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Hemiptera: Cicadomorpha) from the Middle Jurassic of Daohugou, Inner Mongolia, China

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Hemiptera: Cicadomorpha) from the Middle Jurassic of Daohugou, Inner Mongolia, China Eur. J. Entomol. 112(2): 373–380, 2015 doi: 10.14411/eje.2015.044 ISSN 1210-5759 (print), 1802-8829 (online) New fossil Procercopidae (Hemiptera: Cicadomorpha) from the Middle Jurassic of Daohugou, Inner Mongolia, China JUN CHEN 1, BO WANG 2, 3, HAICHUN ZHANG 2, XIAOLI WANG 1 and XIAOTING ZHENG 1 1 Institute of Geology and Paleontology, Linyi University, Shuangling Rd., Linyi 276000, China; e-mails: [email protected]; [email protected]; [email protected] 2 State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, East Beijing Rd., Nanjing 210008, China; e-mails: [email protected]; [email protected] 3 Steinmann Institute, University of Bonn, 53115 Bonn, Germany Key words. Hemiptera, Cicadomorpha, Procercopidae, Anthoscytina, new species, fossil, Middle Jurassic, Daohugou Abstract. Anthoscytina Hong, 1983 is the largest genus within the Mesozoic Procercopidae, the stem group of the superfamily Cerco- poidea. Herein, we describe two new species from the Middle Jurassic of Daohugou, northeast China. Anthoscytina brevineura Chen, Wang & Zhang, sp. n. and Anthoscytina elegans Chen, Wang & Zhang, sp. n. are established on the basis of new well-preserved fossils. Although these two new species are very similar, some stable differences in tegminal venation and colour patterns confirm their species status. Sinotettegarcta longa Hong, 1986 is transferred to Anthoscytina, and to avoid secondary homonymy, a new name Anthoscytina hongi Chen, Wang & Zhang, nom. n. is proposed for that species. In addition, Anthoscytina aphthosa Ren, Yin & Dou, 1998 and An­ thoscytina macula Hu, Yao & Ren, 2014 are transferred from Anthoscytina to Stellularis Chen, Yao & Ren, 2015. INTRODUCTION City, Inner Mongolia, China. The palaeoenvironmental recon- structions indicate a humid and warm-temperate climate in the The Cercopoidea Leach, 1815 is one of the most speci- Middle Jurassic in the Daohugou area (Ren & Krzeminski, 2002; ose superfamilies of Cicadomorpha, with nearly 3000 de- Wang et al., 2013). These deposits are well-known for yielding a scribed species attributed to five modern families and three highly diverse array of insects (e.g., Hou et al., 2012a, b; Yan & extinct families from the Mesozoic (Hamilton, 2001; Di- Wang, 2010; Chen J. et al., 2014), including very large numbers etrich, 2002; Hu et al., 2012; Wang et al., 2012). Most fos- of cercopoids (Wang & Zhang, 2009; Wang et al., 2012; Li et sil and extant cercopoids are small insects, usually shorter al., 2013). All the material collected is housed in the Shandong than 13 mm (Hong, 1983; Wang & Zhang, 2009). Adult Tianyu Museum of Nature (STMN) at Pingyi, Shandong, China. cercopoids are commonly called froghoppers because they The fossils were examined using a stereomicroscope (Zeiss look like tiny frogs and are very adept at jumping (Bur- SteREO Discovery V8). Photographs were taken using a Nikon D800 digital camera. Line drawings were prepared with two im- rows, 2003). Their nymphs, known as spittlebugs, cover age-editing pieces of software (CorelDraw 12.0 and Adobe Pho- themselves with foaming spittle to provide protection from toshop CS3). In the line drawings, faintly seen and hypothesized predation, parasitism and desiccation (Hamilton, 1982; Li regions are indicated by dotted lines, the edges of missing regions et al., 2013). indicated by thin solid lines. All measurements were made us- The family Procercopidae, as the earliest cercopoid ing software ImageJ 1.42q (Wayne Rasband; National Institute group, is recorded from the Early Jurassic to Early Creta- of Health, USA). ceous in Germany, Russia, Central Asia, Southeast Asia and Currently there is no consensus over the interpretation of vein China. Anthoscytina Hong, 1983, the largest genus within nomenclature in Cicadomorpha (Wang et al., 2009). Herein, we the Procercopidae, was erected on the basis of a complete tentatively follow Nel et al. (2012) and Bourgoin et al. (2014) with slight modifications. tegmen from the Middle Jurassic of Beipiao, Liaoning in China (Hong, 1983). It was previously referred to the fam- SYSTEMATIC PALAEONTOLOGY ily Scytinopteridae Handlirsch, 1906 and later transferred Order Hemiptera Linnaeus, 1758 to the Procercopidae (Shcherbakov, 1988). The tegminal Suborder Cicadomorpha Evans, 1946 lengths reported for species of Anthoscytina range from 10 mm to 15 mm. Superfamily Cercopoidea Leach, 1815 Very recently the present authors collected some new gi- Family Procercopidae Handlirsch, 1906 ant procercopids from Daohugou in Northeast China, of Genus Anthoscytina Hong, 1983 which some are assigned to the genus Anthoscytina. We re- Type species. Anthoscytina longa Hong, 1983; Haifanggou port herein on two of these species with tegmens of around Formation, Beipao City, Liaoning, China; Middle Jurassic. 20 mm in length. Species included. A. longa Hong, 1983; A. reducta (Beck- er-Migdisova, 1949); A. liugouensis (Hong, 1983); A. daica MATERIAL AND METHODS Shcherbakov, 1988; A. parallelica Ren, Lu & Guo, 1995; A. tri­ The fossils were collected from the Middle Jurassic Jiulongshan nervus (Ren, Lu & Guo, 1995); A. pustulosus (Ren, Lu & Guo Formation in Daohugou Village, Ningcheng County, Chifeng 1995); A. perpetua Li, Shih & Ren, 2013; A. hongi Chen, Wang & 373 Fig. 1. Holotype of Anthoscytina brevineura Chen, Wang & Zhang, sp. n. A – photograph of part of STMH 48-1782a; B – photograph of part of STMH 48-1782a, under alcohol; C – photograph of the counterpart of STMH 48-1782b; D – illustration based on STMH 48-1782a. All to scale. Zhang, nom. n.; A. brevineura Chen, Wang & Zhang, sp. n.; and 1983 and A. longa Hong, 1983, respectively. Furthermore, A. elegans Chen, Wang & Zhang, sp. n. Wang et al. (2012) treated Mesocercopis Hong, 1983 and Revised diagnosis. Fore femur robust, hind tibia long Sinotettegarcta Hong, 1986 as synonyms of Anthoscy­ and slender, with a short lateral spine; Ovipositor short; tina Hong, 1983, but did not mention the validity of both tegmen with vein R bifurcating near basal 1/3 wing length; Mesocercopis longa Hong, 1983 and Sinotettegarcta longa RA simple or multi-branched; RP simple; M with at most Hong, 1986. For M. longa, the most important characters 3 branches; CuA branching basal of or at the same level as at the specific level are not included in the original descrip- the branch in M; cross vein im absent; hind wing with vein tion, so it is impossible to determine its species status until RA and RP simple, M and CuA two-branched. the type specimen is re-examined. Sinotettegarcta longa is Remarks. Shcherbakov (1988) revised the generic di- similar to A. longa in terms of the pattern in the venation agnosis and transferred Cycloscytina reducta Becker- and size, but distinctly differs from the latter in that on the Migdisova, 1949 to this genus. He suggested that Para­ tegmen R is multi-branched and M branches close to the cicadella Hong, 1983 and Paracicadella beipiaoensis apex of the wing. Hong, 1983 are junior synonyms of Anthoscytina Hong, 374 Fig. 2. Paratypes of Anthoscytina brevineura Chen, Wang & Zhang, sp. n. A – STMH 48-1783; B – hind leg of 48-1783, under alco- hol; C – line drawing of hind wing of 48-1783; D – STMH 48-1784a; E – STMH 48-1785; F – STMH 48-1786a. Scale bars = 2 mm. Anthoscytina hongi Chen, Wang & Zhang, nom. n. is similar to A. daica and A. parallelica in having a tegmen Anthoscytina hongi Chen, Wang & Zhang, nom. n. for An­ with a multi-branched R and differs in its M1+2 and M3+4 are thoscytina longa (Hong, 1986: 14–15, Fig. 5, Pl. II: 3) comb. n. longitudinal and nearly parallel. (described as Sinotettegarcta longa), a junior secondary homo- Anthoscytina brevineura Chen, Wang & Zhang, sp. n. nym of Anthoscytina longa Hong, 1983: 61–62, Fig. 47, Pl. 10: 3. (Figs 1, 2) Etymology. The specific epithet is named after Youchong Hong who described the species. Etymology. The specific epithet is derived from the Latin “bre- vis” (meaning short) and “neurus” (meaning vein), which refers Remarks. This species is very similar to the type species to the extremely short cross vein m­cua. of Anthoscytina, but differs from the latter in having a teg- Holotype. STMN48-1782, sex unknown, adult in ventral view men with cross veins ir and r­m almost at the same level. It with tegmen preserved. 375 Fig. 3. Holotype of Anthoscytina elegans Chen, Wang & Zhang, sp. n. A – photograph of part of STMH 48-1787a; B – photograph of part of STMH 48-1787a, under alcohol; C – photograph of the counterpart of STMH 48-1787b; D – illustration based on STMH 48-1787a. All to scale. Type locality. Daohugou Village, Shantou Township, Ning- CuA2; CuA2 short and oblique. Hind wing, M1+2 and CuA cheng County, Inner Mongolia, China. nearly straight. Type horizon. Jiulongshan Formation, Middle Jurassic. Description. Body 21.0–23.2 mm long including teg- Paratypes. STMN48-1783, sex unknown, adult in dorsal view men in repose (Figs 1, 2A, D–F). Postclypeus swollen, with tegmen and hind wing preserved; STMN48-1784, adult fe- about 2.0 mm long, with weak oblique grooves and a dis- male in lateral aspect with tegmen preserved; STMN48-1785, adult male in lateral aspect with tegmen and hind wing preserved; tinct median groove. Compound eyes large, nearly round STMN48-1786, sex unknown, adult in lateral aspect with tegmen (Figs 1, 2F). Pronotum well-developed, approximately 3.0 preserved. mm wide. Fore femur extremely strong, about 2.5 mm long and 0.8 mm wide; fore tibia about 2.4 mm long (Fig 2D). Diagnosis. Body large. Tegmen broad; membrane Middle femur slenderer than fore femur, approximately tinged with longitudinal stripes, and apical margin darkly 0.6 mm wide; middle tibia slightly longer than fore tibia, stained; RA simple, connected with M by cross vein r­m 1+2 approximately 2.7 mm long; middle tarsus about 1.0 mm just distal of junction with cross vein ir; stem M branch- long, with three tarsomeres; apical tarsomere longer than ing into M and M near base at 0.9 wing length; M 1+2 3+4 1+2 mid tarsomere and basitarsomere; two claws and an aro- short, transverse; M vertical, rectangular at junction with 3+4 lium visible (Figs 2A, D).
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