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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 0195-6671/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2009.06.004 Author's personal copy 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 Æ 0.18 basal crossvein at most]; crossvenation over remigium irregular for the lowest beds (Chen et al., 2004), 126.1 Æ1.7 to 124.6 Æ 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 Æ0.2 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,