Chorilingia (: Grammolingiidae): a new genus of lacewings with four species from the Middle of Inner Mongolia, China

CHAO-FAN SHI, YONG-JIE WANG, QIANG YANG and DONG REN

SHI, C.F., WANG, Y.J., YANG,Q.&REN, D., September 2012. Chorilingia (Neuroptera: Grammolingiidae): a new genus of lacewings with four species from the of Inner Mongolia, China. Alcheringa 36, 309–318. ISSN 0311-5518.

A new grammolingiid genus, Chorilingia containing four new species (C. euryptera, C. parvica, C. translucida and C. peregrina) is described and illustrated from the Jiulongshan Formation at Daohugou, Inner Mongolia, China. The new genus is differentiated mainly on the first branch of vein Rs (Rs1) separating distal to the forks of both CuA and CuP. A key to species of the genus is provided.

Chaofan Shi, Yongjie Wang, Qiang Yang, Dong Ren [[email protected]], Key Lab of Evolution & Environmental Changes, College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, 100048, PR China. Received 31.8.2011, revised 2.11.2011, accepted 25.11.2011.

Key words: Grammolingiidae, Middle Jurassic, Jiulongshan Formation, China.

GRAMMOLINGIIDAE, an extinct family of Neu- lingiidae with ten species have been described from the roptera, has been found only in upper Mesozoic strata Mesozoic of China and Mongolia (Ren 2002, Khra- of northeastern China and Mongolia. This remarkable mov 2010, Liu et al. 2011, Shi et al. 2011; Table 1). The family, together with its contemporaneous relatives genera were mainly differentiated by the termination Kalligrammatidae, Panfiloviidae, Aetheogrammatidae positions of vein 1A, the separation of the most and Saucrosmylinae (an extinct subfamily of Osmyli- proximal branch of Rs, and the position of the forks of dae), represent a special group of Mesozoic Neurop- CuA and CuP. The species are separated by the tera having large body size, complex venation and configuration of veins CuA, CuP, Rs and wing narrow geographic distribution. However, the phylo- markings. geny of these families remains unresolved. To date, Most species of Grammolingiidae in this study only a few phylogenetic analyses have included both were collected from the outcrop at Daohugou, Inner extant and extinct lacewings. Based on phylogenetic Mongolia, China. This deposit is well known for its relationships resolved among some living and fossil rich insect fauna incorporating 19 orders, e.g., lineages of Neuropterida (Grimaldi & Engel 2005, Ephemeroptera (Huang et al. 2008), Odonata (Zhang Engel & Grimaldi 2008), Grammolingiidae has been et al. 2008), Plecoptera (Liu et al. 2007), Blattodea assigned to the suborder Myrmeleontiformia. This (Liang et al. 2009), Heteroptera (Yao et al. 2006),

Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012 family is clearly distinguished by the following Homoptera (Wang et al. 2007), Neuroptera (Ren characters: biareolate costal area in the forewings; 2002), Coleoptera (Tan et al. 2006), Hymenoptera Sc, R1 and Rs stem parallel with the costal margin for (Shih et al. 2009) and Diptera (Zhang et al. 2010). almost the whole length, terminating freely; subcostal Additionally, spiders (Selden et al. 2008), freshwater area and space between R1 and Rs as broad as the diplostracans (Zhang et al. 1987), salamanders (Gao & costal area, with numerous crossveins; cubital area Shubin 2003), feathered dinosaurs (Xu & Zhang 2005), broad, CuA with numerous branches; and two or pterosaurs (Ji & Yuan 2002) and mammals (Ji et al. three rows of cells present between the basal part of 1A 2006) have been recovered from this locality. Accord- and 2A. ing to 40Ar/39Ar and SHRIMP U–Pb dating and Compared with related families of Neuroptera, biostratigraphic interpretations, the fossil-bearing Grammolingiidae appears to be less diverse in beds are of Middle Jurassic age (165 Ma: Shen morphology. To date, only three genera of Grammo- et al. 2003, Chen et al. 2004, Rasnitsyn & Zhang 2004, Gao & Ren 2006, Liu et al. 2006). The Capital Normal University collection hosts ISSN 0311-5518 (print)/ISSN 1752-0754 (online) Ó 2012 Association of Australasian Palaeontologists more than 700 grammolingiid specimens. Here, we http://dx.doi.org/10.1080/03115518.2012.644994 describe a new genus, Chorilingia, with four new 310 CHAO-FAN SHI et al. ALCHERINGA

Species Citations Age (Ma) Epoch Localities Grammolingia boi Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China Leptolingia calonervis Shi et al. (2011) 161.2–167.7 Middle Jurassic Inner Mongolia, China Leptolingia imminuta Liu et al. (2011) 161.2–167.7 Middle Jurassic Inner Mongolia, China Leptolingia jurassica Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China Leptolingia shartegica Khramov (2010) 150.8–145.5 Late Jurassic Mongolia Leptolingia tianyiensis Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China Litholingia eumorpha Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China Litholingia polychotoma Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China Litholingia ptesa Shi et al. (2011) 161.2–167.7 Middle Jurassic Inner Mongolia, China Litholingia rhora Ren (2002) 161.2–167.7 Middle Jurassic Inner Mongolia, China

Table 1. Distribution of the published genera and species of Grammolingiidae.

species: C. euryptera, C. parvica, C. translucida and Diagnosis. Wings narrowly elliptical. Forewing: tri- C. peregrina from Daohugou, Inner Mongolia, China. chosors present at distal part of costal and outer margins; costal region with double rows of cells; Rs with six to nine branches; the most proximal branch of Material and methods Rs separating far from the origin of Rs, about at the The specimens described here were collected from the basal one-third of the wing length, distal to the forks of Jiulongshan Formation of Middle Jurassic age at CuA and CuP; CuA and CuP separating at wing base, Daohugou, Inner Mongolia, China. They were exam- both forked in the proximal one-third to one-fourth of ined using a Leica MZ 7.5 dissecting microscope and the wing length. Hindwing: costal area narrower than illustrated with the aid of a drawing tube attachment. that of forewing, with only one row of cells; CuP Partially magnified images of the specimens were taken pectinate, with fewer branches than forewing. using a Nikon SMZ1000 camera attachment. Line drawings were prepared with Adobe Illustrator CS5 Remarks. According to Lawrence et al. (1991), RA and Adobe Photoshop CS5 graphics software. The (corresponding to R1 in this paper) and CuA are the photographs were taken with a Nikon D100 Digital strongest veins in the wings; the positions and patterns Camera. Specimens of the new species are deposited in of radius and cubitus forking may play an important the Key Lab of Insect Evolution and Environmental part in the wing structure. In Myrmeleontidae, the Changes, the College of Life Sciences, Capital Normal differences in configuration of Rs and CuA in the University, Beijing, China (CNU, Ren Dong, Cura- forewings are usually used for the identification of tor). tribes (Wang et al. 2003). Given the similarity between Myrmeleontidae and Grammolingiidae in wing shape, broad cubital area, numerous branches and dense Venation abbreviation crossveins, we consider that it is reasonable to employ 1A, 2A, 3A ¼ anal veins; CuA ¼ anterior cubitus; the positions of the first fork of the Rs and the forks of CuP ¼ posterior cubitus; MA ¼ anterior media; CuA and CuP as diagnostic features among genera in MP ¼ posterior media; R1 ¼ the first branch of radius; the Grammolingiidae. Therefore, the new genus is Rs ¼ radial sector; Sc ¼ subcosta. The basic terminol- distinguished from the three other genera of Gram-

Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012 ogy of the wing venation follows the schemes of New molingiidae by the first branch of Rs separating from (1989) and Ren (2002). the Rs stem distal to the forks of CuA and CuP, about at one-third of the wing length from base. Within the new genus, most species bear two sets of cells in the Systematic palaeontology costal area along whole length of the wing (with Order NEUROPTERA Linnaeus, 1758 the exception of Chorilingia parvica sp. nov., which has Family GRAMMOLINGIIDAE Ren, 2002 double costal cells only in the basal part of the costal area). This phenomenon also occurs in Leptolingia Chorilingia gen. nov. (Grammolingiidae: L. tianyiensis Ren, 2002). Groups with biareolate costal regions are rare among Neu- Etymology. The new genus name is a combination of roptera, only occurring in Myrmeleontidae (Acantha- the Greek chori- (meaning ‘distant’) and the Chinese clisinae; New 1985). ‘Ling’ (meaning lacewing) referring to the Rs being forked distally in the forewing. The gender is feminine. Stratigraphic and geographic range. The genus is known only from the Jiulongshan Formation (Middle Type species. Chorilingia euryptera sp. nov. Jurassic) at Daohugou, Inner Mongolia, China. ALCHERINGA JURASSIC LACEWINGS FROM CHINA 311

Key to the species of Chorilingia gen. nov. dilated and isometric. Mesonotom with fully devel- oped sclerites, scutum large, scutellum triangular. 1. Forewing with costal cells doubled along all, or Metanotum consisting of complete sclerites, scutelum most of its length ...... 2 slightly smaller and triangular. Abdomen with eight Costal cells doubled in the proximal half of the segments visible (Fig. 3). forewing ...... C. parvica sp. nov. Wings elongated, somewhat falcate, hindwings 2. In the forewing, the MP is forked distal to longer than forewings. Posterior margin convex and separation of MA from Rs ...... 3 outer margin concave in hindwing. All wings hyaline, In the forewing, the MP is forked before with fuscous stripes parallel across the whole wings separation of MA from Rs ...... C. translucida (Figs 1A, 2A–D, 3). The fuscous stripes are repre- sp. nov. sented from the wing base on the forewings, and are 3. MA forked in the distal part of the also present at the apex. Hindwing hyaline at base, forewing ...... C. euryptera sp. nov. stripes ranging across the whole wings and present at MA forked before the separation of the first the apex. Small pale spots are sparsely present within branch of Rs in forewing . . . C. peregrina sp. nov. the stripes. Trichosors present along distal part of costal and outer margin (Fig. 3). Chorilingia euryptera sp. nov. (Figs 1A, B, 2A–D, 3, 4) Crossveins dense over entire forewing. Costal area relatively broad in the basal part, tapering slightly Etymology. The name derives from Latin ‘euryp- toward apex and not broadening in the pterostigmatic terus’, meaning broad wing, referring to the broad area. Costal area biareolate along wing length, cross- wings of the holotype. veins branched near apex. Rs arising from R close to wing base. Both R1 and Rs stem parallel to Sc almost Holotype. CNU-NEU-NN2010513. The specimen along the full wing length. R1 unforked until apex. Rs consists of a relatively complete body with four wings. pectinate, with seven branches, running parallel, The left forewing and hindwing are spread, whereas curved apically towards outer margin, forked distally. the right forewing and hindwing overlap. Venation is MA coalesced basally with Rs for a very short distance evident in all four wings. and then separating from it, running parallel to branches from Rs; dichotomously branched near Paratypes. CNU-NEU-NN2010542, CNU-NEU- distal one-third of wing length. MP forked shortly NN2010724P/C, CNU-NEU-NN2010818P/C, CNU- after separation of MA. The anterior branch of MP NEU-NN2010854P/C, CNU-NEU-NN2010855, spe- dichotomously branched almost at the same level as cimens represented by isolated forewings; body MA. The posterior pectinate branched in the middle not preserved. CNU-NEU-NN2010853P/C, specimen region, with four main branches forming a narrow consisting of almost a complete body with one triangular area. CuA and CuP separated at wing base. forewing and one hindwing overlapping and another CuA forked distally to CuP. CuA with abundant hindwing. CNU-NEU-NN2010684, CNU-NEU- branches terminating in the middle part of the poster- NN2010715, specimens with incomplete forewings. ior margin, forming a triangular area broader than that of MP. CuP dichotomously branched, with fewer Type locality. Daohugou Village, Wuhua Township, branches than CuA. 1A long, arched shortly after Ningcheng County, Inner Mongolia, China. origin, then entering wing margin. 2A slightly shorter than 1A, running parallel to posterior margin. The Type unit. Jiulongshan Formation, Middle Jurassic. basal area between 1A and 2A broad, two to three

Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012 rows of cells present. 3A short, about half length of 2A Diagnosis. Forewing: costal cells doubled along (Fig. 3). whole, or most of wing length; MA separated from Hindwing venation similar to forewing except: Rs stem near wing base, forked in the distal part of narrower costal area with cells simple; first branch of forewing; MP dichotomously branched shortly Rs separated at around the basal one-fifth of wing after MA separated from Rs stem; CuP forked before length, more proximal than fork of CuA, and that of CuA. forewing; CuP parallel to posterior margin and arched before termination (Fig. 3). Measurements. Body 41 mm long (excluding anten- nae). Forewing 40–61 mm long, 11–21 mm wide. Remarks. The coloration of the wings varies Hindwing 45–63 mm long, 19 mm wide as preserved. significantly within the new species. Most specimens have fuscous stripes on the wings (Figs 1A, 2A–D, Description. Moderate to large lacewing. Compound 3), whereas the whole forewing of the paratype eyes large. Antennae filiform with only three segments (CNU-NEU-NN2010542) is fuscous, with no ob- visible. Prothorax shorter than wide, possibly due to vious marking (Figs 1B, 4). However, the highly distortion when preserved. Meso- and metathorax consistent venation pattern confirms that it belongs 312 CHAO-FAN SHI et al. ALCHERINGA Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012

Fig. 1. Photographs of new Chorilingia species from the Jiulongshan Formation, Inner Mongolia, China. A, Photograph of Chorilingia euryptera gen. et sp. nov., holotype CNU-NEU-NN2010513; B, Photograph of Chorilingia euryptera gen. et sp. nov., paratype CNU-NEU- NN2010542; C, Chorilingia parvica sp. nov., holotype CNU-NEU-NN2010584; D, E, Chorilingia translucida sp. nov., holotype CNU-NEU- NN2010839P/C; F, G, Chorilingia peregrina sp. nov., holotype CNU-NEU-NN2010852P/C. Scale bars ¼ 10 mm. ALCHERINGA JURASSIC LACEWINGS FROM CHINA 313 Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012

Fig. 2. Photographs of new Chorilingia species from the Jiulongshan Formation, Inner Mongolia, China. A, B, Photograph of Chorilingia euryptera sp. nov., paratype CNU-NEU-NN2010853P/C; C, D, Photograph of Chorilingia euryptera sp. nov., paratype CNU-NEU- NN2010854P/C; E, F, Photograph of Chorilingia peregrina sp. nov., paratype CNU-NEU-NN2010859P/C; G, Photograph of Chorilingia translucida sp. nov., paratype CNU-NEU-NN2010546. Scale bars ¼ 10 mm. 314 CHAO-FAN SHI et al. ALCHERINGA

Fig. 5. Line drawing of Chorilingia parvica sp. nov., holotype CNU- NEU-NN2010584, showing venation and wing marking. Scale bar ¼ 10 mm.

Description. Forewing narrowly elliptical, posterior margin slightly convex, apex pointed. Trichosors Fig. 3. Line drawing of Chorilingia euryptera gen. et sp. nov., holotype CNU-NEU-NN2010513, showing body, venation and present along distal part of costal and outer margins. wing markings. Scale bar ¼ 10 mm. Crossveins densely distributed over the entire wing. Costal area broad in the basal one-fourth of the wing, with cells doubled. From mid-region to apex, costal area narrower than the basal part, with costal cells simple. Costal veinlets unforked along nearly the whole wing length. Subcostal area slightly broader than costal area, space between R1 and Rs stem almost as broad as subcostal area. Sc, R1 and Rs stem running parallel to costal margin. Rs pectinate, with eight branches running parallel towards outer margin. The first branch of Rs separated, at about basal one- fourth of wing length. At the distal one-third of wing Fig. 4. Line drawing of Chorilingia euryptera gen. et sp. nov., length, the first branch of Rs fused with the second paratype CNU-NEU-NN2010542, showing venation. Scale branch and bifurcating before entering wing margin. bar ¼ 10 mm. All branches of Rs curved apically towards outer margin. MA coalesced basally with Rs for a very short distance then separating from it, running parallel to to this new species. Moreover, the variation in branches from Rs, unforked until terminus. MP colour patterning may be influenced by preserva- forked at base, even before separation of MA from tional processes. Rs. The anterior branch of MP dichotomously branched in distal one-third of wing length. The Chorilingia parvica sp. nov. (Figs 1C, 5) posterior branch pectinate, with three main branches, forming a narrow triangular space. CuA incurved Etymology. The name derives from the Latin ‘par- after origination, then arched and pectinately forked, vus’, meaning small, referring to the small forewing of with three strong main branches forming a broad the species. triangular space. CuP similar to CuA, but dichot- omously branched before the fork of CuA, with much

Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012 Holotype. CNU-NEU-NN2010584. The specimen fewer branches than CuA. 1A simple, terminating comprises a single forewing, body not preserved. before CuP forked. Basal part between 1A and 2A broad. 2A shorter than 1A; 3A undetected (Fig. 5). Type locality. Daohugou Village, Wuhua Township, Ningcheng County, Inner Mongolia, China. Remarks. This new species differs from others in having a narrower costal area with one row of cells, Type unit. Jiulongshan Formation, Middle Jurassic. which are similar to most hindwings of this family. We identified it as a forewing based on the following Diagnosis. Forewing small, costal area biareolate characters: the basal part of the costal area is broad, only in the basal part; MA separated from Rs stem with double rows of cells, whereas the costal area in near wing base, not forked until wing margin; MP the hindwings is narrow along the whole wing length, forked shortly before the separation of MA from Rs with one row of cells; the CuP, dichotomously forked, stem; CuP forked before fork of CuA. is remote from the posterior margin in the basal part but approaches it near the terminus in C. parvica sp. Measurements. Forewing 33 mm long, 12.5 mm nov., whereas the CuP of the hindwings runs parallel wide as preserved. and close to the posterior margin and is convex ALCHERINGA JURASSIC LACEWINGS FROM CHINA 315

slightly before the termination, then curves to the Type locality. Daohugou Village, Wuhua Township, posterior margin, becoming pectinate with branches Ningcheng County, Inner Mongolia, China. parallel to the crossveins; 1A and 2A are well developed in the new specimen, but reduced in most Type unit. Jiulongshan Formation, Middle Jurassic. hindwings of this group. Diagnosis. Forewing: costal area biareolate almost Chorilingia translucida sp. nov. (Figs 1D, E, 2G, along the entire wing length; MA separated from Rs 6A–C) stem significantly distant from the separation of R1 and Rs, nearly at the one-fifth of forewing length; MA Etymology. The name derives from the Latin ‘trans- separated from Rs stem near wing base, forked in the lucidus’, meaning translucent, referring to the translu- distal part of forewing; MP forked before the cent wings with clear venation of the species. separation of MA from Rs stem.

Holotype. CNU-NEU-NN2010839P/C. The speci- Measurements. Body 26 mm long from the apex of men is represented by an almost complete body with the head to the abdomen. Forewing 47 mm long, four overlapping wings. Most venation of one fore- 16 mm wide. Hindwing 48 mm long, 14 mm wide. wing is clear, whereas the other wings are mostly indistinct. Description. The vertex of the head is domed. Compound eyes rounded and large, occupying almost Paratype. CNU-NEU-NN2010546. This specimen half of the head. Antennal sockets distinct and consists of an incomplete body with the basal half of rounded, and mandibles falciform, with blunt apices. the right forewing. Prothorax small. Mesothorax dilated, with small Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012

Fig. 6. Line drawings of Chorilingia translucida sp. nov. A, Holotype CNU-NEU-NN2010839C, showing body, main venation and wing marking; B, Holotype CNU-NEU-NN2010839P, showing details of body; C, Holotype CNU-NEU-NN2010839C, showing details of forewing. Scale bars ¼ 10 mm in Figs 6A, C; 2 mm in Fig 6B. 316 CHAO-FAN SHI et al. ALCHERINGA

scutum visible. Legs slender and hairy, with two pretarsal claws (Figs 6A, B). Wings elongated, somewhat falcate, hindwing narrower than forewing. All wings fuscous, with three parallel hyaline stripes across whole wings. The base and apex of wings fuscous. Trichosors present at distal part of costal and outer margins (Fig. 6A). Forewing crossveins densely distributed over the whole wing. Costal area biareolate along wing length, Fig. 7. Line drawing of Chorilingia peregrina sp. nov., holotype veinlets branched near apex. Rs arising close to the CNU-NEU-NN2010852C, showing venation and wing marking. Scale bar ¼ 10 mm. base of wing, pectinate with more than four branches. The first branch arising far distal to the forks of CuA and CuP. All branches running parallel and curved Chorilingia peregrina sp. nov. (Figs 1F, G, 2E, F, 7) apically towards wing margin, with few distal forks. MA coalesced basally with Rs and separating from it Etymology. The name derives from the Latin ‘pere- at one-fifth of wing length, far distal to the separation grinus’, meaning strange, referring to the rare venation of Rs from R1. MA dichotomously branched at distal of the species. one-fourth of wing length. MP forked near wing base, midway between separations of Rs from R1 and MA Holotype. CNU-NEU-NN2010852P/C, a specimen from Rs. Both anterior branch of MP and MA parallel consisting of a single forewing, body not preserved. to branches of Rs. The anterior branch of MP forked almost at the same level as MA. The posterior branch Paratype. CNU-NEU-NN2010859P/C. The specimen dichotomously forked at the middle of the wing length. is an almost complete forewing; body not preserved. The area formed by branches of the posterior branch of MP small. CuA forked slightly before CuP. CuA Type locality. Daohugou Village, Wuhua Township, incurved after origination, then arched and forked Ningcheng County, Inner Mongolia, China. trichotomously, with numerous branches forming a broad triangular area. CuP dichotomously forked near Type unit. Jiulongshan Formation, Middle Jurassic. outer margin. 1A simple, terminating slightly before forking of CuP. 1A arched after origination and Diagnosis. Forewing: costal area biareolate almost forming a broad space with 2A in the basal part, two along the whole wing length; MA deeply forked at rows of cells present between them. 2A simple, shorter wing base, even before separation of the first branch of than 1A. 3A undetected (Figs 6A, C). Rs from the Rs stem; the anterior branch of MA Hindwing venation similar to forewing, but dif- unforked until the terminus; the posterior branch ferent in having a narrower costal area with one row of dichotomously branched at distal one-fourth of wing cells, narrower than subcostal area; CuP parallel with length; MP forked distal to the separation of the MA posterior margin and convex before termination from the Rs stem. (Fig. 6A). Measurements. Forewing 37–53 mm long, 14 mm Remarks. Thisnewspecieshasseveralspecialand wide. uncommon characters, such as the MA being separa- ted from the Rs stem rather distal to the separation Description. Forewing narrowly elliptical, posterior

Downloaded by [Museum fuer Naturkunde], [Chaofan Shi] at 07:45 22 October 2012 of the R1 and Rs, whereas in all other species of margin slightly convex, apex pointed. Forewing this family, the MA separates shortly after the fuscous, one narrow pale stripe across wing in the separation of the R1 and Rs; the MP is dichot- basal part, small pale spots sparsely present on the omously branched before the separation of MA from wing. Trichosors present along distal part of costal and the Rs stem. Furthermore, some additional features outer margins. Crossveins densely distributed over the of this species are distinct even within the family: (1) whole wing. Costal area biareolate along most of wing the posterior branch of the MP is dichotomously length, crossveins rarely forked. Costal area broad in forked, and the space formed by the branches of the the basal part, narrowed slightly towards apex and not posterior branch of the MP is much smaller than broadening in the pterostigmatic area. Sc, R1 and Rs that in any other species; (2) the CuA is forked stem parallel to costa. Subcostal space and area trichotomously, which is rare in Grammolingiidae between R1 and Rs stem the same breadth as costal (to date, this feature has only been found in area. Rs pectinate with seven branches running Litholingia polychotoma Ren, 2002 and Chorilingia parallel and curved apically towards outer margin. translucida sp. nov.); (3) the CuA is forked slightly The first branch of Rs arising after one-third of wing before the CuP and the 1A vein terminates shortly length, far distal to the forks of CuA and CuP. MA before the CuP is forked. coalesced basally with Rs and separating shortly after ALCHERINGA JURASSIC LACEWINGS FROM CHINA 317

rise of Rs. MA deeply forked in the basal part of GAO,K.Q.&REN, D., 2006. Radiometric dating of ignimbrite from forewing, about midway between separation of MA Inner Mongolia provides no indication of a post-Middle Jurassic age for the Daohugou beds. Acta Geologica Sinia 80,42–45. from Rs and first fork of Rs. The anterior branch of GAO, K.Q. & SHUBIN, N.H., 2003. Earliest known crown-group MA incompletely preserved, no fork evident before its salamanders. Nature 422, 424–428. loss near the wing margin. The posterior branch of GRIMALDI, D.A. & ENGEL, M.S., 2005. The Holometabola. In Evolution of the .GRIMALDI, D.A. & ENGEL M.S., eds, MA dichotomously forked at distal one-fourth of wing Cambridge University Press, Cambridge, 335–356. length. Both branches of MA parallel to branches of HUANG, J.D., REN, D., SINITSHENKOVA, N.D. & SHIH, C.K., 2008. Rs. MP forked slightly distal to separation of MA New fossil mayflies (Insecta: Ephemeroptera) from the Middle from Rs stem, but before fork of MA. The anterior Jurassic of Daohugou, Inner Mongolia, China. Insect Science 15, 193–198. branch of MP forked dichotomously in the distal part JI,Q.&YUAN, C.X., 2002. Discovery of two kinds of proto- of forewing, before the fork of the posterior branch of feathered pterosaurs in the Mesozoic Daohugou Biota in the MA. The posterior branch of MP pectinate in the mid- Ningcheng region and its stratigraphic and biologic signifi- region of wing length, with three main branches, cance. Geological Review 48, 221–224. (in Chinese with English abstract) forming a narrow triangular space. CuA and CuP JI, Q., LUO, Z.X., YUAN, C.X. & TABRUM, A.R., 2006. A swimming separating at wing base. CuA dichotomously forked mammaliaform from the Middle Jurassic and ecomorphological at basal one-third of wing length, with numerous diversification of early mammals. Science 311, 1123–1127. branches, forming a triangular space broader than KHRAMOV, A.V., 2010. A new lacewing (Insecta: Neuroptera: Grammolingiidae) from the Upper Jurassic of Mongolia. that of MP. CuP dichotomously forked before fork Paleontological Journal 44, 188–191. of CuA, less branches and smaller space than CuA. LAWRENCE, J.F., NIELSEN, E.S. & MACKERRAS, I.M., 1991. Skeletal 1A long, arched after origination, forming a broad anatomy and key to orders. In The Insects of Australia, 2nd space with 2A in the basal part. Two rows of cells edition. NAUMANN, I.D., CARNE, P.B., LAWRENCE, J.F., NIELSEN, E.S., SPRADBERY, J.P., TAYLOR, R.W., WHITTEN,M.J.& present in the basal area between 1A and 2A. 1A LITTLEJOHN, M.J., eds, Melbourne University Press, Carlton, terminating distal to fork of CuP. 2A slightly Victoria, 14–17. shorter than 1A, running parallel to posterior LIANG, J.L., VRSˇ ANSKY´, P., REN,D.&SHIH, C.K., 2009. A new Jurassic carnivorous cockroach (Insecta, Blattaria, Raphidio- margin. 3A distinct and short (Fig. 7). mimidae) from the Inner Mongolia in China. Zootaxa 1974, 17– 30. Remarks. All previously described species of Gram- LINNAEUS, C., 1758. Systema naturæ per regna tria naturæ, secundum molingiidae bear a vein MA that is dichotomously classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1, 10th edition, Salvius, Stockholm, 1– forked in the distal part, about three-fourths to two- 824. thirds of the forewing length. The species described LIU, Y.Q., LIU, Y.X., JI, S.A. & YANG, Z.Q., 2006. U–Pb zircon age here is the first to show the MA forked in the basal for the Daohugou Biota at Ningcheng of Inner Mongolia and one-fourth of the forewing, adding one more second- comments on related issues. Chinese Science Bulletin 51, 2634– 2644. ary longitudinal vein to the basal part of the forewing. LIU, Y.S., REN, D., SINITSHENKOVA, N.D. & SHIH, C.K., 2007. 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