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Cretaceous Research 45 (2013) 306e313

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Cretaceous Research

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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,**

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

a r t i c l e i n f o

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

Keywords:RaphidiopteraPriscaenigmatomorphaNew genusYixian Formation

* Corresponding author. Institute of Biology and Soil Sthe Russian Academy of Sciences, Vladivostok 690022, R** Corresponding author. Tel.: þ86 10 68980851.

E-mail addresses: vnmakarkin@mail.ru (V.mail.cnu.edu.cn (D. Ren).

0195-6671/$ e see front matter � 2013 Elsevier Ltd.http://dx.doi.org/10.1016/j.cretres.2013.06.001

a b s t r a c t

Chrysoraphidia relicta gen. et sp. nov. is described from the Early Cretaceous (Barremian) of the YixianFormation, Liaoning Province, China. Its venation is characterized by a mixture of character states thatoccur mainly in the Neuroptera and Raphidioptera. The assignment of Chrysoraphidia gen. nov. toRaphidioptera is supported by the presence of its distinct pterostigma and the configuration of the wingvein ‘subcosta anterior’. It is interpreted as the first record of a group of basal snakeflies (Priscae-nigmatomorpha) from the Cretaceous of Asia, hitherto known only from the Early Jurassic of Europe.

� 2013 Elsevier Ltd. All rights reserved.

1. Introduction

Raphidioptera (snakeflies) are a distinctive, minor holometab-olous order, belonging to the superorder Neuropterida (Aspöcket al., 1991; Grimaldi and Engel, 2005). The adults of Raphi-dioptera are characterized by a prognathous head, narrowly elon-gate prothorax, and a long ovipositor in females, although manyextinct Mesozoic snakeflies had a relatively broad and short pro-thorax (see e.g., Engel and Ren, 2008). About 230 valid describedspecies of recent snakeflies are currently known, all restricted tothe Northern Hemisphere (Haring et al., 2011; Aspöck et al., 2012).The western Palaearctic has the richest species diversity of extantRaphidioptera, while East and Southeast Asia are becoming knownas another diversity centre of snakeflies due to the rapidlyincreasing number of species described there within the last fouryears (Liu et al., 2009, 2010a,b, 2012; Aspöck et al., 2011).

The phylogenetic position of the order within Neuropterida isstill debated. Traditionally, it is considered to be the sister to Mega-loptera, which is supported by molecular data from nuclear genes(Wiegmannet al., 2009),while it is found to be sister to the remainingtwo orders of Neuropterida (Megaloptera þ Neuroptera) based onboth morphological and molecular data (mainly frommitochondrial

ciences, Far Eastern Branch ofussia. Tel.:þ7423 2311133.

N. Makarkin), rendong@

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genes) (Aspöck et al., 2001; Haring and Aspöck, 2004; Cameron et al.,2009; Wang et al., 2012). Its estimated divergence time based onmolecular sequence data indicate that Raphidioptera might haveoriginated ca. 250Maago 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 intheMesozoic, with high species diversity and global distribution, butunderwent significant extinction at the end of theCretaceous, leavingextant 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 33genera of three families, i.e. Priscaenigmatidae, Baissopteridae, andMesoraphidiidae (Engel, 2002; Engel and Ren, 2008; Jepson et al.,2011; Pérez-de la Fuente et al., 2012). In China, 14 genera and 25species of the two latter families are known from the MiddleJurassic to Early Cretaceous, representing nearly 1/4 of thedescribed fossil snakefly species (see a list in Ren et al., 2010).

Two enigmatic monotypic genera are known from the EarlyJurassic, PriscaenigmaWhalley, 1985 from Sinemurian of Charmouth,England, andHondelagiaBode,1953 fromearly ToarcianofHondelage,Germany. These genera are now treated as comprising the familyPriscaenigmatidae and the suborder Priscaenigmatomorpha, andconsidered to be the basal-most known lineage of Raphidioptera(Engel, 2002; Bechly andWolf-Schwenninger, 2011). An undescribedraphidiopteran species from the early Toarcian of Schandelah (Ger-many) also belongs to Priscaenigmatidae (see Ansorge, 2003, fig. 3k).It is much smaller than other Early Jurassic species: the forewing is

X. Liu et al. / Cretaceous Research 45 (2013) 306e313 307

7.2 mm and ca. 11e12.6 mm long, respectively. In this paper, wedescribe a new genus and species from the Early Cretaceous YixianFormation of Huangbanjigou (China), which we treat as belonging tothis group. These new fossils represent the first record of Priscae-nigmatomorpha in Asia and indicate their presence in the EarlyJurassic to Early Cretaceous of Eurasia.

2. Material and methods

Three specimens described herein come from the Huangbanji-gou locality. It is situated in the Sihetun area, approximately 21 kmsouth of Beipiao in the western Liaoning Province of China, andbelongs to the Jianshangou Member (Bed) of the lower YixianFormation (Wang and Zhou, 2008). A late Barremian age for thisfossil-bearing stratum is considered to be well supported byradiometric dating, from 126.1 � 1.7 to 124.6 � 0.1 Ma (Swisheret al., 1999, 2002; Wang et al., 2001b; Chen et al., 2004; Yanget al., 2007); the uppermost beds of Huangbanjigou locality areearly Aptian, 123.3 � 0.5e122.8 � 1.6Ma (Wang et al., 2001a; Yanget al., 2007). Raphidioptera from the Yixian Formation weredescribed by Ren (1994,1997) and Ren et al. (1995); its neuropteranassemblage was analyzed by Makarkin et al. (2012).

The fossils were examined using a Leica M165C dissecting micro-scope and illustrated with the aid of a drawing tube. Photos of allspecimens were taken by Nikon D90 and Leica DFC500 digital cam-eras. All fossil specimens described herein are deposited in the KeyLaboratory of Insect Evolution & Environmental Changes, College ofLife Sciences, Capital Normal University, Beijing (CNUB, D. Ren,curator).

The wing venation of Raphidioptera has been quite poorlystudied. There are three concepts of the raphidiopteran wingvenation (similarly that of other insect orders), differing from eachother mainly in treatment of the median vein: (1) a traditionalconcept considering M as having a common stem, with its anteriorbranches not fused with R or Rs (e.g., Comstock, 1918, figs. 167, 168;Withycombe, 1923, figs. 1, 2); (2) a concept based on the hypothesisof Martynov (1928), which considers M also having a common stembut 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 conceptwhich considers MA and MP as having no common stem, and MAfully fused with R and Rs (e.g., Kukalová-Peck and Lawrence, 2004,fig. 9). We preliminarily accept the concept (2), pending moredetailed examination of raphidiopteran venation, and thereforebroadly follow the terminology of Aspöck et al. (1991), except in thetreatment 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, 1916Suborder: Priscaenigmatomorpha Engel, 2002Family: UncertainGenus Chrysoraphidia gen. nov.

Type species. Chrysoraphidia relicta sp. nov.

Derivationofname. FromChrysopa andRaphidia, genera-groupnames,in reference to superficial resemblance of Chrysoraphidia wings tothose of Chrysopidae (Neuroptera). Gender: feminine.

Diagnosis. Medium-sized raphidiopterans (forewing 12.1e13.5 mmlong). Wings ovoid, relatively broad; pterostigma distinct, elongate,

open proximally, closed distally by veinlet of R; Sc running withinpterostigma to its distal ending, connecting with R by distal obliquecrossvein; MA simple; MP deeply forked, with simple anteriorbranch and distally forked posterior branch; CuA pectinatelybranched, with two branches; CuP simple; two series of gradatecrossveins in radial to mediocubital areas. In forewing, costal areabasally with short ‘ScA’ terminating on C; 1A long, pectinatelybranched, with two branches; 2A simple.

Remarks. The placement of the new genus in Priscaenigmatomor-pha is based on the following character states: the long Scextending the pterostigma; the similar configuration CuA and CuP;RsþMA separating from R near thewing base; and the narrowanalcell between 1A and 2A.

Chrysoraphidia gen. nov. can be distinguished from the twoJurassic genera by its more distinct pterostigma in both wings; Scrunning more closely to C within the pterostigma; not so stronglyzigzagging of Rs and its branches; more distal origin of Rs þ MA;the simple MA in both wings; and its pectinate 1A. The new genusdiffers from Priscaenigma also by relatively broad wings, thedecreasing number of costal veinlets (�10), and the absence ofcrossveins between branches of CuA. The differences between thenew genus and Hondelagia refer to the shape of costal area in theforewings and the branching number of Rs, MP, and 1A (seeWillmann, 1994).

The new genus represents the first record of the basal suborderPriscaenigmatomorpha from China and in general outside Europeand shows that this group occurred not only in the Early Jurassicbut also persisted into the Early Cretaceous.

Chrysoraphidia relicta sp. nov.Figs. 1e7

Derivation of name. From the Latin relictus, abandoned,relinquished.

Type material. Holotype, CNU-RAP-LB-2012001, a well-preservedincomplete specimen in dorsal aspect. Paratype, CNU-RAP-LB-2012002, a rather well-preserved incomplete specimen in lateralaspect. Paratype CNU-RAP-LB-2012003, a poorly preservedincomplete specimen in lateral aspect.

Type locality and horizon. All collected from the Huangbanjigoulocality (41�3604400N, 120�4904800E), Beipiao City, Liaoning Province,China; Early Cretaceous (Yixian Formation).

Diagnosis. As for the genus.

Description. Holotype (Figs. 1e3). Body robust; head, prothorax notpreserved; 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, detailsnot visible. Abdomen rather broad (maximal width 2.5 mm indorsal view); abdominal segments appear rather narrow, fuscous,with lighter intersegmental membrane; terminal segment caudallyrounded.

Forewing ovoid, relatively broad; 12.3 mm long, 4.6 mm wide.Costal area rather narrow basally, then dilated, again narrowedbefore pterostigma, with seven or eight veinlets. Basally, ‘ScA’rather clearly visible in right forewing. Sc long, running rather closeto C within pterostigma, terminating near its distal ending, con-necting with R by distal crossvein (Fig. 2B; distal-most part of Scpoorly preserved). R long, terminating before wing apex, distallywith twowidely spaced short veinlets. Pterostigmawell developed,elongate (ca. 2.5 mm long), dark; area within pterostigma posteriorto Sc slightly paler than area between Sc and C. Six crossveinspresent between R and Rs, forming six radial cells; 2r over twicelength of 1r, about 1.5 times length of 4r; 3r, 5r, 6r together nearly

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

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

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equal length of 1r. Rs þ MA origins from R approximately 1.9 mmfrom wing base. Rs with four simple branches. Two discal cellspresent between proximal branch of Rs, MA; 1dc about twicelength of 2dc. Two parallel series of gradate crossveins presentbetween branches of Rs to CuA: inner series incomplete with fivecrossveins between second branch of Rs and CuA, outer seriescomplete, with eight crossveins between R, CuA. MA simple,proximally coalesced with Rs for distance nearly 1/3 length offorewing. Four crossveins present between MA, MP, forming threemedial cells. Basal part of MP not clearly visible; MP deeply forked,with anterior branch simple, posterior branch forked distally. Threediscoidal cells present between branches of MP; two present be-tweenMP, CuA. Cu dividing into CuA, CuP rather far fromwing base,slightly distal to origin of Rs. CuA zigzagged distally, pectinatelybranched, with two simple branches. Two crossveins present be-tween CuA, CuP. 1A long, pectinately branched, with two (rightwing) or three (left wing) simple branches; 2A simple. Anal cellbetween 1A, 2A narrow. Two crossveins between CuP, 1A; one

crossvein each between 1A, 2A, between 2A, 3A. 3A incompletelypreserved. Membrane probably colourless, transparent except fordark pterostigma.

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

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 sub-costa; Sc(h), hind wing subcosta; sc-r, distal crossvein between Sc and R. Scale barsrepresent 2 mm (A), 1 mm (B).

Fig. 3. Drawings of Chrysoraphidia relicta gen. et sp. nov., holotype CNU-RAP-LB-2012001. A, left forewing; B, right forewing; C, left hind wing; D, right hind wing.Scale bar represents 5 mm (all to scale).

X. Liu et al. / Cretaceous Research 45 (2013) 306e313 309

proximal portion neither forked nor branched. CuA pectinatelybranched. CuP thinner, probably simple. 1A running close to CuP.Membrane probably colourless, transparent except for darkpterostigma.

Paratype CNU-RAP-LB-2012002. (Figs. 4e6). Head and antennaeonly fragmentarily preserved. Pronotum not visible but mostprobably slightly elongate judging from position of head andmesothorax. Mesothorax, metathorax poorly preserved; details notvisible. Legs fragmentary preserved. Abdomen appears very broadin lateral view (possibly compressed post mortem); relatively shorttergites, sternites dark; intersegmental membrane light.

Forewing 11.2 mm long, 4.5 mm wide. Costal area as in holo-type, with seven veinlets. Basally, ‘ScA’ clearly visible in left fore-wing (Fig. 5B). Sc long, running not close to C within pterostigmaand terminating probably near its distal end (distal-most part of Scnot preserved), connecting distally with R by distinct obliquecrossvein (visible in right forewing; Fig. 4B). R long, terminatingbefore wing apex, distally with two widely spaced short veinlets.Pterostigma well developed, elongate, uniformly darkish. Fivecrossveins preserved between R, Rs; radial cell 2r over twicelength of 1r. Rs with five simple branches (right wing). Two discalcells present between Rs, MA; 1dc about 1/5 length of 2dc. Twoparallel series of gradate crossveins present between branches ofRs to CuA: inner series incomplete with six crossveins preserved

between first branch of Rs and CuA, outer series complete, withnine crossveins between R, CuA. MA simple, proximally coalescedwith Rs for distance nearly 1/3 length of forewing. Three cross-veins preserved between MA, MP. Basal part of MP not clearlyvisible but most probably origins from R (i.e., MP appears fusedwith R for considerable distance); two basal crossveins each be-tween R, MP, between MP, CuA. MP deeply forked, with anteriorbranch simple, posterior branch forked distally. Three crossveinspresent between branches of MP, and between MP, CuA. Cudividing into CuA, CuP rather far from wing base, slightly distal toorigin of Rs. CuA zigzagged distally, pectinately branched, withtwo simple branches. Two crossveins present between CuA, CuP.CuP simple. 1A long, pectinately branched, with two simplebranches; 2A simple. 3A probably deeply forked, with crossveinbetween branches (alternately, posterior branch of 3A is 4A;incompletely preserved). Anal cell between 1A, 2A narrow.Crossveins between CuP, 1A not preserved. Membrane probablycolourless, transparent except for dark pterostigma.

Hind wing ca. 9 mm long. Sc long, running not close to C withinpterostigma; terminating near its distal end; distal crossvein notpreserved (Fig. 4B). R long, terminating near wing apex, with twodistal veinlets. Pterostigma well developed, elongate, uniformlydark. Five crossveins preserved between R, Rs. Rs probably withfour simple branches. Two series of gradate crossveins presentbetween branches of Rs and CuA: inner series with two preservedcrossveins between MA, second branch of Rs; outer series withseven crossveins between R, CuA. MA simple. MP basally unclear;deeply forked slightly proximal origin MA from Rs; anterior branchsimple, posterior branch forked distally. CuA pectinately branched,with two simple oblique branches. Basal portion of wings poorlypreserved.

Fig. 6. Drawings of Chrysoraphidia relicta gen. et sp. nov., paratype CNU-RAP-LB-2012002. 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 ofthe paratype CNU-RAP-LB-2012002. A, dry specimen; B, wetted with ethanol. Scale barrepresents 1 mm (both to scale).

X. Liu et al. / Cretaceous Research 45 (2013) 306e313310

Paratype CNU-RAP-LB-2012003 (Fig. 7). Head, pronotum not pre-served. Mesothorax, metathorax, legs poorly preserved, probablycrumpled. Abdomen very short, broad in lateral view; details notvisible.

Forewing ca. 13.5 mm long, 5.3 mmwide. Preserved venation isvery similar to that of holotype right forewing, but 1A is configuredas in left wing. Pterostigma nearly unicolorous. Sc within pter-ostigma; distal crossvein between Sc, R clearly visible; terminationof Sc not preserved; distal sc-r almost perpendicular to Sc (Fig. 7B).

Hind wing venation very poorly preserved. In right wing, Scwithin pterostigma, distal crossvein between Sc, R clearly visible; Scentering wing margin slightly distal to distal sc-r, which isperpendicular to Sc. R with two closely spaced veinlets immedi-ately distal to pterostigma (Fig. 7B).

Remarks. All the specimens lack an ovipositor, and are assumed tobe males.

4. Distinctive morphological characters of Chrysoraphidiagen. nov.

The general venational pattern of the forewing of Chrysoraphidiagen. 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 Priscae-nigma and Hondelagia, and Megaloptera, e.g., the Late Eocene Cor-ydasialis inexpectatus Wichard et al., 2005 (Corydasialidae).Chrysoraphidia gen. nov. is clearly distinguished from these Neu-roptera and Megaloptera taxa by the presence of distinctpterostigma.

The dark pterostigma is present in some taxa of Neuroptera(especially distinct in Mantispidae, Mesochrysopidae, and Ascala-phidae) and all Raphidioptera. The structure of the pterostigma(and in general the costal area) of Chrysoraphidia gen. nov. is su-perficially most similar to those of Mantispinae, the most advancedsubfamily of Mantispidae. However, the pterostigma in thosefamilies of Neuroptera and Raphidioptera was evolved indepen-dently with certainty as their venation is otherwise stronglydifferent.

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 sub-costa; Sc(h), hind wing subcosta; sc-r, distal crossvein between Sc and R. Scale barsrepresent 2 mm (A), 1 mm (B).

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The basal part of the stem ofMP has a tendency to reduction. It isinvisible in the holotype of Chrysoraphidia relicta gen. et sp. nov, butin one of the paratypes it is relatively well visible as originatingfrom R (Figs. 5 and 6). This character state is plesiomorphic here, asit occurs in most taxa of Neuropterida.

The pectinate 1A is characteristic of this genus. This characterstate often occurs in the majority taxa of Neuroptera, and themegalopteran family Corydasialidae, but is absent in all highersnakeflies (Raphidiomorpha), and not detected in the Early JurassicPriscaenigmatidae (although in known specimens of these taxa, theforewing proximal portion is poorly preserved).

The well developed, stick-shaped, erected humeral plate in thehind wing as found in Chrysoraphidia gen. nov. was hitherto un-known in Raphidioptera, nor in Megaloptera. Such a humeral plateis characteristic of some families of Neuroptera, e.g., Psychopsidae,Ithonidae, Parakseneuridae (Tjeder, 1960, fig. 327; Makarkin andMenon, 2007, fig. 2; Yang et al., 2012, figs. 13, 23, 26, 30C). Unfor-tunately, the basal portions of hind wings of the early Jurassicgenera of Priscaenigmatidae specimens are not preserved. Thehumeral plate of other extant Raphidioptera and Megaloptera donot 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) isone of the most characteristic features of this genus. Some extantRaphidioptera (especially Raphidiidae) possess the vestigial,remnant ‘ScA’ in the base of the forewing costal area. In these, ashort convex fold obliquely runs from the wing base near Sc to thecostal margin, where a very short vein lacking hairs is present(interpreted here as the remnant ‘ScA’). A similar ‘ScA’ is shown tobe primitively present in the extant Corydalidae (see Kukalová-Peck, 1991, fig. 6.16). In Chrysoraphidia gen. nov., ‘ScA’ is poorlypreserved in the holotype, but very clearly present in the paratypesCNU-RAP-LB-2012002. ‘ScA’ of extant Raphidioptera, Chrysor-aphidia gen. nov. and Megaloptera (at least Corydalinae) are simi-larly constructed and probably homologous, due to their similarposition in the wing. Such a form of ‘ScA’ is characteristic of some

insect orders (most developed in Orthoptera and Palae-odictyoptera), and is thus plesiomorphic. ‘ScA’ of Neuroptera (whenpresent) is very differently configured, always terminating on Sc,not C (see Yang et al., 2012).

5. Order and family affinity of Chrysoraphidia gen. nov.

This short analysis of the distinctive characters of Chrysoraphidiagen. nov. shows that its venation possesses a mixture of conditionsoccurring in Neuroptera and Raphidioptera, and to lesser extent inMegaloptera. The configuration of ‘ScA’ excludes Neuroptera fromconsideration (see above), and the presence of distinct pterostigmaexcludes Megaloptera. Therefore, Raphidioptera is the most prob-able assignment of this genus based on the presence of the pter-ostigma and ‘ScA’ running to the costal margin. Besides, thepresence of other typical snakefly features (i.e., elongate prothoraxand female ovipositor) in Chrysoraphidia gen. nov. cannot beeliminated until a completely preserved female of this genus isfound. Actually, judging from the position of the head and meso-thorax in the paratype CNU-RAP-LB-2012002 of Chrysoraphidiarelicta sp. nov., the pronotum is likely to be elongated in thisspecies.

The venation of this Cretaceous genus is most similar to that ofthe Early Jurassic genera Priscaenigma and Hondelagia, whichcomprise the family Priscaenigmatidae and the suborder Priscae-nigmatomorpha. Its diagnosis includes the following forewingcharacter states: Sc is long, running within the pterostigma,extending nearly to the wing apex [Sc is short, terminating on Cbefore the pterostigma in all other Raphidioptera]; Rs þ MA origi-nates near the wing base [Rs þ MA originates very far from wingbase in other Raphidioptera]; MP is basally fused with Rs þ MA orCuA (current knowledge does not allow a decision onwhich variantis correct); Cu is continuous with CuA [Cu is continuous with CuP;CuA appears proximally as a crossvein between Cu and M in otherRaphidioptera]; the cell between 1A and 2A is narrow [the cellbetween 1A and 2A is broadly ovoid in other Raphidioptera]. Mostof these conditions are plesiomorphic, at least in Neuropterida,except for the basal structure of MP. The same configuration of basalMP in Priscaenigma and Hondelagia may be interpreted as the onlydistinct synapomorphy of Priscaenigmatidae supporting itsmonophyly. Both variants (MP basally fused with Rs þ MA or CuA)are apomorphic relatively to the plesiomorphic condition whereMP (and thenM) is basally either running freely to the wing base orfused for a short distance with R in most other Neuropterida(including Chrysoraphidia gen. nov.). Although the basal fusion ofMP and CuA is characteristic of all Raphidioptera (if MP is inter-preted as basally fused with CuA), the mode of their fusion is quitedifferent in Priscaenigmatomorpha and other Raphidioptera(Raphidiomorpha): there, MP and CuA run as a single vein towardsthe wing base in the former, and these veins are widely separate atthe wing base in the latter. Therefore, it is likely that the basalfusion of MP and CuA was independently evolved in these groups.

Chrysoraphidia gen. nov. shares all of these character states withPriscaenigmatidae, except the configuration of MP, which is basallynot fused with either CuA or Rs þ MA in this genus. Additionally,the genus rather greatly differs from Priscaenigma and Hondelagiaby other characters, i.e., the origin of Rs þ MA is located relativelyfar from the wing base; many veins are mostly not zigzagged,especially Rs and its branches; and 1A is pectinate. Therefore, theassignment of the new genus to Priscaenigmatidae cannot beconfirmed confidently, but its assignment to Priscaenigmatomor-pha is very probable. The new genus cannot be assigned to a newfamily, as the vast majority (or all) of the diagnostic character statesof this potential new family would be plesiomorphic in Neuro-pterida. As a result, we do not assign Chrysoraphidia gen. nov. to any

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family, pending the description of other genera of Priscaenigma-tomorpha clarifying the situation.

In general, the order affinity of the family Priscaenigmatidae(and Priscaenigmatomorpha) is debatable. Its type genus (Priscae-nigma) was described by Whalley (1985) as belonging to the orderRaphidioptera, to which it has usually been considered to belongsince 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 un-available name). Ponomarenko (1980) assigned it to the Raphi-dioptera. Carpenter (1992) considered this genus as Neuropteraincertae sedis, however, possibly belonging to Raphidioptera.Willmann (1994) reexamined the type material of these twogenera, and concluded they might be regarded as the basal-mostgroup(s) of Raphidioptera. Engel (2002) described the family Pris-caenigmatidae (and the suborder Priscaenigmatomorpha),comprised of these two genera. Priscaenigmatomorpha is consid-ered to be sister of Raphidiomorpha, i.e., all other Raphidioptera(Engel, 2002; Bechly and Wolf-Schwenninger, 2011). However, thisplacement was questioned by Ponomarenko (2002) and Aspöckand Aspöck (2004), who excluded Priscaenigma and Hondelagiafrom Raphidioptera, but did not assign them to any other order.

Aspöck and Aspöck (2004) proposed four autapomorphies ofRaphidioptera (excluding Priscaenigmatidae, i.e., in fact of Raphi-diomorpha): (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 the9th abdominal segment in the male form a ring, and (4) elongationof the female ovipositor. The Sc in the forewing is obviously long inPriscaenigmatidae, and the other three autapomorphies are notpreserved in all known fossils of this family.

Recently, Bechly and Wolf-Schwenninger (2011) accepted theplacement of Priscaenigmatidae in Raphidioptera, and listed thefollowing venational synapomorphies shared by taxa of this order:(A) MP is developed as an oblique veinlet between MA and CuA inthe hind wing; (B) the elongate open cell between the basal parts ofRs and M/MA in the hind wing; and (C) the pterostigma is devel-oped (at least weakly). However, the character states (A) and (B) areplesiomorphic in Neuropterida. The character condition (A) ispresent in most extant Neuropterida species and primitively char-acteristic of all Neoptera (known as the so called ‘M5’; see Ren andMakarkin, 2009; Yang et al., 2012); the character condition (B) is aplesiomorphy of Neuropterida and present in half of the extantRaphidioptera species. Only the condition (C) may be regarded as asynapomorphy shared by all taxa of the order Raphidioptera.

The placement of Priscaenigmatidae in Raphidioptera is thusbased on only one synapomorphy, i.e., the pterostigma is at leastweakly developed. Although the developed pterostigma is alsopresent in some Neuroptera families, it seems plausible to interpretthat this character is independently evolved in various lineages ofNeuropterida rather than symplesiomorphic. Rigorous study on thehigher phylogeny of Raphidioptera should be made in the futurebased on comprehensive and detailed studies of fossil taxa andcorrect 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 fortheir critical review of the manuscript. This research was supportedby the National Natural Science Foundation of China (No. 41271063,31071964, 31230065, 41272006), the National Basic Research Pro-gram of China (973 Program) (2012CB821906), the Foundation forthe Author of National Excellent Doctoral Dissertation of PR China(No. 201178), Project of GreatWall Scholar andKEYproject of Beijing

Municipal Commission of Education (grants KZ201310028033), andChina Geological Survey (1212011120115).

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