A Remarkable New Genus of the Snakefly Family Mesoraphidiidae

Cretaceous Research 89 (2018) 119e125

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Cretaceous Research

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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 diversification 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 female 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 2017095, a poorly preserved male specimen compressed laterally; snakeflies due to the narrowly prolonged occiput. Conversely, in all CNU-RAP-LB-2017096, an incomplete specimen (sex unknown) other known extinct or extant snakeflies, the occiput is much compressed laterally, apex of abdomen and wings not preserved. shorter or not visible. Owing to the failure to consider the character Remarks. With respect to body size (forewing length ca. of the distinctive occiput, S. obliquivenatica was previously assigned 13.0e17.0 mm) and wing venation, S. obliquivenatica resembles to Mesoraphidia Martynov, 1925, based on the similar wing vena- some Mesoraphidia species described from the Yixian Formation, tion as Mesoraphidia. However, it differs from the latter genus in i.e. M. longistigmosa (Ren, 1994), M. shangyuanensis (Ren, 1994) having the exceptionally narrow and prolonged occiput and the and M. polyphlebia (Ren, 1994). However, the latter three species elongate prothorax, while M. grandis and a number of species are barely distinguishable from each other and lack convincing currently placed in Mesoraphidia possess a short prothorax. diagnostic characters. Moreover, these species may even be conspecific with S. obliquivenatica. The following characters previ- The head shape of snakeflies is a useful diagnostic character for ously used for distinguishing these species require reconside- distinguishing genera, but it is often overlooked or not preserved in ration, i.e. the pterostigmal crossvein (oblique in S. obliquivenatica, fossil snakeflies. Based on our examination of more than 20 fossils unclear in M. polyphlebia, but vertical in M. shangyuanensis and of Stenoraphidia gen. nov., the narrowly prolonged occiput appears M. longistigmosa), and the number of RA veinlets and RP branches in Y.-n. Lyu et al. / Cretaceous Research 89 (2018) 119e125 121

Fig. 1. Stenoraphidia obliquivenatica (Ren, 1994), comb. nov. AeG Photographs, HeI Drawings. A. Habitus of CNU-RAP-LB-2017079, female, lateral view; B. Habitus of CNU-RAP-LB- 2017083, female, lateral view; C. Photo of head and prothorax of CNU-RAP-LB-2017094; D. Photo of head and prothorax of CNU-RAP-LB-2017085; E. Pterostigma of forewing of CNU- RAP-LB-2017094; F. Pterostigma of forewing of CNU-RAP-LB-2017081; G. Photo of head and prothorax of CNU-RAP-LB-2017095; H. Wing venation of CNU-RAP-LB-2017079; I. Wing venation of CNU-RAP-LB-2017083. Scale bar ¼ 2.0 mm. Arrow in F indicates pterostigmal crossvein.

both fore- and hind wings (four RA veinlets and five RP branches M. shangyuanensis and M. longistigmosa, and the four RA veinlets present in M. polyphlebia, but three RA veinlets and four RP and five RP branches that are present in M. polyphlebia. Therefore, it branches present in S. obliquivenatica, M. shangyuanensis, and is more appropriate to identify these additional fossils as M. longistigmosa). Unfortunately, the holotypes of the four above S. obliquivenatica. Whether M. shangyuanensis, M. longistigmosa and mentioned species are presently unavailable. Nevertheless, the M. polyphlebia belong to Styporaphidia gen. nov. is a question specimens of S. obliquivenatica that we examined are nearly iden- requiring further consideration upon future examination of the tical to the original description and illustration of this species. types, as well as additional fossils of these species. Among all the examined materials from the same desposit, we did Stenoraphidia longioccipitalis sp. nov. not find the vertical pterostigmal crossvein that is present in (Fig. 2) 122 Y.-n. Lyu et al. / Cretaceous Research 89 (2018) 119e125 urn:lsid:zoobank.org:act:F2D6A6A3-B5A3-4872-B026- Abdomen 3.2 mm long, 1.7 mm wide at maximum in lateral F4D94A6606FD view; color of dorsal part of abdomen paler than venter; ovipositor strong, 4.7 mm long. Type materials. Holotype: CNU-RAP-LB-2017071, a well preserved female specimen compressed laterally. Paratypes: CNU-RAP-LB- Description of paratypes. Body length 10.1e11.4 mm (measured 2017141, a well preserved female specimen compressed laterally; without ovipositor). Head length 2.8e3.4 mm. Antenna partially CNU-RAP-LB-2017142, a well preserved female specimen com- preserved, more than 3.2 mm long, with at least 34 flagellomeres. pressed dorsoventrally; CNU-RAP-LB-2017144, a poorly preserved Prothorax length 1.6e2.2 mm. Meso- plus metathorax combined specimen (sex unknown) compressed laterally, without abdomen; length 1.6e2.6 mm. Legs slender. Forewing length 8.2e9.3 mm CNU-RAP-LB-2017145, a well preserved specimen (sex unknown) long, maximum width 2.3e2.4 mm. Hind wing length 8.2 mm. compressed laterally; CNU-RAP-LB-2017146, a poorly preserved Abdomen length 4.0e4.5 mm (measured without ovipositor). A specimen (sex unknown) compressed laterally. weak oblique pterostigmal crossvein present at subdistal portion of Locality and horizon. The Yixian Formation (upper Barremian, Lower pterostigma. Ovipositor incomplete. Configuration of body and Cretaceous), Huangbanjigou locality (413604400N, 1204904800E), wings same to that of holotype female. Beipiao city, Liaoning Province, northeastern China. Etymology. The specific epithet longioccipitalis is a combination of Male. Unknown. the Latin adjective longus and the Latin noun occipitalis, referring to the prolonged occiput. Remarks. The new species can be distinguished from Diagnosis. Medium-sized snakeflies (body length 10.1e11.4 mm, S. obliquivenatica by the relatively small body-size (body length forewing length 8.2e9.3 mm). Head elongate, dark-colored, with an 10.1e11.4 mm, forewing length 8.2e9.3 mm) [body-size large in extremely prolonged occiput nearly half the length of the head. S. obliquivenatica (body length 14.1e23.0 mm, forewing length Prothorax elongate, pale-colored throughout. In both fore- and 13.8e16.9 mm)] and the uniformly pale-colored pronotum (Fig. 2A, hind wings, uniformly colored pterostigma long, with an oblique B, E, F) [pronotum bicolored in S. obliquivenatica (Fig. 1C, D, G)]. pterostigmal crossvein; RP with three branches; MA with two simple branches. Description of holotype female. Body 10.1 mm long. Head elongate, 3.0 mm long, dark-colored, with occiput nearly half of head length. 4. Discussion Preserved antenna with more than 23 flagellomeres. Prothorax rectangular in lateral view, 1.9 mm long, nearly twice as long as The present description of a new genus, Stenoraphidia gen. nov., wide, pale-colored. Meso- and metathorax combined 2.5 mm long. highlights the morphological diversification of the head in snake- Legs slender, uniformly dark-colored. flies. Four types of head shapes (Fig. 3) are recognized in the various lineages of extinct and present-day Raphidioptera. 1) Ovoid head Forewing: 9.3 mm long, 2.4 mm wide. Costal space with four with short vertex and short occiput (Fig. 3A) (only known in the crossveins preserved; ScP terminating into costal margin nearly at mesoraphidiid tribe Nanoraphidiini) (Grimaldi, 2000; Engel, 2002; midpoint of wing; two RA veinlets present; pterostigma uniformly Bechly and Wolf-Schwenninger, 2011; Liu et al., 2016). 2) Ovoid colored, starting beyond termination of ScP, probably closed by a head with prolonged vertex and short occiput (Fig. 3B, D) (known in crossvein proximally, distally ended at anterior veinlet of RA; Mesoraphidiidae, Baissopteridae, Raphidiidae, and Juroraphidiidae) pterostigmal crossvein invisible; two radial crossveins present be- (Martynova, 1961; Aspock€ et al., 1991; Liu et al., 2014). 3) Rectan- tween RA and RP,1r nearly as long as 2r; 2ra-rp crossvein connected gular head with prolonged vertex and short occiput (Fig. 3C) posteriorly to the anterior branch of RP; RP forked nearly at its (known in Mesoraphidiidae, Baissopteridae, Raphidiidae and Ino- midpoint, with a bifurcate anterior branch and a simple posterior celliidae) (Martynov, 1925; Aspock€ et al., 1991; Ponomarenko, 1993; branch; a single rp-ma crossvein connected posteriorly to the Lyu et al., 2017). 4) Elongate head with prolonged vertex and an anterior branch of MA, forming a discal cell; MA with stem origi- extremely prolonged occiput (Figs. 1, 2, 3E) (only known in the nating near first branching point of MP; two crossveins present mesoraphidiid genus Stenoraphidia gen. nov.). between MA and MP, forming two short medial cells; MP deeply Unfortunately, the head shape in Chrysoraphidiidae, Priscae- forked, with six terminal branches and three discoidal cells; CuA nigmatidae and Metaraphidiidae remain unknown. The juror- and CuP simple; two cua-cup crossveins present; A1 simple and A2 aphidiid species has a common ovoid head with prolonged vertex bifurcate. but short occiput. The head of the second predominant extinct Hind wing: 8.4 mm long, 2.3 mm wide. Costal space with four group, Baissopteridae, exhibits two main types, i.e., an ovoid head crossveins preserved; ScP terminating into costal margin near with prolonged vertex but short occiput, and a rectangular head midpoint of wing; two simple RA veinlets present; pterostigma with prolonged vertex but short occiput (see Lyu et al., 2017: fig. 1). uniformly colored, starting beyond termination of ScP, proximally The head of the Mesoraphidiidae displays a rich diversity of all four closed by a crossvein, distally ended at anterior veinlet of RA; types herein recognized in snakeflies. Except for the tribe Nano- pterostigmal crossvein absent; three radial crossveins present raphidiini with a short vertex, most mesoraphidiid species exhibit between RA and RP, 3r nearly as long as 1rþ2r; 1ra-rp crossvein an ovoid head with prolonged vertex and short occiput. Never- connected posteriorly to the stem of RP þ MA; 3ra-rp crossvein theless, some mesoraphidiid species, such as, M. grandis Martynov, connected posteriorly to the anterior branch of RP; RP forked 1925, M. glossophylla (Ren, 1997), possess a rectangular head (see nearly at its midpoint, with a bifurcate anterior branch and a Martynov, 1925: fig. 1; Ren, 1997: fig. 11). Most notably, in Stenor- simple posterior branch; a single rp-ma crossvein connected aphidia gen. nov. the occiput is narrowly prolonged. The head in the posteriorly to the first bifurcation of MA, forming a discal cell; MA majority of the extant group Raphidiidae is the common ovoid type stem diverging from R distad diverging point between MP and R, with prolonged vertex but short occiput, while in a few raphidiid with two simple branches; two crossveins present between MA species it is rectangular (see Aspock€ et al., 1991: fig. 180e199). All and MP, forming a long medial cell and a short one; MP deeply inocelliid species presently known have a rectangular head. forked, with five terminal branches and a discoidal cell; CuA The combination of the variously shaped prothoraces and head bifurcate and CuP simple. (i.e., general shape, vertex and occiput) that are characteristic of Y.-n. Lyu et al. / Cretaceous Research 89 (2018) 119e125 123

Fig. 2. Stenoraphidia longioccipitalis sp. nov. AeF Photographs, GeH Drawings. A. Habitus photo of holotype CNU-RAP-LB-2017071, female, lateral view; B. Habitus photo of paratype CNU-RAP-LB-2017141, female, lateral view; C. Photo of head and prothorax of holotype CNU-RAP-LB-2017071, lateral view; D. Photo of head of paratype CNU-RAP-LB-2017142, dorsal view; E. Habitus photo of paratype CNU-RAP-LB-2017142, female, dorsal view; F. Habitus photo of paratype CNU-RAP-LB-2017145, sex unknown, lateral view; G. Draw- ing of wing venation of holotype CNU-RAP-LB-2017071; H. Drawing of wing venation of paratype CNU-RAP-LB-2017141. Scale bar ¼ 2.0 mm. 124 Y.-n. Lyu et al. / Cretaceous Research 89 (2018) 119e125

5. Concluding remarks

The present finding of new fossil snakeflies with extremely prolonged occiput increases our knowledge on the morphology and palaeodiversity of Raphidioptera. The new genus Stenoraphidia gen. nov. with its remarkably narrow and prolonged occiput and elongate prothorax is highly interesting since the specialized head and prothorax could be adaptive traits related to feeding or hunting behaviors. Aside from using wing venational characters, it is important to explore and evaluate more non-wing characters for the identification of fossil snakeflies and the reconstruction of their phylogenetic relationships.

Acknowledgements

This research was supported by the National Natural Science Foundation of China (Nos. 31672322, 31672323, 31322501 and 41688103 and 31730087), the Beijing Natural Science Foundation (No. 5162016), and the Chinese Universities Scientiﬁc Fund (No. 2017TC031), Project of High-level Teachers in Beijing Municipal Universities (IDHT20180518) and Program for Changjiang Scholars and Innovative Research Team in University (IRT-17R75). We thank Dr. Davide Badano and an anonymous reviewer for their valuable comments which improved the manuscript, and Dr. John D. Plant for checking the English.

References

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