SYSTEMATICS Two New of Kalligrammatidae () From the of China, With Comments on Venational Homologies

1,2 1,3 1 VLADIMIR N. MAKARKIN, DONG REN, AND QIANG YANG

Ann. Entomol. Soc. Am. 102(6): 964Ð969 (2009) ABSTRACT Two new fossil species of Kalligrammatidae (Neuroptera) are described from the Middle Jurassic locality at Daohugou, Inner Mongolia, China: Limnogramma hani n. sp. and Limno- gramma mongolicum n. sp. Some venational homologies in Neuroptera are discussed brießy.

KEY WORDS Neuroptera, Kalligrammatidae, Jurassic, Daohugou, China

The Mesozoic Kalligrammatidae are the most striking venational homologies of the hind wings of Neurop- neuropterans, due to their large, presumably bright- tera. colored wings with eye-spots similar to those of but- terßies (e.g., saturniids). Their resemblance to Lepi- Materials and Methods doptera may be greater if the hypothesis is conÞrmed that kalligrammatids were specialized pollinators This study is based on two specimens collected near feeding on pollen of generative organs of gymno- Daohugou Village (Shantou Township, Ningcheng sperms (Zherikhin 1978, 1980; Rasnitsyn 1980). County, Inner Mongolia, China) and housed in the The family has never been revised, and is in need of fossil collection of the Key Laboratory of Insect comprehensive taxonomic and phylogenetic treat- Evolution & Environmental Changes, College of Life ment. The diagnoses of Handlirsch (1906Ð1908), Mar- Science, Capital Normal University, Beijing, China tynova (1962), and Carpenter (1992) are obsolete and (CNUB; D.R., curator). These insect-bearing beds are incomplete. Hitherto, 26 species were described from here considered as belonging to the Jiulongshan For- the Middle Jurassic to Early of present-day mation and dated Middle Jurassic (alternatively con- Eurasia (Walther 1904; Handlirsch 1919; Martynova sidered to be Late Jurassic by Zhang 2003). 1947; PanÞlov 1968, 1980; Ponomarenko 1992; Lam- Specimens were examined using an MZ12.5 dissect- bkin 1994; Ren and Guo 1996; Jarzembowski 2001; Ren ing microscope (Leica, Wetzlar, Germany), illus- and Oswald 2002; Ren 2003; Zhang 2003; Zhang and trated with the aid of a drawing tube and Illustrator Zhang 2003; Engel 2005). In addition, there are un- CS3 and Photoshop CS3 (Abobe Systems, Mountain described kalligrammitids from the Early Cretaceous View, CA). of Kazakhstan (Zherikhin 1978), Spain (Martõ´nez- We use the traditional (sensu Wootton 2003) vena- Delclo`s and Nel 1995), and England (Ross and Jar- tional terminology of Comstock (1918) following the zembowski 1996). The oldest known species of Kal- recent interpretations of Oswald (1993), Archibald ligrammatidae (undescribed) is from Early Jurassic and Makarkin (2006), and Wedmann and Makarkin (Early [Late Liassic]) of Kerkhofen, Ger- (2007). many (called ÔPsychopsidaeÕ in Schmidt-Kaler et al. 1992: Þg. 50). It is assumed that the Makarkinia Systematic Palaeontology Martins-Neto, 1992 from the Late Crato For- Order Neuroptera Linnaeus, 1758 mation of Brazil is kalligrammatid (Makarkin and Family Kalligrammatidae Handlirsch, 1906 Archibald 2003), and an undescribed specimen of typ- Genus Limnogramma Ren, 2003 ical kalligrammatid appearance is also known from this formation. Therefore, the known temporal occur- Type Species. Limnogramma mirum Ren, 2003, by rence of this family ranges from Early Toarcian to Late original designation. Aptian. Diagnosis. May be easily separated from other kal- In this article, we describe two small kalligrammma- ligrammatid genera by CuP very deeply forked in hind tid species from the Middle Jurassic locality at Dao- wings [CuP shallowly forked in other genera]. hugou, in Inner Mongolia, China, and we discuss some Species Included. Three species, L. mirum, L. hani n. sp., and L. mongolicum n. sp., from Middle Jurassic 1 College of Life Science, Capital Normal University, Beijing to Early Cretaceous of China. 100048, China. Comments. The gender of Limnogramma is neuter, 2 Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, 690022, Russia. from the gender of the Greek noun gramma, drawing, 3 Corresponding author, e-mail: [email protected]. letter (ICZN 1999: article 30.1.1), and therefore the

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