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This Article Appeared in a Journal Published by Elsevier. the Attached This article appeared in a journal published by Elsevier. The attached copy is furnished to the author for internal non-commercial research and education use, including for instruction at the authors institution and sharing with colleagues. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. 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) 1325–1338 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes New lacewings (Insecta: Neuroptera) from the Lower Cretaceous Wealden supergroup of Southern England James E. Jepson a,*, Vladimir N. Makarkin b, Edmund A. Jarzembowski c a School of Earth, Atmospheric and Environmental Sciences, University of Manchester, Williamson Building, Oxford Road, Manchester, M13 9PL, UK b Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russian Federation c Maidstone Museum and Bentlif Art Gallery, St Faith’s Street, Maidstone, Kent ME14 1LH, UK article info abstract Article history: Eight new genera and thirteen new species of lacewings (Neuroptera) are described from the Lower Received 5 February 2009 Cretaceous Wealden Supergroup, Weald Sub-basin: Principiala rudgwickensis sp. nov. (Ithonidae), Sten- Accepted in revised form 29 July 2009 omylina medialis gen. et sp. nov., Protosmylina bifasciata gen. et sp. nov. (Osmylidae), Cretapsychops corami Available online 5 August 2009 gen. et sp. nov., Micropsychops parallelus gen. et sp. nov., Valdipsychops logunovi gen. et sp. nov., V. brigidae gen. et sp. nov., V. maculosus gen. et sp. nov., V. proudlovei gen. et sp. nov., V. minimus gen. et sp. nov., Keywords: Psychopsites rolandi gen. et sp. nov. (Psychopsidae), Actinophlebioides valdensis gen. et sp. nov. and Pro- Ithonidae tohemerobius perexiguus gen. et sp. nov. (Neuroptera familia incertae sedis). Psychopsidae are shown to Kalligrammatidae Neuropterida dominate the neuropteran fauna, while the presence of Brongniartiellidae has not been confirmed in the Osmylidae Wealden, due to previously thought representatives of this family being in fact psychopsids. This is the Psychopsidae first fossil record of Ithonidae and Osmylidae in Britain. The range of Kalligrammatidae has been Weald Sub-basin extended in the Cretaceous of Britain from the mid-Valanginian to early Barremian. Ó 2009 Elsevier Ltd. All rights reserved. 1. Introduction been found in the Triassic (Tillyard, 1933), Jurassic (Whalley, 1988), Cretaceous (e.g. Jarzembowski, 1984,1993) and Palaeogene Neuroptera (lacewings) is one of three orders that make up the (Jarzembowski, 1980). superorder Neuropterida (Insecta: Holometabola) (Grimaldi and Engel, 2005) and consists of approximately 6000 described extant species in 17 families (Aspo¨ck, 2002). They are widespread globally, 2. Geological setting but tend to be most abundant in the tropics, with several families having restricted distributions (New, 1991). Sixty-six extant species The Lower Cretaceous Wealden Supergroup (c. 140–125 Ma: Gradstein et al., 2005) of southern England is subdivided into the from six families: Chrysopidae (green lacewings), Coniopterygidae (wax flies), Hemerobiidae (brown lacewings), Myrmeleontidae (ant Weald and Wessex Sub-basins (Allen and Wimbledon, 1991). The non-marine Weald Sub-basin has yielded the most insect remains lions), Osmylidae and Sisyridae (spongillaflies) occur in Britain (Plant, 1997). Prior to this study, the presence of three psychopsoid and it is from this sub-basin that the specimens described herein have been collected (Fig. 1). The Weald Sub-basin consists of the neuropteran families were recorded in the Wealden: Psychopsidae (silky lacewings), and the extinct Brongniartiellidae and Kalli- Hastings Beds Group (Ashdown Beds, Wadhurst Clay, Lower and Upper Tunbridge Wells Sand, and the Grinstead Clay formations) grammatidae (Jarzembowski, 1984; Ross and Cook, 1995) and one species of kalligrammatid has been formally described: Kalli- and the Weald Clay Group (Upper and Lower Weald Clay forma- tions) (Allen and Wimbledon, 1991; Rasnitsyn et al., 1998). The gramma roycrowsoni (Jarzembowski, 2001). Neuroptera have a long and diverse fossil record from the majority of insects and the specimens described herein are from the Weald Clay Group, which are dated as Hauterivian–Aptian? (Allen Permian to Recent with all the major family radiations having occurred prior to the Lower Cretaceous (Ponomarenko, 2002; and Wimbledon, 1991; Rasnitsyn et al., 1998). The Weald Clays Grimaldi and Engel, 2005). In Britain, fossil Neuroptera have represent a lacustrine–lagoonal environment (Allen, 1981,1989; Radley, 2005). The insect orders that have so far been recorded in the Weald Sub-basin are Odonata, ‘‘Blattaria’’, Orthoptera, Hemi- ptera, Neuroptera, Coleoptera, Mecoptera, Diptera, Trichoptera, * Corresponding author. Hymenoptera, Isoptera, Raphidioptera (Jarzembowski, 1984,1995) E-mail address: james.jepson@postgrad.manchester.ac.uk (J.E. Jepson). and Plecoptera (Sinitshenkova, 1998). The insects are preserved as 0195-6671/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2009.07.012 Author's personal copy 1326 J.E. Jepson et al. / Cretaceous Research 30 (2009) 1325–1338 Fig. 1. Geological map of the Wealden Supergroup, southeast England, with locations (brickworks) where Neuroptera have been found marked on. (Redrawn and modified from Gallois, 1978; Ross and Cook, 1995; Rasnitsyn et al., 1998). uncrushed and unoxidized, disarticulated skeletal material, with Oswald (1993a) and Archibald & Makarkin (2006). Wing vein isolated wings being the most common (Jarzembowski, 1984). They abbreviations are as follows: C, costa; Sc, subcosta; R, radius; R1, are found in the calcareous siltstone developments or phosphatic first branch of R; Rs, radial sector; Rs1, most proximal branch of Rs; or sideritic concretions in the clay formations. The climate has been M, media; MA, media anterior; MP, media posterior; Cu, cubitus; interpreted as subtropical/warm temperate with seasonal rainfall CuA, cubitus anterior; CuP, cubitus posterior; 1A–3A, first to third (Allen, 1998; Haywood et al., 2005). The occurrence of wet and dry anal veins. (commonly with wild fires) seasons has been proposed, as has year- round precipitation with little evidence for prolonged drought but 4. Systematic palaeontology high evaporation (Batten, 1998; Haywood et al., 2005). Wildfire evidence exists in the form of fusainized coleopteran elytra and Order: Neuroptera Linnaeus, 1758 plant remains (Watson and Alvin, 1996; Batten, 1998; Jarzembow- Family: Ithonidae Newman, 1853 (¼ Rapismatidae Kru¨ ger, 1922) ski, 2003). Genus Principiala Makarkin et Menon, 2007 Principiala rudgwickensis sp. nov. 3. Material and methods Fig. 2 Derivation of name. After the Rudgwick Brickworks locality The neuropteran material described herein was collected from where the specimen was collected. the Weald Clay Group in various brickworks: Clockhouse (Lower Weald Clay Formation, Hauterivian), Auclaye, Rudgwick and Material. Holotype BMB 025005, -6 (part and counterpart), an Smokejacks (Upper Weald Clay Formation, Barremian) (locations incomplete forewing. Rudgwick Brickworks, Upper Weald Clay, labelled in fig. 1). The material is deposited in the following insti- Lower Cretaceous, Barremian. tutions: Booth Museum of Natural History, Brighton (BMB), Natural Diagnosis. May be distinguished from Principiala incerta by the History Museum, London (BMNH Pal.) and Maidstone Museum and following forewing character states [alternative character states are Bentlif Art Gallery, Kent (MNEMG). given in brackets]: most proximal branch of Rs terminated on MA The material was studied using an Olympus SZH stereomicro- long [very short in P. incerta]; numerous crossveins between scope with a camera lucida attachment for producing drawings. proximal 5 branches of Rs: some veins having a ‘zig-zag’ appear- Some drawings are composites, merging the part and counterpart ance [not detected in P. incerta]; proximal 6 fused branches of Rs are to establish a more complete venation; this was achieved using straight [curved in P. incerta]. a light box. 10% ethanol was used to enhance vein detail. Photo- graphs were taken using a Nikon D1X digital SLR camera mounted Description. Forewing shape indeterminate, preserved wing on a rostrum stand, except Psychopsites rolandi gen. et sp. nov. length 9.4 mm (estimated length about 11–12 mm). C not preserved. which was drawn by EAJ using a camera lucida attachment on Few subcostal veinlets preserved. Gradate series of crossveins in a M4A Wild microscope and photographed by Tony Mitchell costal space partially preserved. Sc partially preserved appearing to (volunteer Maidstone Museum) using a Nikon Coolpix 5000 digital fuse with R1 distally: these veins at least, closely approach each other camera. towards wing apex. 5 subcostal crossveins preserved. R incomplete, We follow the traditional (sensu Wootton, 2003) venational origin and termination not preserved. 2 r-rs crossveins preserved. Rs terminology of Comstock (1918) with the recent interpretation
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