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From the Middle Jurassic of Daohugou, Inner Mongolia, China

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From the Middle Jurassic of Daohugou, Inner Mongolia, China Two new kalligrammatids (Insecta, Neuroptera) from the Middle Jurassic of Daohugou, Inner Mongolia, China QING LIU*, DARAN ZHENG, QI ZHANG, BO WANG, YAN FANG and HAICHUN ZHANG LIU, Q., ZHENG, D.R., ZHANG, Q., WANG, B., FANG,Y.&ZHANG, H.C., iFirst article. Two new kalligrammatids (Insecta, Neuroptera) from the Middle Jurassic of Daohugou, Inner Mongolia, China. Alcheringa 38, XX–XX. ISSN 0311-5518. A new genus and two new species of kalligrammatid lacewings (Insecta, Neuroptera), Kalligramma paradoxum sp. nov. and Huiyingogramma formosum gen. et sp. nov., are described and figured, based on two well-preserved forewings from the Middle Jurassic of Daohugou, Inner Mongolia, China. Kalligramma paradoxum sp. nov. can be distinguished from other known Kalligramma species based on forewing characters (e.g., wing shape, costal space, branches of Rs, eye-spot). Huiyingogramma gen. nov. is characterized by a distinct humeral recurrent vein, relatively broad costal space with well-forked costal veinlets, well-developed eye-spot and dense crossveins over the entire wing. Qing Liu (corresponding author) [[email protected]; [email protected]], Daran Zheng [[email protected]], Qi Zhang [[email protected]], Bo Wang [[email protected]], Yan Fang [[email protected]] and Haichun Zhang [[email protected]], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, 210008, PR China; secondary address of Daran Zheng & Qi Zhang, University of Chinese Academy of Sciences, Beijing, 100049, PR China. Received 15.4.2013, revised 9.6.2013, accepted 24.6.2103. Key words: Neuroptera, Kalligrammatidae, Kalligramma, Huiyingogramma, eye-spot, Middle Jurassic, China. KALLIGRAMMATIDAE, an extinct family of tinct eye-spot in many species. They are commonly Neuroptera (Insecta) known from the Jurassic to the referred to as the ‘butterflies of the Jurassic’ (Engel Early Cretaceous, have been widely reported from 2005). However, kalligrammatids may have had a pred- Europe and Asia (Walther 1904, Handlirsch 1906, atory feeding habit like most extant neuropterans (Fang 1919, Martynova 1947, Panfilov 1968, 1980, et al. 2010). Here, a new genus and two new species of Ponomarenko 1984, 1992, Whalley 1988, Carpenter Kalligrammatidae are described from the Middle Juras- 1992, Lambkin 1994, Ren & Guo 1996, Jarzembowski sic of Daohugou, Inner Mongolia, China. 2001, Ren & Oswald 2002, Zhang 2003, Zhang & Zhang 2003, Engel 2005, Yang et al. 2009, 2011). Until the present study, 14 genera have been attributed to the Materials and methods Kalligrammatidae: Kalligramma Walther, 1904; The two neuropteran forewings described in this paper Meioneurites Handlirsch, 1906; Palparites Handlirsch, were collected from the Middle Jurassic Haifanggou 1906; Kalligrammula Handlirsch, 1919; Lithogramma Formation at Daohugou Village, Wuhua Township, Downloaded by [Capital Normal University] at 17:50 22 September 2013 Panfilov, 1968; Kalligrammina Panfilov, 1980; Ningcheng County, Chifeng City, Inner Mongolia, Angarogramma Ponomarenko, 1984; Sophogramma China. The Daohugou deposits, consisting of grey tuff, Ren & Guo, 1996; Kallihemerobius Ren & Oswald, tuffaceous siltstone and mudstone, are now considered 2002; Limnogramma Ren, 2003; Oregramma Ren, to be one of the most important insect Lagerstätten 2003; Sinokalligramma Zhang, 2003; Protokalligramma (Rasnitsyn & Zhang 2004). The fossil insects at this Yang, et al. 2011 and Apochrysogramma Yang, et al. locality are commonly preserved as compressions in 2011. Most of these genera were documented from grey tuffaceous siltstones and are found together with Asia, and eight have been reported from China. On this small freshwater branchiopods (Wang et al. 2009). basis, several authors have proposed that Kalligrammat- All photographs were taken using a Canon EOS 5D idae might have originated in Asia (e.g., Zhang 2003, digital camera. Line drawings were prepared on photo- Yang et al. 2009). The enigmatic Kalligrammatidae are graphs using CorelDRAW 12 image-editing software. characterized by large wings, presumably bright colora- The traditional venation terminology of Comstock tion, dense crossveins over the entire wing and a dis- (1918; sensu Wootton 2003) with the recent interpreta- tion of Oswald (1993) and Archibald & Makarkin (2006) is used in this study. Wing vein abbreviations are as follows: C, costa; Sc, subcosta; Vr, humeral Ó 2013 Association of Australasian Palaeontologists recurrent vein; R, radius; R1, first branch of R; Rs, http://dx.doi.org/10.1080/03115518.2013.828251 2 QING LIU et al. ALCHERINGA radial sector; Rs1, most basal branch of Rs; M, media; & Zhang, 2003, from the Middle Jurassic Haifanggou MA, media anterior; MP, media posterior; Cu, cubitus; Formation of Liaoning, China. CuA, cubitus anterior; CuP, cubitus posterior; 1A–3A, first to third anal veins. All specimens are housed at the Kalligramma paradoxum sp. nov. (Fig. 1A–C) Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (prefix NIGP). Holotype. A nearly complete and well-preserved left forewing (NIGP 156188), part and counterpart. Type locality, formation and age. Daohugou Village, Systematic palaeontology Wuhua Township, Ningcheng County, Chifeng City, Inner Mongolia, China; Haifanggou Formation, Middle Class INSECTA Linnaeus, 1758 Jurassic. Order NEUROPTERA Linnaeus, 1758 Family KALLIGRAMMATIDAE Handlirsch, 1906 Etymology. From the Latin ‘paradoxus’ for marvellous or strange. Kalligramma Walther, 1904 Diagnosis. Forewing elongate oval. Eye-spot well Type species. Kalligramma haeckeli Walther, 1904, from developed. Crossveins dense on entire wing. Costal the Upper Jurassic Solnhofen Plattenkalk of Germany. space moderately broad and slightly narrowed towards Other species. Kalligramma multinerve Panfilov, 1968, apex; costal veinlets sinuous and mostly forked; Rs K. flexuosum Panfilov, 1968, K. sharovi Panfilov, 1968 with 16 primary branches, each forked distally; MA and K. turutanovae Panfilov, 1968, from the Upper Juras- simple and forked distally; MP with at least seven pec- sic Karabastau Formation of Kazakhstan; K. liaoningense tinate branches; CuP branched distally; 1A forked Ren & Guo, 1996, from the Lower Cretaceous Yixian beyond its mid-length; 2A well developed; 1A and 2A Formation of Liaoning, China; K. jurarchegonium Zhang stems both parallel to CuP; 3A short with few branches. Downloaded by [Capital Normal University] at 17:50 22 September 2013 Fig. 1. Kalligramma paradoxum sp. nov. Holotype, a nearly complete and well-preserved left forewing (NIGP 156188). A, Part. B, Counterpart. C, Line drawing of part. Scale bar = 20 mm. ALCHERINGA JURASSIC KALLIGRAMMATID INSECTS FROM CHINA 3 Description. Left forewing elongate oval and broad, ing apices). MA running just posterior to and forked 84 mm long, 39 mm wide. Anterior margin partially dichotomously just apical to eye-spot. MP with seven damaged, most of posterior margin and apex missing. branches preserved that are dichotomously forked (some Eye-spot fuscous and rounded, located slightly beyond several times over); MP1 forked slightly beyond first wing mid-length and slightly closer to costal margin forking of MA; MP2 forked basad of eye-spot and than to posterior margin, with one unpigmented dot more basad of any remaining branches. Preserved CuA subcentrally located and four chains of small unpig- section simple, running close to MP stem and forked mented dots attached apically. Wing decorated with two apically. CuP damaged apically with four primary roughly circular markings surrounding eye-spot and branches preserved. 1A forked basad of first branches some irregular band-like markings near apex along of CuA and CuP. 2A densely and pectinately branched posterior margin. Costal space most dilated near basal along length. 3A very short, with no more than five one-third of its length and slightly narrowed towards branches. Maximum width between CuA and CuP both wing base and apex. Costal veinlets sinuous with about twice as broad as that between MP and CuA, several basal branches simple and all others forked; about 1.5 times as broad as between CuP and 1A, and costal space with dense crossveins. Subcostal space slightly broader than between 1A and 2A (Fig. 1A–C). distinctly wider than R1 space, and both regions have widely spaced crossveins. Sc and R1 fused within Huiyingogramma gen. nov. apical one-eighth of wing length. Rs damaged distally, Type species. Huiyingogramma formosum gen. et sp. nov. but with 16 primary branches preserved; each branch running rectilinearly and forked distally several times Etymology. ‘Huiyingo-’ is dedicated to a great lady, (13th, 14th and 16th branches unknown owing to miss- Mrs Huiying Liu, who is the mother of the lead Downloaded by [Capital Normal University] at 17:50 22 September 2013 Fig. 2. Huiyingogramma formosum gen. et sp. nov. Holotype, a nearly complete and well-preserved right forewing (NIGP 156189). A, Part. B, Counterpart. C, Line drawing of part. Scale bar = 20 mm. 4 QING LIU et al. ALCHERINGA author, ‘-gramma’ is a traditional ending of generic Discussion names in Kalligrammatidae. Among known kalligrammatids, well-preserved and Diagnosis. Forewing broad. Eye-spot well developed. complete specimens consisting of both forewings and Crossveins dense over entire wing. Humeral recurrent hindwings are extremely rare (e.g., Yang et al. 2009, fi vein (Vr) present. Costal space broad; crossveins pres- g. 1). Most kalligrammatid genera and species
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