New fossil hangingflies (Mecoptera, Raptipeda, Bittacidae) from the Middle Jurassic to Early Cretaceous of Northeastern China Xiaoguang YANG Dong REN Chungkun SHIH College of Life Sciences, Capital Normal University, 100048 Beijing (China) [email protected] [email protected] [email protected] Yang X.-G., Ren D. & Shih C.-K. 2012. — New fossil hangingflies (Mecoptera, Raptipeda, Bit- tacidae) from the Middle Jurassic to Early Cretaceous of Northeastern China. Geodiversitas 34 (4): 785-799. http://dx.doi.org/10.5252/g2012n4a4 ABSTRACT A new genus with a new species, Exilibittacus lii n. gen., n. sp., and a new spe- cies, Megabittacus spatiosus n. sp., are described and illustrated. Both of them belong to the family Bittacidae (Mecoptera). These specimens were collected respectively from the Middle Jurassic of Jiulongshan Formation at Daohugou in Inner Mongolia and from the Late Jurassic to Early Cretaceous of Yixian Formation at Dawangzhangzi in Liaoning, China. In Exilibittacus n. gen., both RP+MA and MP have four branches in forewings, while both have three KEY WORDS branches in hind wings, and RP1+2 and MP3 are not forked. This is the first Mecoptera, Bittacidae, time that the character ‘MP with three branches in hind wings’ is reported for Insect fossil, the Bittacidae. Based on data on Megabittacus spatiosus n. sp., the diagnosis of Jiulongshan Formation, Megabittacus is revised and a key to the known species is provided. Morphologi- Yixian Formation, new species, cal changes in hangingflies from Middle Jurassic to present are discussed based new genus. on body length, wing length, and forewing/body length ratio. GEODIVERSITAS • 2012 • 34 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 785 Yang X.-G. et al. RÉSUMÉ De nouveaux fossiles de « mouches-scorpion » (Mecoptera, Raptipeda, Bittacidae) du Jurassique Moyen au Crétacé inférieur du nord-est de la Chine. Exilibittacus lii n. gen., n. sp. et Megabittacus spatiosus n. sp. sont décrits et appar- tiennent à la famille des Bittacidae (Mecoptera). Les spécimens proviennent, respec- tivement, du Jurassique moyen de la Formation Jiulongshan (Daohugou, Mongolie intérieure) et du Jurassique supérieur au Crétacé inférieur de la Formation Yixian (Dawangzhangzi, Liaoning, Chine). Chez Exilibittacus n. gen., RP+MA et MP ont chacune quatre branches aux ailes antérieures, tandis que ces nervures ont trois branches aux ailes postérieures, et RP1+2 et MP3 ne sont pas branchées. C’est la première fois que le caractère « MP avec trois branches aux ailes postérieures » est MOTS CLÉS observé chez les Bittacidae. Basée sur des données sur Megabittacus spatiosus n. sp., Mecoptera, la diagnose de Megabittacus est révisée et une clé des espèces connues est fournie. Bittacidae, Les changements morphologiques chez les « mouches-scorpion » du Jurassique insectes fossiles, Formation Jiulongshan, Moyen à l’actuel sont discutés sur la base de la longueur du corps, la longueur des Formation Yixian. ailes, et du rapport de longueur ailes antérieures/corps. INTRODUCTION Novokshonov 1997; Petrulevičius & Jarzembowski 2004; Petrulevičius et al. 2007). Fossil records show Bittacidae, a large family of Mecoptera which is one a broadest generic diversity occurring in the Jurassic of the less diverse insect Orders, is considered to be (Novokshonov 2002). a sister group of Cimbrophlebiidae. Together, Bit- To date, eight fossil genera of hangingflies from tacidae and Cimbrophlebiidae form the infraorder the Middle Jurassic to the Early Cretaceous have of Raptipeda (Willmann 1977, 1989). Bittacids been recorded in China: Liaobittacus Ren, 1993 and cimbrophlebiids share the same special tarsi from the Haifanggou Formation; Jichoristella Ren, structure: the fifth tarsomere can be folded against 1994 from the Lushanfeng Formation; Megabit- the fourth and tarsus has only one claw. Their wing tacus Ren, 1997, and Sibirobittacus Sukatsheva, venations are similar, but cimbrophlebiids can be 1990 from the Yixian Formation; Mongolbittacus easily identified by long and branched 2A, which is Petrulevičius, Huang & Ren, 2007, Formosibittacus short and unbranched in bittacids (Willmann 1977, Li, Ren & Shih, 2008, Jurahylobittacus Li, Ren & 1989). Usually bittacids wings are slender and both Shih, 2008, and Decoribittacus Li & Ren, 2009 RP+MA and MP veins have four branches, except from the Jiulongshan Formation. for Orobittacus RP+MA with three branches. They A new hangingfly, Exilibittacus lii n. gen., n. sp., are called hangingflies because they commonly hang is described in this paper. The specimen was col- their bodies by two fore legs from branches among lected from the Jiulongshan Formation at Daohugou low vegetation, while using free middle and hind Village of Ningcheng County in Inner Mongolia, legs to catch insects (Petrulevičius et al. 2007). At China. The age of the Daohugou fossil-bearing beds present, there are 16 extant genera including about is considered to be the Middle Jurassic (Ren et al. 270 species of Bittacidae (Krzemiński 2007). Extant 1995; Ren & Krzemiński 2002; Ren et al. 2002; bittacids are distributed globally in both temperate Shen et al. 2003; Chen et al. 2004; Liu et al. 2004; and warm tropical climates. In addition to extant Gao & Ren 2006; Huang et al. 2006; Ji et al. 2006). bittacids, about 29 fossil genera have been described A new species, Megabittacus spatiosus n. sp., was (Tillyard 1933; Handlirsch 1939; Martynov 1951; collected from the Yixian Formation at Dawang- Byers 1979; Villegas & Byers 1981; Ren 1993, 1997; zhangzi in Lingyuan, western Liaoning, China. The 786 GEODIVERSITAS • 2012 • 34 (4) Fossil hangingflies from China age of the Yixian Formation remains debatable. TABLE 1. — Comparison of venation nomenclatures between Three different opinions about the age: the Late Kukalová-Peck and Byers. Jurassic (Ren 1997; Zheng et al. 2003), transition from the Late Jurassic to Early Cretaceous (Chen Kukalová-Peck system et al. 2004; Wang et al. 2004, 2005), and the Early (1983, 1991) Byers system (1979) Cretaceous (Swisher et al. 1999; Li et al. 2001; Pang ScP Sc RA R1 et al. 2002; Zhou et al. 2003), have been proposed RP RS based on both biostratigraphic and radiometric MA R5 geochronologies. By comparing the Yixian biota MP M MP4 + CuAl + 2 M4 with the Solnhofen biota of Germany, the Purbeck CuA3 + 4 CuA biota in England and the Late Jurassic Terori-type CuP CuP and Ryoseki-type floras in Japan, Wang et al. (2005) AA3 + 4 1A AP1 + 2 2A considered the synthetic age of the Yixian Forma- AP3+4 3A tion as the Late Jurassic-Early Cretaceous (the Late Tithonian to the Berriasian). SYSTEMATICS MATERIAL AND METHODS Order MECOPTERA Packard, 1886 Infraorder RAPTIPEDA Willmann, 1977 The fossil specimens were examinated by a LEICA Family BITTACIDAE Handlirsch, 1906 MZ12.5 dissecting microscope and illustrated with the aid of a camera lucida attached to the Genus Exilibittacus n. gen. microscope. All specimens are deposited in the Key Laboratory of Insect Evolution & Environmental TYPE SPECIES. — Exilibittacus lii n. sp. Changes, the College of Life Sciences, Capital ETYMOLOGY. — The generic name is derived from the Normal University, Beijing, China (CNU, Ren Latin word “exil-” (weak) and bittacus (a recent genus Dong, Curator). Photo graphs of whole speci- of Bittacidae). Gender masculine. mens were taken by Nikon D100 digital camera DIAGNOSIS. — One crossvein between ScP and RA; RA coupled to a Nikkor 105 mm macro lens, and sharply bent distally; RP+MA forking before the fork of detailed photographs were made by using Nikon MP; “Kreuz der Bittaciden” aligned, MP3+MP4+CuA1+2 SMZ1000 stereomicroscope. The terminology forking before posterior part of “Kreuz der Bittaciden” for wing venation follows Kukalová-Peck (1983, (if existing); one crossvein between MP4+CuA1+2 and CuA3+4; AA3+4 short; Forewing both RP+MA and 1991), “Kreuz der Bittaciden” is defined as the MP with four branches, but hind wing, three branches. cross-veins of [RP3+4+MA-MP1+2, MP1+2-MP3] (Novokshonov 1993). The venational nomenclature REMARKS of Kukalová-Peck is compared with that of Byers Exilibittacus n. gen. can be assigned to the infraorder (1979) in Table 1 for easy reference. Raptipeda (Willmann 1977, 1989) on the char- acteristics of legs: long and slender, with a single, predatory claw, the fifth tarsomere folding back ABBREVIATIONS Venation nomenclature toward fourth. Furthermore, it can be classified RA anterior radius; into the Bittacidae by AA3+4 short and AP1+2 RP posterior radius; absent on a well-preserved forewing, in contrast MA anterior media; to cimbrophlebiids with AA3+4 long and AP1+2 MP posterior media; very long and branched. CuA anterior cubitus; AA anterior anal; Forewing of Exilibittacus n. gen. is similar to AP posterior anal; Antiquanabittacus Petrulevičius & Jarzembowski, ScP posterior subcosta. 2004 and Mongolbittacus Petrulevičius, Huang & GEODIVERSITAS • 2012 • 34 (4) 787 Yang X.-G. et al. TABLE 2. — A list of fossil and extant bittacid species examined. Abbreviation: fm., formation; LR, Length ratio of forewing/body. Species Distribution Age Sex Body (mm) Forewing (mm) LR Liaobittacus longanteenatus Ren, 1993 Haifanggou Fm. J2 Male 20 21 1.05 Mongolbittacus daohugouensis Jiulongshan J2 ? 9.5 12.5 1.32 Petrulevičius, Huang & Ren, 2007 Fm. Formosibittacus macularis Jiulongshan J2 Female 21 23 1.10 Li, Ren & Shih, 2008 Fm. Jurahylobittacus astictus Jiulongshan J2 Male 18 12 0.67 Li, Ren & Shih, 2008 Fm. Exilibittacus lii