This Article Appeared in a Journal Published by Elsevier. the Attached Copy Is Furnished to the Author for Internal Non-Commerci

Home , Berothidae, Grammolingiidae

This article appeared in a journal published by Elsevier. The attached copy is furnished to the author for internal non-commercial research and education use, including for instruction at the authors institution and sharing with colleagues. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/copyright Author's personal copy

Cretaceous Research 30 (2009) 1217–1222

Contents lists available at ScienceDirect

Cretaceous Research

journal homepage: www.elsevier.com/locate/CretRes

Ascalochrysidae – a new lacewing family from the Mesozoic of China (Insecta: Neuroptera: Chrysopoidea)

Dong Ren a, Vladimir N. Makarkin b,* a College of Life Science, Capital Normal University, Beijing 100048, China b Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russian Federation article info abstract

Article history: Ascalochrysa megaptera gen. et sp. nov. is described from the Yixian Formation (Lower Cretaceous) of Received 3 March 2009 China. It is treated as belonging to the new family Ascalochrysidae fam. nov., closely related to Meso- Accepted in revised form 19 June 2009 chrysopidae but easily distinguished from it in the hindwing venation features: e.g., numerous subcostal Available online 1 July 2009 crossveins; branches of Rs widely, irregularly spaced and deeply branched; crossvenation rich, irregular; convex vein-like fold (‘M5’) before concave CuA present. The presence of well-developed ‘M5’ in the hind Keywords: wing is interpreted to be plesiomorphic condition within the order. Ascalochrysidae are considered the Ascalochrysidae sister group of Mesochrysopidae (s.l.). Mesochrysopidae Cretaceous Ó 2009 Elsevier Ltd. All rights reserved. Yixian Formation China

1. Introduction 2. Materials and methods

The taxonomic diversity of the Order Neuroptera was greatest in The specimen examined was collected near Chaomidian Village the Mesozoic. Several new neuropteran families have been in Liaoning Province, NE China from the deposits of the Yixian described from this era in the last decade, i.e., Makarkiniidae Formation. All photographs were taken with a Nikon Digital (Martins-Neto, 2000), Grammolingiidae (Ren, 2002), Limaiidae, Camera DXM1200C attached to a Leica MZ12.5 stereomicroscope. Tachinymphidae, Liassochrysidae (Nel et al., 2005), Aetheog- Drawing was done directly from the fossil with a camera lucida rammatidae (Ren and Engel, 2008a). Although at least two of these mounted on a Leica MZ12.5 stereomicroscope. are probably synonyms of other families (e.g., Makarkiniidae is We follow here the traditional (sensu Wootton, 2003) venational a probable synonym of Kalligrammatidae: Makarkin and Archibald, terminology of Comstock (1918) with the current interpretation of 2003; Liassochrysidae is a synonym of Mantispidae: Wedmann and Oswald (1993), Makarkin and Menon (2005), Archibald and Makarkin, 2007), the description of new Mesozoic families should Makarkin (2006) and Wedmann and Makarkin (2007). Terminology be expected in the future. of wing spaces follows Oswald (1993). In this paper we describe a new genus and species of a large Venational abbreviations are as follows: Sc, subcosta; R1, ﬁrst neuropteran from the Lower Cretaceous of China. We interpret this branch of radius (R); Rs, radial sector; Rs1, most proximal branch of as a representative of the new family Ascalochrysidae fam. nov. Rs; M, media; MA, media anterior; MP, media posterior; CuP, poste- belonging to Chrysopoidea. This superfamily was recently estab- rior cubitus; CuA, anterior cubitus; 1A–3A, ﬁrst to third anal veins. lished to include several families, mainly extinct (Nel et al., 2005). Institutional abbreviation. CNU, Capital Normal University, We discuss the systematic position of Ascalochrysidae fam. nov. Beijing, China. among chrysopoids, the family composition of this superfamily, and some important character states found in Ascalochrysa gen. 3. Locality and stratigraphy nov., e.g., the presence of a convex vein-like fold before CuA. Fossils occurring in the Yixian and Jiufotang Formations of northeastern China constitute the Jehol Biota, an assemblage which is thought to have been widely distributed in eastern Asia during * Corresponding author. the Early Cretaceous (Chen, 1988; Chen et al., 1998; Zhou et al., E-mail address: [email protected] (V.N. Makarkin). 2003). The Yixian Formation consists mainly of lacustrine

1218 D. Ren, V.N. Makarkin / Cretaceous Research 30 (2009) 1217–1222 sediments intercalated with volcaniclastics (Ren et al., 1995). It is crossveins throughout wing remigium numerous, irregular, not well known by a large number of exceptionally well-preserved forming gradate series or regular reticulation. fossils, including numerous insects, conchostracans, ostracods, gastropods, bivalves, fish, salamanders, turtles, lizards, feathered Comparison. The venational pattern of Ascalochrysidae fam. nov. dinosaurs, primitive birds (Confuciusornis Hou et al. and Liao- is most similar to that of Mesochrysopidae (s.l.) and strongly ningornis Hou) and angiosperm (Archaefructus Sun et al.) (Sun et al., different from that of all other families of the order. The new family 1998; Hou et al., 1999; Ding et al., 2001; Zhou et al., 2003). Some shares with Mesochrysopidae (s.l.) the following hind wing apo- Yixian Neuroptera have been described by Ren et al. (1995), Ren morphic character states: (1) the reduction of CuP to a part of and Guo (1996), Ren and Yin (2002), Ren (2003a), Nel et al. (2005) apparent basal crossvein between CuA and 1A; (2) the entire loss and Ren and Engel (2008a, b). Palaeobotanical data from paly- (or strong reduction) of 2A and 3A as a result of the reduction of nomorphs and macrofossils provide contradictory evidence on the cubito-anal area as a whole; (3) the loss or poor development of the climatic conditions in the Jehol times. On the one hand, the data humeral lobe and frenulum of coupling apparatus (the latter shared indicate a warm and humid Yixian climate (Ding et al., 2001), and with Chrysopidae). Other shared apomorphic character states are on the other, many plants suggest arid or semi-arid climatic the absence of trichosors, nygmata and the basal sinuous crossvein conditions (Zhou et al., 2003; Barrett and Hilton, 2006). Fu¨ rsich r-m, and the distal fusion of Sc and R1, but these features occur in et al. (2007) suggest that the climate was semi-arid with dry and other families of different lineages. Moreover, both families have wet seasons. distinctive structure of M and CuA. Ascalochrysidae fam. nov. are The age of the Yixian Formation was controversial for a long easily distinguished from Mesochrysopidae (s.l.) by the following time, with estimates varying from Late Jurassic to Early Cretaceous features [character states of Mesochrysopidae are given in (Sha, 2007a,b for summary). Now, an Early Cretaceous age of the brackets]: Sc þ R1 entering wing margin well beyond apex [at or Yixian Formation is considered to be well supported by radiometric (rarer) before wing apex]; subcostal crossveins numerous [only one dating; using different radioactive decay series, from 133.46 0.18 basal crossvein at most]; crossvenation over remigium irregular for the lowest beds (Chen et al., 2004), 126.1 1.7 to 124.6 0.1 Ma [regular, forming gradate series or reticulation]; ‘M5’ present for the second Member containing insect fossils (Swisher et al., [absent]; M branched very close to the wing base [far to the wing 1999, 2002; Wang et al., 2001; Chen et al., 2004; He et al., 2006)to base]; Banksian folds absent [two of these folds present]. 121 0.2 Ma for overlying lava layers and intrusive volcanics Superficially, the genus Ascalochrysa gen. nov. resembles the (Smith et al., 1995), i.e., ranging from the late Hauterivian to early genera of Ascalaphidae in general appearance, but they are easily Aptian (geologic time scale of Gradstein et al., 2005). Recently, an distinguished from the new genus in the absence of subcostal Early Cretaceous age of the Yixian Formation was supported by new crossveins and ‘M5’, the distal position of the origin of Rs, and the evidence (Li and Batten 2007; Yang et al., 2007; Zhu et al., 2007). configuration of M. Determination of the exact age of the Yixian Formation is very Comments on some character states.Scþ R1 entering wing important as it bears great evolutionary consequences, for example, margin beyond apex as found in Ascalochrysa gen. nov. is charac- the origin of angiosperms and the early radiation of birds (Barrett, teristic of the Myrmeleontoidea families (i.e., Myrmeleontidae, 2000). Some palaeontologists interpret this formation (treated as Ascalaphidae, Nymphidae, Palaeoleontidae, Nemopteridae, Babin- Early Cretaceous in age) as a refugium of Jurassic terrestrial relicts skaiidae) for which this character state is certainly a synapomor- in East Asia, based on the occurrences of compsognathid thero- phy. This condition is probably convergent in Ascalochrysidae fam. pod dinosaurs, ‘rhamphorhynchoid’ pterosaurs, and primitive nov. and Myrmeleontoidea. However, Ponomarenko (1992) and mammalians (Luo, 1999; Matsukawa et al., 2006). Makarkin and Menon (2005) suggested that the chrysopoid lineage The first author (DR) is of the opinion that at the current stage of (Mesochrysopidae and Chrysopidae) and myrmeleontoid lineage knowledge, the Yixian Formation should be considered Late might have had a common ancestor, and form a monophylum. So, Jurassic to earliest Early Cretaceous in age. there is a probability that this condition may be a synapomorphy of the two lineages. The irregular and widely spaced branches of Rs is an unusual 4. Systematic paleontology feature occurring very rarely in Neuroptera. In most taxa of the order, these branches are more or less regular pectinate, including Class: Insecta Linnaeus, 1758 all Permian and Triassic taxa; exclusions are those taxa which Order: Neuroptera Linnaeus, 1758 possess distinctly derived venation (e.g., Allopterus Zhang, 1991). Superfamily: Chrysopoidea Schneider, 1851 The condition as found in Ascalochrysa gen. nov. may be compared Family: Ascalochrysidae fam. nov. to some extent with only a few osmylids (e.g., Gumillinae: Menon and Makarkin, 2008) and undescribed taxa from the Middle Type and only genus. Ascalochrysa gen. nov., from the Lower Jurassic of China (Ren, 2003b). Cretaceous of China. The CuP in this genus is strongly reduced. We interpret it to be represented by only its basal-most portion fusing distally with Diagnosis. Large neuropterans (hind wing approximately 60 mm a basal crossvein between CuP and 1A (Fig. 3) in a similar way as long) of myrmeleontoid appearance, distinguished from other seen in some recent Hemerobiidae (e.g., Hemerobius Linnaeus, families by the following combination of hind wing character 1758; see Oswald, 1993: fig. 34). Makarkin and Menon (2005) had states: trichosors and nygmata absent; Sc and R1 fused distally; the opinion that CuP is absent in Mesochrysopidae (s.l.), whereas subcostal crossveins numerous, rather regular; Sc þ R1 enter wing Nel et al. (2005) considered this to be present as a basal ‘crossvein’ margin well beyond apex; Rs with irregular, widely spaced between CuA and 1A. Actually, the structure of the hind wing CuP in branches, some deeply forked; no distinct longitudinal folds Mesochrysopidae (s.l.) is probably the same as in Ascalochrysa gen. throughout wing; M branched very close to wing base; convex nov. CuP is often lost in the hind wings of other Hemerobiiformia: vein-like fold anterior to CuA (‘M5’) well developed, not reaching in some Hemerobiidae, many Berothidae, and all Mantispidae wing margin; CuA relatively short, concave; CuP almost entirely except Symphrasinae (Lambkin, 1986). reduced and represented by part of apparent crossvein between Ascalochrysa gen. nov. possesses several widely spaced, oblique CuA and 1A; 1A well-developed; 2A and 3A apparently lost; crossveins forming a medial longitudinal series slightly suggestive Author's personal copy

D. Ren, V.N. Makarkin / Cretaceous Research 30 (2009) 1217–1222 1219 of the ‘Banksian line’ of Mesochrysopidae and Myrmeleontidae but genus-group names, from the Greek chryseos or chrysos, gold), in not accompanied by a fold. The hind wings of Mesochrysopidae reference to superficial resemblance of the type species to ascala- possess two well-developed longitudinal folds throughout the phids and its actual affinity with chrysopoids. Gender feminine. wing, which are converged distally (‘Banksian folds’: Menon and Makarkin, 2008), see for example Nel et al. (2005: figs. 8.3, 8.5). Diagnosis. As for the family. These folds may be reduced in the most advanced (specialized) genera (e.g., Allopterus). Description. As for Ascalochrysa megaptera gen. et sp. nov.

Sclerotized convex fold before CuA in hind wings (‘M5’). In the hind Ascalochrysa megaptera gen. et sp. nov wing of some endopterygotes there is a strongly convex sclerotized Figs. 1–3 (vein-like) fold before the concave CuA, which does often not reach the wing margin. It seems to occur mainly in mecopteroid insects, in Derivation of name. The specific epithet from Greek megas, large, particular in the hind wing of the earliest Holometabolous insect and pteron [ ¼ pteryx], wing, in reference to large size of the species. Westphalomerope maryvonneae Nel et al., 2007 (Mecopteroids: Pro- Material. Holotype CNU LB2006001-1 (part), LB2006001-2 tomeropidae) from the Upper Carboniferous (Lower Pennsylvanian) (counterpart). A beautifully preserved, almost complete hind wing. (Nel et al., 2007), in ‘mecopteroid-like’ insects from Lower Permian of China, Liaoning Province, Beipiao City, near Chaomidian Village; Obora, Czech Republic (Kukalova´-Peck and Willmann,1990: figs. 4, 5, Yixian Formation, Lower Cretaceous. 14), in many Permian Mecoptera and Trichoptera (Novokshonov, 1997), and in Thaumatomerope Rasnitsyn in Ponomarenko et Ras- Diagnosis. As for the genus. nitsyn, 1974 (Mecoptera: Meropeidae) from the Triassic of Madygen Formation, Kyrgyzstan (Ponomarenko and Rasnitsyn, 1974). Descriptions. Hind wing 57 mm long, 17 mm wide; elongate sub- This vein-like fold is usually named M5 (Novokshonov,1997). The triangular, with distinct tornus nearly at mid-point of wing. Costa term M5 was introduced by Tillyard (1919) who considered the Media very prominent all around wing. Trichosors absent. Nygmata not of ‘mecopteroid’ orders (including Neuropterida) to be originally five detected, probably absent. Costal space rather broad, slightly branched. M5 in the extant taxawas identified by him in most cases as dilated in proximal portion, and strongly expanded distally. Pter- a basal crossvein-like vein fused distally with CuA. This hypothesis ostigma not detected (this area not preserved), probably absent. All was rejected by Kukalova´-Peck (1991) who considered ‘M5’ as one of subcostal veinlets simple, straight, rather widely spaced; humeral the ‘‘secondary veinal elements . frequently re-formed from veinlet crossvein-like, neither branched nor recurrent. Sc fused membrane.’’ (p. 149). Various authors are now in disagreement with with R1 apically. Sc þ R1 entering wing margin apparently well each other about the nature of M5: see for example a discussion on beyond apex (not completely preserved). Some (?all) apical-most the issue by Be´thoux (2007, 2008) and Rasnitsyn (2007). We name veinlets of Sc and Sc þ R1 deeply forked (one with additional here a convex fold before CuA as ‘M5’ irrespective of what it turns out marginal fork), several irregularly arranged crossveins between ultimately to be, a branch of M or a secondary veinal element. them. Subcostal space relatively broad, with numerous subcostal Anyway, in the hind wing of the Upper Carboniferous Westphalome- crossveins (>30) arranged regularly. R1 space (between R1 and Rs) rope maryvonneae a strongly convex vein-like fold before CuA is approximately as broad as costal space, maximum width at its mid- clearly visible as a branch of M reaching the wing margin (see Nel point. Crossveins in R1 space numerous (>31), rather regularly et al., 2007: fig. 1A, although the authors treat it as a fold). spaced. Rs concave, smooth (only slightly zigzagged in distal part), In the hind wings of extant Neuroptera ‘M5’ is detected only in originating at acute angle; its length from origin to origin of Rs1 Nevrorthidae. In Nipponeurorthus pallidinervis Nakahara, 1958 this relatively short (ca. 4 mm). Branches of Rs widely and irregularly is a sclerotized, vein-like structure lacking hairs, originating at M spaced, 9–10 in number, dichotomously branched. Basal-most part but not reaching the wing margin (VM, pers. obs., see also Naka- of M not clearly preserved, apparently not fused with R basally. hara, 1958: pl. 8, figs. 1, 2). ‘M5’ seems to occur in more ancient M forked very close to wing base. MA concave, dichotomously groups. The Nevrorthidae are considered the sister group of other branched, similar to branching of proximal branches of Rs; extant Neuroptera (Aspo¨ck et al., 2001; Haring and Aspo¨ck, 2004). entering margin nearly at wing mid-point. MP convex, occupying In the beautifully preserved hind wing of Permopsychops saurensis large area; its posterior trace with 4 branches directed anteriorly. Novokshonov et Vilesov (Permithonidae) from the Upper Permian Convex sclerotized (vein-like) fold anterior to CuA (‘M5’) well locality of Karaungir, Kazakhstan ‘M5’ is well-developed as a true developed, not reaching wing margin; originated apparently at M posterior branch of M basally continuing distally as an unsclero- before its fork (this portion is poorly preserved). CuA short, tized convex fold before CuA (Vilesov and Novokshonov, 1994: fig. concave, with single marginal fork. CuP reduced, represented by 3; Novokshonov, 1997: pl. 1, fig. 1). In the Permian/Triassic Per- proximal part of apparent crossvein between CuA and 1A. 1A well- morapisma gori Ponomarenko et Shcherbakov, 2004 (Permithoni- developed, convex, with 4 simple branches. 2A, 3A not detected, dae) a convex furrow runs anterior to the concave CuA probably lost. Crossvenation throughout wing dense, irregular, not (Ponomarenko and Shcherbakov, 2004: p. S198). Therefore, the forming gradate series or reticulation, except for a medial longi- presence of a well-developed ‘M5’ in the hind wing of Ascalochrysa tudinal series formed by 5 (preserved) oblique crossveins con- gen. nov. may be interpreted as a plesiomorphic character state. nected branches of Rs. ‘M5’ is not detected in the closely related Mesochrysopidae. In particular, it is certainly absent in the well-preserved hind wing of 5. Discussion Tachinymphes ascalaphoides (Ponomarenko, 1992: fig. 3). The current higher classification and phylogeny of the extant Genus Ascalochrysa gen. nov. Neuroptera is mainly based on characters of larvae, genitalia of imagoes (e.g., Aspo¨ck, 1995, 2001; Aspo¨ck et al., 2001), and Type and only species. Ascalochrysa megaptera gen. et sp. nov. molecular data (Winterton, 2003; Haring and Aspo¨ck, 2004). The incorporation of the data from the fossil taxa into such phylogenies Derivation of name. Ascalo- (from Ascalaphidae, a neuropteran is problematic, as the majority of these taxa are represented only by family-group name), and -chrysa (a traditional ending of chrysopid wings. There are only a few, very preliminary phylogenetic Author's personal copy

1220 D. Ren, V.N. Makarkin / Cretaceous Research 30 (2009) 1217–1222

Fig. 1. Ascalochrysa megaptera gen. et sp. nov., photograph of the holotype. A, part (LB2006001-1). B, counterpart (LB2006001-2). Scale bar represents 5 mm. hypotheses which comprise both extant and extinct families; the The superfamily Chrysopoidea was erected by Nel et al. (2005) most recent of those are of New (1989, redrawn from Schlu¨ ter, in the superorder Hemerobiiformia. The phylogenetic relationships 1986), Grimaldi and Engel (2005), and Engel and Grimaldi (2008). between chrysopoid groups proposed by these authors are as Given that the majority of neuropteran taxa are extinct, ‘phyloge- follows: (Cratochrysa Martins-Neto, 1994 þ (Liassochrysidae netic integration’ of the entire order will probably be done in the þ (Paralembochrysa Nel et al., 2005 þ ((Allopteridae þ Meso- very distant future. chrysopidae þ Tachinymphidae þ Mesotermes Haase, 1890) þ (Lim- Formerly, the neuropteran families were classified into aiidae þ Chrysopidae))))). a number of superfamilies. Their number and familial composition In our opinion, however, the family Liassochrysidae does not are different for various authors. For example, both Martynova belong to this superfamily; it was recently considered to be (1949) and Henry (1982) recognized six superfamilies in Neuro- a synonym of Mantispidae (Wedmann and Makarkin, 2007). The ptera but their names and the composition (especially) are very family affinity of Cratochrysa is not entirely unknown yet, but it is different: Ithonoidea, Coniopterygoidea, Polystoechotoidea, Psy- surely distant from Chrysopoidea. Paralembochrysa belongs to chosidoidea, Hemerobioidea, Myrmelentoidea in Martynova Chrysopidae with certainty. The taxon [Limaiidae þ Chrysopidae] is (1949), and Hemerobiiformia (Ithonoidea, Coniopterygoidea, accepted here to be a single family, Chrysopidae, as it has been Mantispoidea, Osmyloidea, Hemerobioidea), Myrmeleontiformia previously treated (e.g., Makarkin, 1997). Nel et al. (2005) found ‘‘no (Myrmelentoidea) in Henry (1982). Now, the order is divided into clear autapomorphy that would characterize the Limaiidae’’ (p. 54). three suborders (Nevrorhtiformia, Myrmeleontiformia and Hem- The taxon [Allopteridae þ Mesochrysopidae þ Tachinymphidae þ erobiiformia), and a superfamily category is not usually used (e.g., Mesotermes]ofNel et al. (2005) was treated by Makarkin & Menon Haring and Aspo¨ck 2004; Grimaldi and Engel 2005, see however (2005) as the family Mesochrysopidae (s.l.). Both Makarkin and Engel and Grimaldi 2008). However, until a coherent phylogeny of Menon (2005) and Nel et al. (2005) considered this taxon mono- the whole order is worked out, the recognition of groups of closely phylous. The classification of Mesochrysopidae (s.l.) is outside the related families (i.e., superfamilies) is justified, especially when scope of this paper, but at least Allopteridae are treated by Martins- dealing with fossil taxa. Neto et al. (2008) as a separate family.

Fig. 2. Ascalochrysa megaptera gen. et sp. nov., holotype, drawing of hind wing. Scale bar represents 5 mm. Author's personal copy

D. Ren, V.N. Makarkin / Cretaceous Research 30 (2009) 1217–1222 1221

Acknowledgements

We thank James Jepson (University of Manchester, UK) for cor- recting the English. This research is supported by the National Natural Science Foundation of China (No.30430100, 40872022), the Nature Science Foundation of Beijing (No.5082002) and Scientiﬁc Research Key Program and PHR Project of Beijing Municipal Commission of Education.

References

Archibald, S.B., Makarkin, V.N., 2006. Tertiary giant lacewings (Neuroptera: Poly- stoechotidae): revision and description of new taxa from western North America and Denmark. Journal of Systematic Palaeontology 4, 119–155. 307 [errata]. Aspo¨ck, U., 1995. Neue hypothesen zum System der Neuropterida. Mitteilungen der Deutschen Gesellschaft fu¨r Allgemeine und Angewandte Entomologie 10, 633–636. Aspo¨ck, U., 2001. Phylogeny of the Neuropterida (Insecta: Holometabola). Zoologica Scripta 31, 51–55. Aspo¨ck, U., Plant, J.D., Nemeschkal, H.L., 2001. Cladistic analysis of Neuroptera and their systematic position within the Neuropterida (Insecta: Holometabola: Neuropterida: Neuroptera). Systematic Entomology 26, 73–86. Barrett, P.M., 2000. Evolutionary consequences of dating the Yixian Formation. Trends in Ecology and Evolution 15, 99–103. Fig. 3. Ascalochrysa megaptera gen. et sp. nov., holotype (counterpart), basal portion of Barrett, P.M., Hilton, J., 2006. The Jehol Biota (Lower Cretaceous, China): new hind wing showing CuP; cu-a1, basal crossvein between CuP and 1A. Scale bar discoveries and future prospects. Integrative Zoology 1, 15–17. represents 1 mm. Be´thoux, O., 2007. Archaeorthoptera wing venation nomenclature: a reply to Gor- okhov. Paleontological Journal 41, 338–340. Be´thoux, O., 2008. Groundplan, nomenclature, homology, phylogeny, and the Therefore, the Chrysopoidea of Nel et al. (2005) comprises only question of the insect wing venation pattern. Alavesia 2, 219–232. two families: Mesochrysopidae (s.l.) and Chrysopidae (s.l.). Asca- Chen, P.J., 1988. Distribution and migration of the Jehol Fauna with reference to lochrysidae fam. nov. is most closely related to Mesochrysopidae non-marine Jurassic-Cretaceous boundary in China. Acta Palaeontologica Sinica 27, 659–683 [in Chinese, English abstract]. (s.l.) with certainty, and consequently it also belongs to this Chen, P.J., Dong, Z.M., Zhen, S.N., 1998. An exceptionally well-preserved theropod superfamily. The main putative synapomorphies in the hind wing dinosaur from the Yixian Formation of China. Nature 391, 147–152. venation of these two families are (1) the strong reduction of CuP Chen, S.W., Jin, C.Z., Zhang, Y.P., Zhang, L.D., Guo, S.Z., 2004. Discussion on the structural-volcanic activities and biological events during the Early Cretaceous and (2) the loss (or strong reduction) of 2A and 3A (other shared in the Sihetun Area, Liaoning Province, China. Tikhookeanskaya Geologiya 23 features see above, in comparison section). The Ascalochrysidae (3), 52–59. fam. nov. are probably the sister group of the Mesochrysopidae (s.l.) Comstock, J.H., 1918. The Wings of Insects. Comstock Publication Company, Ithaca, NY, 430 pp. [ ¼ the taxon Allopteridae þ Mesochrysopidae þ Tachinymphidae], Ding, Q.H., Zhang, L.D., Guo, S.Z., Zhang, C.J., Peng, Y.D., Jia, B., Chen, S.W., Xing, D.H., as the venation of Chrysopidae (s.l.) differs from that of these two 2001. The stratigraphic sequence and fossil-bearing horizon of the Yixian families quite strongly (e.g., in particular, CuP, 2A, and 3A are well Formation in western Liaoning, China. Geology and Resources 10, 193–198 [in Chinese, English abstract]. developed in the chrysopid hind wing). Engel, M.S., Grimaldi, D.A., 2008. Diverse Neuropterida in Cretaceous amber, with Some character states may be provisionally considered autapo- particular reference to the paleofauna of Myanmar (Insecta). Nova Supplementa morphies of Ascalochrysidae fam. nov: (1) Sc þ R1 entering wing Entomologica 20, 1–86. margin well beyond apex (probably homoplastic with Myrme- Fu¨ rsich, F.T., Sha, J.G., Jiang, B.Y., Pan, Y.H., 2007. High resolution palaeoecological and taphonomic analysis of Early Cretaceous lake biota, western Liaoning lontoidea), and (2) numerous subcostal crossveins (probably homo- (NE-China). Palaeogeography, Palaeoclimatology, Palaeoecology 253, 434–457. plastic with some families in other superfamilies). On the other hand, Grimaldi, D.A., Engel, M.S., 2005. Evolution of the Insects. Cambridge University Ascalochrysidae fam. nov. possess some plesiomorphic conditions Press, Cambridge, xv þ 755 pp. Gradstein, F.M., Ogg, J.G., Smith, A.G. (Eds.), 2005. A Geologic Time Scale 2004. with regard to Mesochrysopidae, e.g., the presence of ‘M5’; the Cambridge University Press, Cambridge, 589 pp. branching of M very close to the wing base; the absence of Banksian Haase, E., 1890. Bemerkungen zur Palaeontologie der Insecten. Neues Jahrbuch fu¨ r folds. It is unclear yet if the presence of irregular crossvenations over Mineralogie. Geologie und Palaöntologie 2, 1–33. Handlirsch, A., 1906–1908. Die fossilen Insekten und die Phylogenie der rezenten the remigium and irregular, widely spaced branches of Rs found in Formen. Ein Handbuch fu¨ r Palaëontologen und Zoologen. W. Engelmann, Ascalochrysidae fam. nov. are primary or secondary conditions with Leipzig, ix þ 1430 pp. [Issued in 1906 (pp. 1–640), 1907 (pp. 641–1120), 1908 regard to the regular venation of Mesochrysopidae (s.l.). (pp. 1120–1430)]. Haring, E., Aspo¨ck, U., 2004. Phylogeny of the Neuropterida: a first molecular The mesochrysopids known from the Lower Cretaceous of China approach. Systematic Entomology 29, 415–430. (Yixian and Laiyang formations), i.e., Allopterus, Mesascalaphus Ren He, H.Y., Wang, X.L., Zhou, Z.H., Jin, F., Wang, F., Yang, L.K., Ding, X., Boven, A., in Ren et al., 1995, Tachinymphes Ponomarenko, 1992, are very Zhu, R.X., 2006. 40Ar/39Ar dating of Lujiatun Bed (Jehol Group) in Liaoning, northeastern China. L04303. Geophysical Research Letters 33. doi:10.1029/ different from Ascalochrysa gen. nov. All these genera belong to 2005GL025274. specialized representatives of the family Mesochrysopidae (s.l.). Of Henry, C.S., 1982. Neuroptera. In: Parker, S. (Ed.), Synopsis and Classification of the Jurassic genera of Mesochrysopidae, which are more ‘primitive’, Living Organisms, vol. 2. McGraw-Hill Book Co., New York, pp. 470–482. the only genus with known hind wings is Mesochrysopa Handlirsch, Hou, L.H., Martin, L.D., Zhou, Z.H., Feduccia, A., Zhang, F., 1999. A diapsid skull in a new species of the primitive bird Confuciusornis. Nature 399, 679–682. 1906. It has the venation that most resembles Ascalochrysa gen. Kukalova´-Peck, J., 1991. Fossil history and the evolution of hexapod structures. nov., for example it has a quite similar configuration of M. But all of In: Naumann, I.D. (Ed.), The Insects of Australia. A textbook for Students the Jurassic genera of Mesochrysopidae differ from Ascalochrysa and Research Workers, second ed., vol. 1. Melbourne University Press, Carlton, pp. 41–179. gen. nov. in having the regular pectinate branching of Rs, the Kukalova´-Peck, J., Willmann, R., 1990. Lower Permian ‘mecopteroid-like’ insects crossveins arranged in gradate series, and the absence of subcostal from Central Europe (Endopterygota). Canadian Journal of Earth Sciences 27, crossveins, except for one basal. The Early Jurassic genus Proto- 459–468. Lambkin, K.J., 1986. A revision of the Australian Mantispidae (Insecta: Neuroptera) aristonymphes Nel et Henrotay, 1994 in this respect does not differ with a contribution to the classification of the family. I. General and Drepani- from the Late Jurassic genera. cinae. Australian Journal of Zoology 116 (Supplementary Series), 1–142. Author's personal copy

1222 D. Ren, V.N. Makarkin / Cretaceous Research 30 (2009) 1217–1222

Li, J.G., Batten, D.J., 2007. Palynological evidence of an Early Cretaceous age for the Ren, D., 2003a. Two new Late Jurassic genera of kalligrammatids from Beipiao, Yixian Formation at Sihetun, western Liaoning, China. Cretaceous Research 28, Liaoning (Neuroptera: Kalligrammatidae). Acta Zootaxonomica Sinica 28, 105– 333–338. 109 [in Chinese, English summary]. Linnaeus, C., 1758. Systema natura per regna tria naturae secundum classes, Ren, D., 2003b. Study on the Jurassic fossil Raphidioptera and Neuroptera from ordines, genera, species, cum characteribus, differentiis, synonymis, locis, tenth northeast China. PhD dissertation, Beijing Forestry University, Beijing, 202 pp. ed., vol. 1. Salvii, Holmiae, 824 pp. [in Chinese, English abstract]. Luo, Z.X., 1999. A refugium for relicts. Nature 400, 23–25. Ren, D., Engel, M.S., 2008a. Aetheogrammatidae, a new family of lacewings from the Makarkin, V.N., 1997. Fossil Neuroptera of the Lower Cretaceous of Baisa, East Mesozoic of China (Neuroptera: Myrmeleontiformia). Journal of the Kansas Siberia. Part 3. Chrysopidae (Insecta). Spixiana 20, 107–118. Entomological Society 81, 161–167. Makarkin, V.N., Archibald, S.B., 2003. Family affinity of the genus Palaeopsychops Ren, D., Engel, M.S., 2008b. A second antlion from the Mesozoic of northeastern Andersen with description of a new species from Early Eocene of British China (Neuroptera: Myrmeleontidae). Alavesia 2, 183–186. Columbia (Neuroptera: Polystoechotidae). Annals of the Entomological Society Ren, D., Guo, Z.G., 1996. On the new fossil genera and species of Neuroptera of America 96, 171–180. (Insecta) from the Late Jurassic of northeast China. Acta Zootaxonomica Sinica Makarkin, V.N., Menon, F., 2005. New species of the Mesochrysopidae (Insecta, 21, 461–479. Neuroptera) from the Crato Formation of Brazil (Lower Cretaceous), with Ren, D., Lu, L.W., Guo, Z.G., Ji, S., 1995. Faunae and Stratigraphy of Jurassic-Creta- taxonomic treatment of the family. Cretaceous Research 26, 801–812. ceous in Beijing and the Adjacent Areas. Seismic Publishing House, Beijing, 223 Martins-Neto, R.G., 1994. Neuro´ pteros (Insecta, Planippenia) da Formaçaõ Santana pp. [in Chinese, English summary]. (Cretaćeo Inferior), Bacia do Araripe, nordeste do Brasil. IX. Primeiros resultados Ren, D., Yin, J.C., 2002. A new genus and new species of lacewings in the Jurassic of da composiçaõ da fauna e descriçaõ de novos ta´xons. Acta Geologica Leo- China (Neuroptera: Myrmeleontoidea). Acta Zootaxonomica Sinica 27, 269–273. poldensia 17 (39/1), 269–288. Schlu¨ ter, T., 1986. The fossil Planipennia – a review. In: Gepp, J., Aspo¨ck, H., Ho¨l- Martins-Neto, R.G., 2000. Remarks on the neuropterofauna (Insecta, Neuroptera) zel, H. (Eds.), Recent Research in Neuropterology. Proceedings of the Second from the Brazilian Cretaceous, with keys for the identification of the known International Symposium on Neuropterology. Privately printed, Graz, Austria, taxa. Acta Geologica Hispanica 35, 97–118. pp. 103–111. Martins-Neto, R.G., Heads, S.W., Bechly, G., 2008. Neuropterida: snakeflies, Schneider, W.G., 1851. Symbolae ad monographiam generis Chrysopae, Leach. Apud dobsonflies, and lacewings. In: Martill, D.M., Bechly, G., Loveridge, R.F. (Eds.), Ferdinandum Hirt, Vratislaviae, 178 pp. The Crato Fossil Beds of Brazil. Window into an Ancient World. Cambridge Sha, J.G., 2007a. Current research on Cretaceous lake systems in northeast China. University Press, Cambridge, pp. 328–340. Cretaceous Research 28, 143–145. Martynova, O.M., 1949. Mesozoic lacewings (Neuroptera) and their bearing on Sha, J.G., 2007b. Cretaceous stratigraphy of northeast China: non-marine and concepts of phylogeny and systematics of the order. Trudy Paleon- marine correlation. Cretaceous Research 28, 146–170. tologicheskogo Instituta 20, 150–170 [in Russian]. Smith, P.E., Evensen, N.M., York, D., Chang, M.M., Jin, F., Li, J.L., Cumbaa, S., Matsukawa, M., Ito, M., Nishida, N., Koarai, K., Lockley, M.G., Nichols, D.J., 2006. The Russell, D., 1995. Dates and rates in ancient lakes: 40Ar-39Ar evidence for an Cretaceous Tetori biota in Japan and its evolutionary significance for terrestrial Early Cretaceous age for the Jehol Group, northeast China. Canadian Journal of ecosystems in Asia. Cretaceous Research 27, 199–225. Earth Science 32, 1426–1431. Menon, F., Makarkin, V.N., 2008. New fossil lacewings and antlions (Insecta, Neu- Sun, G., Dilcher, D.L., Zheng, S.L., Zhou, Z.K., 1998. In search of the first flower: roptera) from the Lower Cretaceous Crato Formation of Brazil. Palaeontology 51, a Jurassic angiosperm, Archaefructus, from Northeast China. Science 282, 149–152. 1692–1695. Nakahara, W., 1958. The Neurorthinae, a new subfamily of the Sisyridae (Neuro- Swisher III, C.C., Wang, Y.Q., Wang, X.L., Xu, X., Wang, Y., 1999. Cretaceous age for the ptera). Mushi 32, 19–32. feathered dinosaurs of Liaoning, China. Nature 400, 58–61. Nel, A., Delclo` s, X., Hutin, A., 2005. Mesozoic chrysopid-like Planippenia: a phylo- Swisher III, C.C., Wang, X.L., Zhou, Z.Z., Wang, Y.Q., Jin, F., Zhang, J., Xu, X., Zhang, F., genetic approach (Insecta: Neuroptera). Annales de la Socie´te´ Entomologique Wang, Y., 2002. Further support for a Cretaceous age for the feathered-dinosaur de France 41, 29–68. beds of Liaoning, China: new 40Ar/39Ar dating of the Yixian and Tuchengzi Nel, A., Henrotay, M., 1994. Les Chrysopidae me´sozoı¨ques. E´ tat actuel des con- Formations. Chinese Science Bulletin 47, 135–138. naissances. Description d’un nouveau genre et nouvelle espe`ce dans le Juras- Tillyard, R.J., 1919. The panorpoid complex. Part 3: – The wing-venation. Proceed- sique infe´rieur (Lias) (Insecta: Neuroptera). Annales de la Socie´te´ ings of the Linnean Society of New South Wales 44, 533–718. Entomologique de France 30, 293–318. Vilesov, A.P., Novokshonov, V.G., 1994. New fossil insects (Myrmeleontida, Jurinida) Nel, A., Roques, P., Nel, P., Prokop, J., Steyer, J.S., 2007. The earliest holometabolous from the Upper Permian of eastern Kazakhstan. Paleontologicheskii Zhurnal 1994 insect from the Carboniferous: a "crucial" innovation with delayed success (2), 66–74 [in Russian; English translation: Paleontological Journal 28(2), 81–92]. (Insecta Protomeropina Protomeropidae). Annales de la Socie´te´ Entomologique Wang, S.S., Wang, Y.Q., Hu, H.G., Li, H.M., 2001. The existing time of Sihetun de France 43, 349–355. vertebrate in western Liaoning, China: evidence from U-Pb dating of zircon. New, T.R., 1989. Planipennia, Lacewings. In: Fischer, M. (Ed.), Handbuch der Zoo- Chinese Science Bulletin 46, 779–782. logie. Arthropoda: Insecta. Part 30, Band IV. Walter de Gruyter, Berlin, 132 pp. Wedmann, S., Makarkin, V.N., 2007. A new genus of Mantispidae (Insecta: Neuroptera) Novokshonov, V.G., 1997. Early Evolution of Scorpionflies (Insecta: Panorpida). from the Eocene of Germany, with a review of the fossil record and palae- Nauka Press, Moscow, 140 pp. [in Russian]. obiogeography of the family. Zoological Journal of the Linnean Society 149, 701–716. Oswald, J.D., 1993. Revision and cladistic analysis of the world genera of the family Winterton, S.L., 2003. Molecular phylogeny of Neuropterida with emphasis on the Hemerobiidae (Insecta: Neuroptera). Journal of the New York Entomological lacewings (Neuroptera). Entomologische Abhandlungen, Staatliches Museum Society 101, 143–299. fu¨ r Tierkunde in Dresden 61, 158–160. Ponomarenko, A.G., 1992. Neuroptera (Insecta) from the Lower Cretaceous of Wootton, R.J., 2003. Wings. In: Resh, V.H., Carde, R.T. (Eds.), Encyclopedia of Insects. Transbaikalia. Paleontologicheskii Zhurnal 1992 (3), 43–50 [in Russian; English Academic Press, London, pp. 1186–1192. translation: Paleontological Journal 26(3), 56–66]. Yang, W., Li, S.G., Jiang, B.Y., 2007. New evidence for Cretaceous age of the feathered Ponomarenko, A.G., Rasnitsyn, A.P., 1974. New Mesozoic and Cenozoic Proto- dinosaurs of Liaoning: zircon U-Pb SHRIMP dating of the Yixian formation in mecoptera. Paleontologicheskii Zhurnal 1974 (4), 59–73 [in Russian, English Sihetun, northeast China. Cretaceous Research 28, 177–182. translation: Paleontological Journal 8, 493–507]. Zhang, J.F., 1991. A new family of Neuroptera (Insecta) from the Late Mesozoic of Ponomarenko, A.G., Shcherbakov, D.E., 2004. New Lacewings (Neuroptera) from the Shandong, China. Science in China (Series B) 34, 1105–1111. Terminal Permian and Basal Triassic of Siberia. Paleontological Journal 38 Zhou, Z.H., Barrett, P.M., Hilton, J., 2003. An exceptionally preserved Lower Creta- (Suppl. 2), S197–S203. ceous ecosystem. Nature 421, 807–814. Rasnitsyn, A.P., 2007. On the discussion of the wing venation of (Archae) Orthoptera Zhu, R.X., Pan, Y.X., Shi, R.P., Liu, Q.S., Li, D.M., 2007. Palaeomagnetic and 40Ar/39Ar (Insecta). Paleontological Journal 41, 341–344. dating constraints on the age of the Jehol Biota and the duration of deposition Ren, D., 2002. A new lacewing family (Neuroptera) from the Middle Jurassic of Inner of the Sihetun fossil-bearing lake sediments, northeast China. Cretaceous Mongolia, China. Entomologia Sinica 9, 53–67. Research 28, 171–176.