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(INSECTA: NEUROPTERA: MANTISPIDAE) from the MESOZOIC of NORTH-EASTERN CHINA by JAMES E

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(INSECTA: NEUROPTERA: MANTISPIDAE) from the MESOZOIC of NORTH-EASTERN CHINA by JAMES E [Palaeontology, Vol. 56, Part 3, 2013, pp. 603–613] NEW FOSSIL MANTIDFLIES (INSECTA: NEUROPTERA: MANTISPIDAE) FROM THE MESOZOIC OF NORTH-EASTERN CHINA by JAMES E. JEPSON1,2*, SAM W. HEADS3, VLADIMIR N. MAKARKIN4 and DONG REN1 1College of Life Sciences, Capital Normal University, 105 Xianshianbeilu, Haidian District, Beijing 100048, China; e-mail: [email protected] 2School of Earth, Atmospheric and Environmental Sciences, University of Manchester, Oxford Road, Manchester M13 9PL, United Kingdom; e-mail: [email protected] 3Illinois Natural History Survey, Prairie Research Institute, University of Illinois at Urbana-Champaign, 1816 South Oak Street, Champaign, IL 61820-6960, USA; e-mail: [email protected] 4Institute of Biology and Soil Sciences, Far Eastern Branch, Russian Academy of Sciences, Vladivostok 690022, Russia; e-mail: [email protected] *Corresponding author. Typescript received 14 December 2011; accepted in revised form 21 October 2012 Abstract: Three new genera and four new species of the preserved, including the specialized raptorial forelegs articu- extinct mantidfly subfamily Mesomantispinae (Insecta: Neu- lated to the prothorax anteriorly, an autapomorphy of the roptera: Mantispidae) are described from the Lower Creta- family. These new taxa further confirm the placement of the ceous Yixian Formation of Liaoning and the Middle Jurassic subfamily Mesomantispinae within the family Mantispidae; Jiulongshan Formation of Inner Mongolia: Archaeodrepanicus however, the monophyly of Mesomantispinae has not been nuddsi gen. et sp. nov., A. acutus gen. et sp. nov., Sinomeso- confirmed, and it is likely that it will prove to be para- mantispa microdentata gen. et sp. nov., (Yixian Formation) phyletic. and Clavifemora rotundata gen. et sp. nov. (Jiulongshan For- mation). The specimens described herein represent the first Key words: Neuropterida, Hemerobiiformia, Jiulongshan Mesozoic mantidfly compression fossils to have body parts Formation, mantisflies, Mesomantispinae, Yixian Formation. Mantispidae, or ‘mantidflies’ (also known as mantis- spider egg cases (Lambkin 1986; Redborg 1998). Little is flies), are a highly specialized group of hemerobiiform known of the life histories of the subfamilies Symphrasinae Neuroptera characterized by their raptorial forelegs, large and Calomantispinae, and the biology of Drepanicinae eyes and elongation of the pronotum behind the forelegs remains almost entirely unknown (Grimaldi and Engel (Lambkin 1986; New 1989; Grimaldi and Engel 2005). 2005). Their specialization extends beyond morphology to their Mantidflies are rare in the fossil record (Wedmann and behaviour and life history, with some species being para- Makarkin 2007), but, nevertheless, have a geological range sites of Araneae or aculeate Hymenoptera; the larvae devel- dating back to the Lower Jurassic (Table 1). All four oping within the egg cases of spiders or inside the nests of extant subfamilies are known from fossil representatives, wasps (Redborg 1998; Wedmann and Makarkin 2007). along with the extinct Mesomantispinae (Wedmann and Indeed, some tropical species are remarkable mimics of Makarkin 2007). The oldest mantidflies are Liassochrysa their hymenopteran hosts, a feature that affords them some stigmatica Ansorge and Schlu¨ter, 1990 from the Lower protection from predators (Grimaldi and Engel 2005). The Jurassic of Dobbertin, Germany, and Promantispa similis family is most diverse and abundant in tropical regions Panfilov, 1980 from the Upper Jurassic of Karatau. In the and comprises approximately 400 extant species divided Cretaceous, the family is represented by Mesomantispa into four extant subfamilies: Symphrasinae, Drepanicinae, sibirica Makarkin, 1997 from Baissa, East Siberia, Gers- Calomantispinae and Mantispinae (Lambkin 1986; Ohl taeckerella asiatica Makarkin, 1990 from Kzyl-Zhar, 2004; Grimaldi and Engel 2005; Wedmann and Makarkin Kazakhstan, and Doratomantispa burmanica Poinar and 2007); and the extinct subfamily Mesomantispinae (Makar- Buckley, 2011 from Burmese amber. Cenozoic Mantispi- kin 1997; Wedmann and Makarkin 2007). Mantispinae dae are known from the Eocene oil shales of Grube Mes- constitutes the most diverse subfamily in terms of numbers sel, Germany, with Symphrasites eocenicus Wedmann and of species and are highly specialized, obligate predators of Makarkin, 2007, an unnamed larva from the Eocene ª The Palaeontological Association doi: 10.1111/pala.12005 603 604 PALAEONTOLOGY, VOLUME 56 TABLE 1. The fossil record of Mantispidae. Taxon Deposit Age Reference Feroseta prisca Dominican amber Miocene Poinar 2006 Dicromantispa moronei Dominican amber Miocene Engel and Grimaldi 2007 Dicromantispa electromexicana Mexican amber Miocene Engel and Grimaldi 2007 Climaciella(?) henrotayi Dauphin, France Oligocene Nel 1989 Prosagittalata oligocenica Ce´reste, France Oligocene Nel 1989 Vectispa relicta Bembridge Marls, Isle of White Eocene Cockerell 1921; Jarzembowski 1980 Unnamed larva Baltic amber Eocene Ohl 2011 Symphrasites eocenicus Messel, Germany Eocene Wedmann and Makarkin 2007 Doratomantispa burmanica Burmese amber Upper Cretaceous Poinar and Buckley 2011 Gerstaeckerella asiatica Kzyl-Zhar, Kazakhstan Upper Cretaceous Makarkin 1990 Sinomesomantispa microdentata Yixian Formation, China Lower Cretaceous Herein gen. et sp. nov. Archaeodrepanicus sp. Yixian Formation, China Lower Cretaceous Herein Archaeodrepanicus acutus gen. Yixian Formation, China Lower Cretaceous Herein et sp. nov Archaeodrepanicus nuddsi gen. Yixian Formation, China Lower Cretaceous Herein et sp. nov. Mesomantispa sibirica Baissa, Russia Lower Cretaceous Makarkin 1997; Wedmann and Makarkin 2007 Promantispa similis Karatau, Kazakhstan Upper Jurassic Panfilov 1980 Clavifemora rotundata gen. Jiulongshan Formation, China Middle Jurassic Herein et sp. nov. Liassochrysa stigmatica Dobbertin, Germany Lower Jurassic Ansorge and Schlu¨ter 1990 Baltic amber (Ohl 2011), the Eocene–Oligocene Bem- It contains an exceptionally preserved diverse insect bridge Marls of the Isle of Wight, England, with Vectispa fauna, composed of complete specimens of Ephemerop- relicta Cockerell, 1921(Jarzembowski 1980), Prosagittalata tera, Odonata, Plecoptera, Blattodea, Orthoptera, Hemip- oligocenica Nel, 1989 from the Late Oligocene of Ce´reste tera, Neuroptera, Coleoptera, Hymenoptera and Diptera and Climaciella? henrotayi Nel, 1989 from Dauphin, described by Hong (1983), Ren et al. (1995), Ren and France, and the youngest from Dominican and Mexican Krzeminski (2002). In addition to the insects, there are ambers (Feroseta priscus Poinar, 2006, Dicromantispa mor- freshwater conchostracans (Zhang and Shen 1987), sala- onei Engel and Grimaldi, 2007 and D. electromexicana manders (Gao and Shubin 2003) and dinosaurs (Ji and Engel and Grimaldi, 2007). Here we describe the first Yuan 2002). The surrounding gymnosperm forests were Mesomantispinae mantidflies from the Mesozoic of dominated by Ginkgopsida, Coniferopsida, Lycoposida, China. The new specimens are remarkably well preserved Sphenopsida, Filicopsida and Cycadopsida (Mi et al. and include some specimens with almost complete bodies, 1996). The wealth of palaeontological data has led to confirming their placement in Mantispidae and providing the interpretation of a humid and warm-temperate cli- additional morphological characters to further character- mate (Tan and Ren 2002). Geochemical dating (using ize the subfamily Mesomantispinae. Ar–Ar and SHRIMP U-Pb techniques) has given the age of intermediate-acid volcanic rocks, which overly the Daohugou fossil-bearing beds as 164–165 Ma, therefore GEOLOGICAL SETTINGS the age of the fossil-bearing beds is older or equal to 165 Ma (Chen et al. 2004). This gives a Middle Jurassic The Jiulongshan Formation is a lacustrine sequence that (Bathonian) age of the Daohugou biota (Zhang and crops out in north-eastern China (Ren et al. 1995; Tan Shen 1987; Ren et al. 1995, 2002; Wang 2000; Shen and Ren 2002). The section at Daohugou Village is et al. 2003; Gao and Ren 2006; Shih et al. 2009; Ren composed of grey tuffaceous sandstone and sandy mud- et al. 2010). stone. The palaeoenvironment has been reconstructed as The Yixian Formation of north-eastern China, together a volcanic region with mountain streams and lakes (Ren with the overlying Jiufotang Formation, has yielded a et al. 2002). General reviews of the Middle Jurassic Yan- remarkable array of exceptionally preserved fossils that liao insect fauna of north China have been given by comprise the ‘Jehol Biota’ (Barrett 2000; Chang et al. Hong (1983), Ren et al. (1995) and Ren and Lu (1996). 2003; Zhou et al. 2003). These two conformable and JEPSON ET AL.: NEW MESOZOIC MANTISPIDAE FROM CHINA 605 lithologically similar formations comprise a sequence of Changes, College of Life Sciences, Capital Normal University, low-energy freshwater lacustrine deposits intercalated with Beijing (CNUB). extrusive basalts and tuffaceous sediments deposited in a geologically complex region close to the Pacific Rim. Both formations are Lower Cretaceous in age, with the Barre- SYSTEMATIC PALAEONTOLOGY mian Yixian Formation and the Aptian Jiufotang Forma- tion (Zhou et al. 2003; Sha 2007). Order NEUROPTERA Linnaeus, 1758 The Jehol Lagersta¨tte achieved international celebrity Suborder HEMEROBIIFORMIA sensu Aspo¨ ck et al., 2001 following the discovery of its diverse vertebrate assem- Family MANTISPIDAE Leach, 1815 blage which includes feathered theropod dinosaurs (Chen Subfamily MESOMANTISPINAE Makarkin, 1997 et al. 1998; Ji et al.
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