1994) comb. nov. and a E-mail addresses: [email protected] (D. Ren), [email protected] new species S. longioccipitalis sp. nov. Additionally, a brief (X. Liu). https://doi.org/10.1016/j.cretres.2018.02.018 0195-6671/© 2018 Elsevier Ltd. All rights reserved. 120 Y.-n. Lyu et al. / Cretaceous Research 89 (2018) 119e125 comparison of the shape of the head and pronotum in Raphi- to be a stable feature of the genus. It is noteworthy that Meso- dioptera is provided. raphidia shangyuanensis (Ren, 1994), which is also from the Yixian Formation, might also possess such a prolonged occiput; however, 2. Material and methods the occiput was probably mistakenly considered as part of the prothorax, since the prothorax of snakeflies cannot bend medially All fossil specimens herein examined are deposited in the Key as illustrated in the original description (see Ren, 1994: fig. 5). As Laboratory of Insect Evolution and Environmental Changes, College such, M. shangyuanensis may be a member of Stenoraphidia gen. of Life Sciences, Capital Normal University (CNU), Beijing. They are nov. and possibly even a synonym of S. obliquivenatica. Until the from the upper Barremian (ca. 126.1 ± 1.7e124.6 ± 0.1 Ma), Lower holotype of M. shangyuanensis can be located and examined, Cretaceous of Jianshangou Member (Bed), Yixian Formation (Wang further taxonomic treatment on this species remains on hold. and Zhou, 2008; Walker et al., 2013), the Huangbanjigou locality (413604400N, 1204904800E) (see Wang et al., 2015: fig. 1), Sihetun Etymology. The generic name is a combination of the Greek noun area, Beipiao City, Liaoning Province, China. The deposit has yielded stenosis, narrowing and Raphidia, a common genus-group name for abundant and exquisitely preserved fossil insects (Ren, 1998; Gao Raphidioptera, in reference to the narrowly elongate occiput. et al., 2013; Wang et al., 2013; Yao et al., 2014). Specimens were Gender: feminine. examined using a Zeiss Discovery V12 stereo microscope, photo- Stenoraphidia obliquivenatica (Ren, 1994), comb. nov. graphs were taken with a Nikon D800 digital camera, and drawings (Fig. 1) were made in Adobe Photoshop CC. The terminology of wing venation generally follows Aspock€ Alloraphidia obliquivenatica Ren, 1994: 134. et al. (1991) and Kukalova-Peck and Lawrence (2004). Abbrevia- tions used for wing veins and spaces are: A, anal; ac, anal cell; C, Revised diagnosis. Large-sized snakeflies (body length 14.1e23.0 mm, costa; Cu, cubitus; CuA, cubitus anterior; CuP, cubitus posterior; dc, forewing length 13.8e16.9 mm). Head elongate, dark-colored, with discal cell; doi, discoidal cell; M, media; MA, media anterior; MP, an extremely prolonged occiput nearly half the length of the head. media posterior; m, medial cell; pt, pterostigma; R, radius; RA, Prothorax elongate, nearly twice as long as wide, anterior half pale- radius anterior; RP, radius posterior; r, radial cell; ScP, subcosta colored, but posterior half dark-colored. Legs slender, uniformly posterior. dark-colored. Pterostigma long, uniformly colored, with an oblique All taxonomic acts established in the present work have been pterostigmal crossvein at subdistal portion of pterostigma (Fig. 1F), registered in ZooBank (see below), together with the electronic its width clearly increases at distal portion; RP with three to four publication under urn:lsid:zoobank.org:pub:14C8F8C9-7843- pectinate branches; MA with three branches. 4E0F-952C-B25DA8B25365. Materials examined. CNU-RAP-LB-2017032c/p, a well preserved fe- male specimen compressed laterally; CNU-RAP-LB-2017079, a well 3. Systematic palaeontology preserved female specimen compressed laterally; CNU-RAP-LB- 2017081, an incomplete specimen (sex unknown) compressed Order Raphidioptera Navas, 1916 laterally, without forewings and apex of abdomen; CNU-RAP-LB- Suborder Raphidiomorpha Engel, 2002 2017082, a poorly preserved female specimen compressed dorso- Family Mesoraphidiidae Martynov, 1925 ventrally; CNU-RAP-LB-2017083, a well preserved female specimen Subfamily Mesoraphidiinae Martynov, 1925 compressed laterally; CNU-RAP-LB-2017084c/p, a well preserved female specimen compressed dorsoventrally; CNU-RAP-LB- Genus Stenoraphidia gen. nov. 2017085, a well preserved male specimen compressed laterally; urn:lsid:zoobank.org:act:511A1CE0-ADD5-4D0D-B3E5- CNU-RAP-LB-2017087, an incomplete female specimen compressed C91BEF88B9FF laterally, with head and prothorax not preserved; CNU-RAP-LB- Type species. Alloraphidia obliquivenatica Ren, 1994; the Lower 2017088, a well preserved male specimen compressed dorsoven- Cretaceous of Yixian Formation of China. trally, with anterior portion of head badly preserved; CNU-RAP-LB- Diagnosis. Head elongate, with an extremely prolonged occiput 2017089, an incomplete specimen (sex unknown) compressed nearly half the length of the head. Ocelli present. Prothoraxelongate, laterally, with head, prothorax and apex of abdomen not preserved; nearly twice as long as wide. Legs slender. Pterostigma long, uni- CNU-RAP-LB-2017092c/p, a well preserved male specimen com- formly colored; an oblique pterostigmal crossvein usually present at pressed laterally; CNU-RAP-LB-2017093, a well preserved female subdistal portion of pterostigma; RP with three or four pectinate specimen compressed laterally; CNU-RAP-LB-2017094, an incom- branches; MA with two or three branches. Forewing: MA with stem plete specimen (sex unknown) compressed dorsoventrally, with originating near MP branching point, three discoidal cells present. hind wings and apex of abdomen not preserved; CNU-RAP-LB- Remarks. The new genus is one of the most distinguishable
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