from the Early Cretaceous of China

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Cretaceous Research 45 (2013) 306e313

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A remarkable new genus of basal snakeflies (Insecta: Raphidioptera: Priscaenigmatomorpha) from the Early Cretaceous of China Xingyue Liu a, Vladimir N. Makarkin b, c, *, Qiang Yang b, Dong Ren b, ** a

Department of Entomology, China Agricultural University, Beijing 100193, China College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China c Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 16 March 2013 Accepted in revised form 3 June 2013 Available online 11 July 2013

Chrysoraphidia relicta gen. et sp. nov. is described from the Early Cretaceous (Barremian) of the Yixian Formation, Liaoning Province, China. Its venation is characterized by a mixture of character states that occur mainly in the Neuroptera and Raphidioptera. The assignment of Chrysoraphidia gen. nov. to Raphidioptera is supported by the presence of its distinct pterostigma and the configuration of the wing vein ‘subcosta anterior’. It is interpreted as the first record of a group of basal snakeflies (Priscaenigmatomorpha) from the Cretaceous of Asia, hitherto known only from the Early Jurassic of Europe. Ó 2013 Elsevier Ltd. All rights reserved.

Keywords: Raphidioptera Priscaenigmatomorpha New genus Yixian Formation

1. Introduction Raphidioptera (snakeflies) are a distinctive, minor holometabolous order, belonging to the superorder Neuropterida (Aspöck et al., 1991; Grimaldi and Engel, 2005). The adults of Raphidioptera are characterized by a prognathous head, narrowly elongate prothorax, and a long ovipositor in females, although many extinct Mesozoic snakeflies had a relatively broad and short prothorax (see e.g., Engel and Ren, 2008). About 230 valid described species of recent snakeflies are currently known, all restricted to the Northern Hemisphere (Haring et al., 2011; Aspöck et al., 2012). The western Palaearctic has the richest species diversity of extant Raphidioptera, while East and Southeast Asia are becoming known as another diversity centre of snakeflies due to the rapidly increasing number of species described there within the last four years (Liu et al., 2009, 2010a,b, 2012; Aspöck et al., 2011). The phylogenetic position of the order within Neuropterida is still debated. Traditionally, it is considered to be the sister to Megaloptera, which is supported by molecular data from nuclear genes (Wiegmann et al., 2009), while it is found to be sister to the remaining two orders of Neuropterida (Megaloptera þ Neuroptera) based on both morphological and molecular data (mainly from mitochondrial * Corresponding author. Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia. Tel.: þ7 423 2311133. ** Corresponding author. Tel.: þ86 10 68980851. E-mail addresses: [email protected] (V.N. Makarkin), rendong@ mail.cnu.edu.cn (D. Ren). 0195-6671/$ e see front matter Ó 2013 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.cretres.2013.06.001

genes) (Aspöck et al., 2001; Haring and Aspöck, 2004; Cameron et al., 2009; Wang et al., 2012). Its estimated divergence time based on molecular sequence data indicate that Raphidioptera might have originated ca. 250 Ma ago in the Late Permian (Winterton et al., 2010), although the earliest fossil snakefly was found in the Early Jurassic (Bode, 1953; Whalley, 1985; Willmann, 1994). Snakeflies prospered in the Mesozoic, with high species diversity and global distribution, but underwent significant extinction at the end of the Cretaceous, leaving extant species as a relict group (Aspöck, 1998; Engel, 2002). Diverse Mesozoic snakeflies have been described from Eurasia, North America, and South America comprising ca. 90 species in 33 genera of three families, i.e. Priscaenigmatidae, Baissopteridae, and Mesoraphidiidae (Engel, 2002; Engel and Ren, 2008; Jepson et al., 2011; Pérez-de la Fuente et al., 2012). In China, 14 genera and 25 species of the two latter families are known from the Middle Jurassic to Early Cretaceous, representing nearly 1/4 of the described fossil snakefly species (see a list in Ren et al., 2010). Two enigmatic monotypic genera are known from the Early Jurassic, Priscaenigma Whalley, 1985 from Sinemurian of Charmouth, England, and Hondelagia Bode,1953 from early Toarcian of Hondelage, Germany. These genera are now treated as comprising the family Priscaenigmatidae and the suborder Priscaenigmatomorpha, and considered to be the basal-most known lineage of Raphidioptera (Engel, 2002; Bechly and Wolf-Schwenninger, 2011). An undescribed raphidiopteran species from the early Toarcian of Schandelah (Germany) also belongs to Priscaenigmatidae (see Ansorge, 2003, fig. 3k). It is much smaller than other Early Jurassic species: the forewing is

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7.2 mm and ca. 11e12.6 mm long, respectively. In this paper, we describe a new genus and species from the Early Cretaceous Yixian Formation of Huangbanjigou (China), which we treat as belonging to this group. These new fossils represent the first record of Priscaenigmatomorpha in Asia and indicate their presence in the Early Jurassic to Early Cretaceous of Eurasia. 2. Material and methods Three specimens described herein come from the Huangbanjigou locality. It is situated in the Sihetun area, approximately 21 km south of Beipiao in the western Liaoning Province of China, and belongs to the Jianshangou Member (Bed) of the lower Yixian Formation (Wang and Zhou, 2008). A late Barremian age for this fossil-bearing stratum is considered to be well supported by radiometric dating, from 126.1  1.7 to 124.6  0.1 Ma (Swisher et al., 1999, 2002; Wang et al., 2001b; Chen et al., 2004; Yang et al., 2007); the uppermost beds of Huangbanjigou locality are early Aptian, 123.3  0.5e122.8  1.6Ma (Wang et al., 2001a; Yang et al., 2007). Raphidioptera from the Yixian Formation were described by Ren (1994, 1997) and Ren et al. (1995); its neuropteran assemblage was analyzed by Makarkin et al. (2012). The fossils were examined using a Leica M165C dissecting microscope and illustrated with the aid of a drawing tube. Photos of all specimens were taken by Nikon D90 and Leica DFC500 digital cameras. All fossil specimens described herein are deposited in the Key Laboratory of Insect Evolution & Environmental Changes, College of Life Sciences, Capital Normal University, Beijing (CNUB, D. Ren, curator). The wing venation of Raphidioptera has been quite poorly studied. There are three concepts of the raphidiopteran wing venation (similarly that of other insect orders), differing from each other mainly in treatment of the median vein: (1) a traditional concept considering M as having a common stem, with its anterior branches not fused with R or Rs (e.g., Comstock, 1918, figs. 167, 168; Withycombe, 1923, figs. 1, 2); (2) a concept based on the hypothesis of Martynov (1928), which considers M also having a common stem but MA is medially fused with R or Rs (e.g., Aspöck et al., 1991, figs. 43, 44; Bechly and Wolf-Schwenninger, 2011, fig. 16); (3) a concept which considers MA and MP as having no common stem, and MA fully fused with R and Rs (e.g., Kukalová-Peck and Lawrence, 2004, fig. 9). We preliminarily accept the concept (2), pending more detailed examination of raphidiopteran venation, and therefore broadly follow the terminology of Aspöck et al. (1991), except in the treatment of ‘ScA’ in which we follow Kukalová-Peck (1991). Wing abbreviations used in the text and figures are as follows: A, anal; C, costa; CuA, cubital anterior; CuP, cubital posterior; dc, discal cell; doi, discoidal cell; m, medial cell; MA, medial anterior; MP, medial posterior; pt, pterostigma; r, radial cell; R, radial; Rs, radial sector; Sc, subcosta; ‘ScA’, ‘subcosta anterior’. 3. Systematic palaeontology Order: Raphidioptera Navás, 1916 Suborder: Priscaenigmatomorpha Engel, 2002 Family: Uncertain Genus Chrysoraphidia gen. nov. Type species. Chrysoraphidia relicta sp. nov. Derivation of name. From Chrysopa and Raphidia, genera-group names, in reference to superficial resemblance of Chrysoraphidia wings to those of Chrysopidae (Neuroptera). Gender: feminine. Diagnosis. Medium-sized raphidiopterans (forewing 12.1e13.5 mm long). Wings ovoid, relatively broad; pterostigma distinct, elongate,

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open proximally, closed distally by veinlet of R; Sc running within pterostigma to its distal ending, connecting with R by distal oblique crossvein; MA simple; MP deeply forked, with simple anterior branch and distally forked posterior branch; CuA pectinately branched, with two branches; CuP simple; two series of gradate crossveins in radial to mediocubital areas. In forewing, costal area basally with short ‘ScA’ terminating on C; 1A long, pectinately branched, with two branches; 2A simple. Remarks. The placement of the new genus in Priscaenigmatomorpha is based on the following character states: the long Sc extending the pterostigma; the similar configuration CuA and CuP; Rs þ MA separating from R near the wing base; and the narrow anal cell between 1A and 2A. Chrysoraphidia gen. nov. can be distinguished from the two Jurassic genera by its more distinct pterostigma in both wings; Sc running more closely to C within the pterostigma; not so strongly zigzagging of Rs and its branches; more distal origin of Rs þ MA; the simple MA in both wings; and its pectinate 1A. The new genus differs from Priscaenigma also by relatively broad wings, the decreasing number of costal veinlets (10), and the absence of crossveins between branches of CuA. The differences between the new genus and Hondelagia refer to the shape of costal area in the forewings and the branching number of Rs, MP, and 1A (see Willmann, 1994). The new genus represents the first record of the basal suborder Priscaenigmatomorpha from China and in general outside Europe and shows that this group occurred not only in the Early Jurassic but also persisted into the Early Cretaceous. Chrysoraphidia relicta sp. nov. Figs. 1e7 Derivation of relinquished.

name.

From

the

Latin

relictus,

abandoned,

Type material. Holotype, CNU-RAP-LB-2012001, a well-preserved incomplete specimen in dorsal aspect. Paratype, CNU-RAP-LB2012002, a rather well-preserved incomplete specimen in lateral aspect. Paratype CNU-RAP-LB-2012003, a poorly preserved incomplete specimen in lateral aspect. Type locality and horizon. All collected from the Huangbanjigou locality (41360 4400 N, 120 490 4800 E), Beipiao City, Liaoning Province, China; Early Cretaceous (Yixian Formation). Diagnosis. As for the genus. Description. Holotype (Figs. 1e3). Body robust; head, prothorax not preserved; meso-, metathorax, abdomen together 9.4 mm long. Mesonotum less than 2 mm long, 2.3 mm wide in dorsal view; prescutum relatively large. Legs fragmentarily preserved, details not visible. Abdomen rather broad (maximal width 2.5 mm in dorsal view); abdominal segments appear rather narrow, fuscous, with lighter intersegmental membrane; terminal segment caudally rounded. Forewing ovoid, relatively broad; 12.3 mm long, 4.6 mm wide. Costal area rather narrow basally, then dilated, again narrowed before pterostigma, with seven or eight veinlets. Basally, ‘ScA’ rather clearly visible in right forewing. Sc long, running rather close to C within pterostigma, terminating near its distal ending, connecting with R by distal crossvein (Fig. 2B; distal-most part of Sc poorly preserved). R long, terminating before wing apex, distally with two widely spaced short veinlets. Pterostigma well developed, elongate (ca. 2.5 mm long), dark; area within pterostigma posterior to Sc slightly paler than area between Sc and C. Six crossveins present between R and Rs, forming six radial cells; 2r over twice length of 1r, about 1.5 times length of 4r; 3r, 5r, 6r together nearly

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Fig. 2. Chrysoraphidia relicta gen. et sp. nov., holotype CNU-RAP-LB-2012001. A, basal portions of left wings; B, pterostigmal areas of left wings; C, pterostigmal area of the right hind wing. All wetted with ethanol. hp, humeral plate of hind wing. Scale bars represent 1 mm.

Fig. 1. Chrysoraphidia relicta gen. et sp. nov., holotype CNU-RAP-LB-2012001. A, specimen as preserved (dry); B, proximal half of right forewing (wetted with ethanol). Scale bars represent 2 mm.

equal length of 1r. Rs þ MA origins from R approximately 1.9 mm from wing base. Rs with four simple branches. Two discal cells present between proximal branch of Rs, MA; 1dc about twice length of 2dc. Two parallel series of gradate crossveins present between branches of Rs to CuA: inner series incomplete with five crossveins between second branch of Rs and CuA, outer series complete, with eight crossveins between R, CuA. MA simple, proximally coalesced with Rs for distance nearly 1/3 length of forewing. Four crossveins present between MA, MP, forming three medial cells. Basal part of MP not clearly visible; MP deeply forked, with anterior branch simple, posterior branch forked distally. Three discoidal cells present between branches of MP; two present between MP, CuA. Cu dividing into CuA, CuP rather far from wing base, slightly distal to origin of Rs. CuA zigzagged distally, pectinately branched, with two simple branches. Two crossveins present between CuA, CuP. 1A long, pectinately branched, with two (right wing) or three (left wing) simple branches; 2A simple. Anal cell between 1A, 2A narrow. Two crossveins between CuP, 1A; one

crossvein each between 1A, 2A, between 2A, 3A. 3A incompletely preserved. Membrane probably colourless, transparent except for dark pterostigma. Hind wing considerably shorter than forewing, 9.8 mm long. Humeral plate well developed, stick-shaped, probably covered with fine short hairs at tip (Fig. 2A). Costal area somewhat dilated before pterostigma, with four or five veinlets preserved. Sc long, running close to C within pterostigma, terminating near its distal ending, connecting to R by poorly preserved oblique crossvein (Fig. 2C). R long, terminating near wing apex, with one veinlet at distal ending of pterostigma. Pterostigma well developed, elongate (ca. 2.2 mm long), dark; area within pterostigma posterior to Sc much paler. Five or six crossveins present between R, Rs, forming five or six radial cells respectively; 1r, 3r conspicuously longer than other radial cells, about 1.5 times length of 2r. Rs origins from R rather close to wing base. Rs with five simple nearly straight branches, directed to outer margin. Two series of gradate crossveins present between branches of Rs, MA (basal part of hind wings poorly preserved): inner series incomplete with one preserved crossveins between branches of Rs, outer series complete, with five crossveins between R, MA. MA incompletely preserved, probably simple, proximally coalesced with Rs. MP incompletely preserved;

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Fig. 4. Chrysoraphidia relicta gen. et sp. nov., paratype CNU-RAP-LB-2012002. A, specimen as preserved; B, pterostigmal areas of left wings. Sc(f), forewing subcosta; Sc(h), hind wing subcosta; sc-r, distal crossvein between Sc and R. Scale bars represent 2 mm (A), 1 mm (B).

Fig. 3. Drawings of Chrysoraphidia relicta gen. et sp. nov., holotype CNU-RAP-LB2012001. A, left forewing; B, right forewing; C, left hind wing; D, right hind wing. Scale bar represents 5 mm (all to scale).

proximal portion neither forked nor branched. CuA pectinately branched. CuP thinner, probably simple. 1A running close to CuP. Membrane probably colourless, transparent except for dark pterostigma. Paratype CNU-RAP-LB-2012002. (Figs. 4e6). Head and antennae only fragmentarily preserved. Pronotum not visible but most probably slightly elongate judging from position of head and mesothorax. Mesothorax, metathorax poorly preserved; details not visible. Legs fragmentary preserved. Abdomen appears very broad in lateral view (possibly compressed post mortem); relatively short tergites, sternites dark; intersegmental membrane light. Forewing 11.2 mm long, 4.5 mm wide. Costal area as in holotype, with seven veinlets. Basally, ‘ScA’ clearly visible in left forewing (Fig. 5B). Sc long, running not close to C within pterostigma and terminating probably near its distal end (distal-most part of Sc not preserved), connecting distally with R by distinct oblique crossvein (visible in right forewing; Fig. 4B). R long, terminating before wing apex, distally with two widely spaced short veinlets. Pterostigma well developed, elongate, uniformly darkish. Five crossveins preserved between R, Rs; radial cell 2r over twice length of 1r. Rs with five simple branches (right wing). Two discal cells present between Rs, MA; 1dc about 1/5 length of 2dc. Two parallel series of gradate crossveins present between branches of Rs to CuA: inner series incomplete with six crossveins preserved

between first branch of Rs and CuA, outer series complete, with nine crossveins between R, CuA. MA simple, proximally coalesced with Rs for distance nearly 1/3 length of forewing. Three crossveins preserved between MA, MP. Basal part of MP not clearly visible but most probably origins from R (i.e., MP appears fused with R for considerable distance); two basal crossveins each between R, MP, between MP, CuA. MP deeply forked, with anterior branch simple, posterior branch forked distally. Three crossveins present between branches of MP, and between MP, CuA. Cu dividing into CuA, CuP rather far from wing base, slightly distal to origin of Rs. CuA zigzagged distally, pectinately branched, with two simple branches. Two crossveins present between CuA, CuP. CuP simple. 1A long, pectinately branched, with two simple branches; 2A simple. 3A probably deeply forked, with crossvein between branches (alternately, posterior branch of 3A is 4A; incompletely preserved). Anal cell between 1A, 2A narrow. Crossveins between CuP, 1A not preserved. Membrane probably colourless, transparent except for dark pterostigma. Hind wing ca. 9 mm long. Sc long, running not close to C within pterostigma; terminating near its distal end; distal crossvein not preserved (Fig. 4B). R long, terminating near wing apex, with two distal veinlets. Pterostigma well developed, elongate, uniformly dark. Five crossveins preserved between R, Rs. Rs probably with four simple branches. Two series of gradate crossveins present between branches of Rs and CuA: inner series with two preserved crossveins between MA, second branch of Rs; outer series with seven crossveins between R, CuA. MA simple. MP basally unclear; deeply forked slightly proximal origin MA from Rs; anterior branch simple, posterior branch forked distally. CuA pectinately branched, with two simple oblique branches. Basal portion of wings poorly preserved.

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Fig. 6. Drawings of Chrysoraphidia relicta gen. et sp. nov., paratype CNU-RAP-LB2012002. A, left forewing; B, right forewing; C, left hind wing; D, right hind wing. Scale bar represents 5 mm (all to scale).

Fig. 5. Chrysoraphidia relicta gen. et sp. nov., photographs of basal portion of wings of the paratype CNU-RAP-LB-2012002. A, dry specimen; B, wetted with ethanol. Scale bar represents 1 mm (both to scale).

Paratype CNU-RAP-LB-2012003 (Fig. 7). Head, pronotum not preserved. Mesothorax, metathorax, legs poorly preserved, probably crumpled. Abdomen very short, broad in lateral view; details not visible. Forewing ca. 13.5 mm long, 5.3 mm wide. Preserved venation is very similar to that of holotype right forewing, but 1A is configured as in left wing. Pterostigma nearly unicolorous. Sc within pterostigma; distal crossvein between Sc, R clearly visible; termination of Sc not preserved; distal sc-r almost perpendicular to Sc (Fig. 7B). Hind wing venation very poorly preserved. In right wing, Sc within pterostigma, distal crossvein between Sc, R clearly visible; Sc entering wing margin slightly distal to distal sc-r, which is perpendicular to Sc. R with two closely spaced veinlets immediately distal to pterostigma (Fig. 7B). Remarks. All the specimens lack an ovipositor, and are assumed to be males.

4. Distinctive morphological characters of Chrysoraphidia gen. nov. The general venational pattern of the forewing of Chrysoraphidia gen. nov. is quite similar to that of some fossil taxa of Neuroptera, e.g., the Early Cretaceous Osmylochrysa anomala Jepson et al., 2012 (Osmylidae), basal Raphidioptera, i.e., the Early Jurassic Priscaenigma and Hondelagia, and Megaloptera, e.g., the Late Eocene Corydasialis inexpectatus Wichard et al., 2005 (Corydasialidae). Chrysoraphidia gen. nov. is clearly distinguished from these Neuroptera and Megaloptera taxa by the presence of distinct pterostigma. The dark pterostigma is present in some taxa of Neuroptera (especially distinct in Mantispidae, Mesochrysopidae, and Ascalaphidae) and all Raphidioptera. The structure of the pterostigma (and in general the costal area) of Chrysoraphidia gen. nov. is superficially most similar to those of Mantispinae, the most advanced subfamily of Mantispidae. However, the pterostigma in those families of Neuroptera and Raphidioptera was evolved independently with certainty as their venation is otherwise strongly different.

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insect orders (most developed in Orthoptera and Palaeodictyoptera), and is thus plesiomorphic. ‘ScA’ of Neuroptera (when present) is very differently configured, always terminating on Sc, not C (see Yang et al., 2012). 5. Order and family affinity of Chrysoraphidia gen. nov.

Fig. 7. Chrysoraphidia relicta gen. et sp. nov., paratype CNU-RAP-LB-2012003. A, specimen as preserved; B, pterostigmal areas of right wings. Sc(f), forewing subcosta; Sc(h), hind wing subcosta; sc-r, distal crossvein between Sc and R. Scale bars represent 2 mm (A), 1 mm (B).

The basal part of the stem of MP has a tendency to reduction. It is invisible in the holotype of Chrysoraphidia relicta gen. et sp. nov, but in one of the paratypes it is relatively well visible as originating from R (Figs. 5 and 6). This character state is plesiomorphic here, as it occurs in most taxa of Neuropterida. The pectinate 1A is characteristic of this genus. This character state often occurs in the majority taxa of Neuroptera, and the megalopteran family Corydasialidae, but is absent in all higher snakeflies (Raphidiomorpha), and not detected in the Early Jurassic Priscaenigmatidae (although in known specimens of these taxa, the forewing proximal portion is poorly preserved). The well developed, stick-shaped, erected humeral plate in the hind wing as found in Chrysoraphidia gen. nov. was hitherto unknown in Raphidioptera, nor in Megaloptera. Such a humeral plate is characteristic of some families of Neuroptera, e.g., Psychopsidae, Ithonidae, Parakseneuridae (Tjeder, 1960, fig. 327; Makarkin and Menon, 2007, fig. 2; Yang et al., 2012, figs. 13, 23, 26, 30C). Unfortunately, the basal portions of hind wings of the early Jurassic genera of Priscaenigmatidae specimens are not preserved. The humeral plate of other extant Raphidioptera and Megaloptera do not project (appearing as the basal-most portion of the Costa), although the humeral lobe is sometimes well developed. The presence of a vein called here the ‘subcosta anterior’ (ScA) is one of the most characteristic features of this genus. Some extant Raphidioptera (especially Raphidiidae) possess the vestigial, remnant ‘ScA’ in the base of the forewing costal area. In these, a short convex fold obliquely runs from the wing base near Sc to the costal margin, where a very short vein lacking hairs is present (interpreted here as the remnant ‘ScA’). A similar ‘ScA’ is shown to be primitively present in the extant Corydalidae (see KukalováPeck, 1991, fig. 6.16). In Chrysoraphidia gen. nov., ‘ScA’ is poorly preserved in the holotype, but very clearly present in the paratypes CNU-RAP-LB-2012002. ‘ScA’ of extant Raphidioptera, Chrysoraphidia gen. nov. and Megaloptera (at least Corydalinae) are similarly constructed and probably homologous, due to their similar position in the wing. Such a form of ‘ScA’ is characteristic of some

This short analysis of the distinctive characters of Chrysoraphidia gen. nov. shows that its venation possesses a mixture of conditions occurring in Neuroptera and Raphidioptera, and to lesser extent in Megaloptera. The configuration of ‘ScA’ excludes Neuroptera from consideration (see above), and the presence of distinct pterostigma excludes Megaloptera. Therefore, Raphidioptera is the most probable assignment of this genus based on the presence of the pterostigma and ‘ScA’ running to the costal margin. Besides, the presence of other typical snakefly features (i.e., elongate prothorax and female ovipositor) in Chrysoraphidia gen. nov. cannot be eliminated until a completely preserved female of this genus is found. Actually, judging from the position of the head and mesothorax in the paratype CNU-RAP-LB-2012002 of Chrysoraphidia relicta sp. nov., the pronotum is likely to be elongated in this species. The venation of this Cretaceous genus is most similar to that of the Early Jurassic genera Priscaenigma and Hondelagia, which comprise the family Priscaenigmatidae and the suborder Priscaenigmatomorpha. Its diagnosis includes the following forewing character states: Sc is long, running within the pterostigma, extending nearly to the wing apex [Sc is short, terminating on C before the pterostigma in all other Raphidioptera]; Rs þ MA originates near the wing base [Rs þ MA originates very far from wing base in other Raphidioptera]; MP is basally fused with Rs þ MA or CuA (current knowledge does not allow a decision on which variant is correct); Cu is continuous with CuA [Cu is continuous with CuP; CuA appears proximally as a crossvein between Cu and M in other Raphidioptera]; the cell between 1A and 2A is narrow [the cell between 1A and 2A is broadly ovoid in other Raphidioptera]. Most of these conditions are plesiomorphic, at least in Neuropterida, except for the basal structure of MP. The same configuration of basal MP in Priscaenigma and Hondelagia may be interpreted as the only distinct synapomorphy of Priscaenigmatidae supporting its monophyly. Both variants (MP basally fused with Rs þ MA or CuA) are apomorphic relatively to the plesiomorphic condition where MP (and then M) is basally either running freely to the wing base or fused for a short distance with R in most other Neuropterida (including Chrysoraphidia gen. nov.). Although the basal fusion of MP and CuA is characteristic of all Raphidioptera (if MP is interpreted as basally fused with CuA), the mode of their fusion is quite different in Priscaenigmatomorpha and other Raphidioptera (Raphidiomorpha): there, MP and CuA run as a single vein towards the wing base in the former, and these veins are widely separate at the wing base in the latter. Therefore, it is likely that the basal fusion of MP and CuA was independently evolved in these groups. Chrysoraphidia gen. nov. shares all of these character states with Priscaenigmatidae, except the configuration of MP, which is basally not fused with either CuA or Rs þ MA in this genus. Additionally, the genus rather greatly differs from Priscaenigma and Hondelagia by other characters, i.e., the origin of Rs þ MA is located relatively far from the wing base; many veins are mostly not zigzagged, especially Rs and its branches; and 1A is pectinate. Therefore, the assignment of the new genus to Priscaenigmatidae cannot be confirmed confidently, but its assignment to Priscaenigmatomorpha is very probable. The new genus cannot be assigned to a new family, as the vast majority (or all) of the diagnostic character states of this potential new family would be plesiomorphic in Neuropterida. As a result, we do not assign Chrysoraphidia gen. nov. to any

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family, pending the description of other genera of Priscaenigmatomorpha clarifying the situation. In general, the order affinity of the family Priscaenigmatidae (and Priscaenigmatomorpha) is debatable. Its type genus (Priscaenigma) was described by Whalley (1985) as belonging to the order Raphidioptera, to which it has usually been considered to belong since then (e.g., Lambkin, 1986; Oswald, 1990; Aspöck et al., 1991; Willmann, 1994). The genus Hondelagia was described by Bode (1953) in Neuroptera as the only genus of ‘Eomantispidae’ (an unavailable name). Ponomarenko (1980) assigned it to the Raphidioptera. Carpenter (1992) considered this genus as Neuroptera incertae sedis, however, possibly belonging to Raphidioptera. Willmann (1994) reexamined the type material of these two genera, and concluded they might be regarded as the basal-most group(s) of Raphidioptera. Engel (2002) described the family Priscaenigmatidae (and the suborder Priscaenigmatomorpha), comprised of these two genera. Priscaenigmatomorpha is considered to be sister of Raphidiomorpha, i.e., all other Raphidioptera (Engel, 2002; Bechly and Wolf-Schwenninger, 2011). However, this placement was questioned by Ponomarenko (2002) and Aspöck and Aspöck (2004), who excluded Priscaenigma and Hondelagia from Raphidioptera, but did not assign them to any other order. Aspöck and Aspöck (2004) proposed four autapomorphies of Raphidioptera (excluding Priscaenigmatidae, i.e., in fact of Raphidiomorpha): (1) the forewing Sc runs into anterior wing margin (i.e., Sc is very short); (2) imaginal tarsi with expanded (bilobed) third tarsomeres; (3) amalgamation of tergite and sternite of the 9th abdominal segment in the male form a ring, and (4) elongation of the female ovipositor. The Sc in the forewing is obviously long in Priscaenigmatidae, and the other three autapomorphies are not preserved in all known fossils of this family. Recently, Bechly and Wolf-Schwenninger (2011) accepted the placement of Priscaenigmatidae in Raphidioptera, and listed the following venational synapomorphies shared by taxa of this order: (A) MP is developed as an oblique veinlet between MA and CuA in the hind wing; (B) the elongate open cell between the basal parts of Rs and M/MA in the hind wing; and (C) the pterostigma is developed (at least weakly). However, the character states (A) and (B) are plesiomorphic in Neuropterida. The character condition (A) is present in most extant Neuropterida species and primitively characteristic of all Neoptera (known as the so called ‘M5’; see Ren and Makarkin, 2009; Yang et al., 2012); the character condition (B) is a plesiomorphy of Neuropterida and present in half of the extant Raphidioptera species. Only the condition (C) may be regarded as a synapomorphy shared by all taxa of the order Raphidioptera. The placement of Priscaenigmatidae in Raphidioptera is thus based on only one synapomorphy, i.e., the pterostigma is at least weakly developed. Although the developed pterostigma is also present in some Neuroptera families, it seems plausible to interpret that this character is independently evolved in various lineages of Neuropterida rather than symplesiomorphic. Rigorous study on the higher phylogeny of Raphidioptera should be made in the future based on comprehensive and detailed studies of fossil taxa and correct interpretation of the evolution of their wing venation. Acknowledgements We thank S. Bruce Archibald (Simon Fraser University, Burnaby, Canada) for correcting the English, and anonymous reviewers for their critical review of the manuscript. This research was supported by the National Natural Science Foundation of China (No. 41271063, 31071964, 31230065, 41272006), the National Basic Research Program of China (973 Program) (2012CB821906), the Foundation for the Author of National Excellent Doctoral Dissertation of PR China (No. 201178), Project of Great Wall Scholar and KEY project of Beijing

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