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The First Fossil Rove Beetle from the Middle Eocene Kishenehn Formation

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The First Fossil Rove Beetle from the Middle Eocene Kishenehn Formation TJSP_A_1266402.3d (TJSP) (215£280mm) 28-12-2016 20:43 Queries are marked in the margins of the proofs, and you can also click the hyperlinks below. AUTHOR QUERIES General points: 1. Permissions: You have warranted that you have secured the necessary written permission from the appropriate copyright owner for the reproduction of any text, illustration, or other material in your article. Please see http://journalauthors.tandf.co.uk/ permissions/usingThirdPartyMaterial.asp. 2. Third-party content: If there is third-party content in your article, please check that the rightsholder details for re-use are shown correctly. 3. Affiliation: The corresponding author is responsible for ensuring that address and email details are correct for all the co-authors. Affiliations given in the article should be the affiliation at the time the research was conducted. 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TJSP_A_1266402.3d (TJSP) (215£280mm) 28-12-2016 20:43 Journal of Systematic Palaeontology, 2016 Vol. 0, No. 0, 1–11, http://dx.doi.org/10.1080/14772019.2016.1266402 The first fossil rove beetle from the middle Eocene Kishenehn Formation (North America) provides evidence for ancient Eocene relicts within the hyperdiverse Staphylinini (Coleoptera: Staphylinidae: Staphylininae) Adam J. Brunkea,b*, Harald Schillhammera and Stylianos Chatzimanolisc 5 aNatural History Museum of Vienna, Zoology Department, Burging 7, A – 1010, Vienna, Austria; bCanadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, 960 Carling Avenue, K.W. Neatby Building, Ottawa, K1A 0C6, Canada; cDepartment of Biology, Geology and Environmental Science, University of Tennessee at Chattanooga, 615 McCallie Avenue, Dept. 2653, Chattanooga, Tennessee 37403, USA (Received 18 April 2016; accepted 14 October 2016) 10 A new rove beetle (Coleoptera: Staphylinidae) is described from the middle Eocene Kishenehn Formation in Montana, USA. yTympanophorus greenwalti Chatzimanolis, Brunke & Schillhammer sp. nov. is the oldest known definitive member of the subtribe Anisolinina (Staphylininae: Staphylinini) and the entire Staphylinini propria clade, which contains the majority of the tribe’s over 5500 described extant species. In order to provide robust justification for the systematic placement of the Kishenehn fossil, all genera of the Tympanophorus lineage are reviewed and redefined. A key to these 15 genera is provided for the first time. Paratympanophorus Lecoq becomes a junior synonym of Trigonopalpus Cameron, with the following new combinations: Trigonopalpus africanus (Lecoq), Tr. peyrierasi (Lecoq), Tr. pubescens (Lecoq), Tr. punctatus (Lecoq) and Tr. steineri (Lecoq). Tympanophorus schenklingi Bernhauer is moved to Trigonopalpus (comb. nov.) and Ty. clavicornis (Lecoq) is moved to Barygnathus (comb. nov.), and thus, the genus Tympanophorus no longer occurs in the Afrotropical region. Tympanophorus is shown to be at least as old as the middle Eocene, and its disjunct New 20 and Old World lineages are hypothesized to have been separated in the early Eocene. http://zoobank.org/urn:lsid:zoobank.org:pub: CA1993B8-1251-45C3-877E-C4604F78E781 Keywords: Palaeogene; Lutetian; biogeography; Montana; Anisolinina; Tympanophorus Introduction of taxa were either described in large polyphyletic genera or incorrectly placed systematically at the genus, subtribe The last decade has seen a renaissance in fossil rove beetle or even subfamily level (Brunke, pers. obs.). One exception 25 systematics, with dozens of new species of Staphylinidae is the diverse Early Cretaceous (Yixian Formation, China) 45 coming to light (for papers summarizing most of these assemblage of Staphylinini described and placed in a phy- fossils see: Chatzimanolis & Engel 2011, 2013; Chatzi- logenetic context by Solodovnikov et al. (2013). However, manolis et al. 2012; Solodovnikov et al. 2013; Peris et al. with the evidence currently available, none of these taxa 2014). One of the major barriers to the detailed integration could be confidently placed within extant lineages and they 30 of palaeontological taxa with extant rove beetle lineages is are probably best treated as early diverging stem groups. 50 that the majority of the supra-specific taxa are poorly Resolving the position and demonstrating monophyly defined and are in need of revision. Recently, there has of the Staphylinini subtribe Anisolinina (sensu Schillham- been renewed interest in reconstructing phylogenetic rela- mer 2004) has been particularly problematical, though a tionships within the tribe Staphylinini of Staphylininae recent analysis of molecular data suggests that it may be a 35 (Chatzimanolis et al. 2010; Brunke & Solodovnikov natural group (Brunke et al. 2016). However, morphologi- 55 2013; Brunke et al. 2016), a hyperdiverse group of more cal character evidence for this group as a whole remains than 5500 relatively large and predatory beetle species. elusive (Brunke & Solodovnikov 2013), making the direct While a consensus topology is emerging for Staphylinini assignment of fossils to Anisolinina difficult. At present, and its higher classification is partially revised (e.g. Chat- Anisolinina is a group of 14 poorly known genera (Schill- 40 zimanolis et al. 2010; Brunke et al. 2016), the fossil hammer & Brunke, in prep.) and just over 200 species, 60 record of this group is in need of revision as the majority which is unknown from the fossil record (A. Newton, *Corresponding author. Email: [email protected] Ó The Trustees of the Natural History Museum, London 2016. All rights reserved. TJSP_A_1266402.3d (TJSP) (215£280mm) 28-12-2016 20:43 2 A. J. Brunke et al. unpublished catalogue). The exclusion of Algon Sharp and Philothalpus Kraatz from the subtribe is supported by morphological (Schillhammer 2004) and molecular evi- 65 dence (Chatzimanolis 2014), though their sister groups remain unknown. The systematic position of Rientis Sharp is also unknown but is thought to be near Algon (Schill- hammer 2006). Based on morphological characters of the mesoventrite, 70 and maxillary and labial palpi, Schillhammer (2004) defined two groups of Anisolinina: the Tympanophorus and Anisolinus lineages. While the genera of the latter are diagnosable, the rarely collected genera of the Tympano- phorus lineage are poorly defined and have never been 75 critically re-evaluated. Nearly all genera of Anisolinina are restricted to the Afrotropical and Oriental regions (i.e. the Old World subtropics and tropics). Misantlius and Tympanophorus are exceptional in that they exhibit a dis- junct subtropical to tropical distribution between the Old 80 and New World: Oriental-Neotropical, or Oriental C east Palearctic (Japan and Korea) ¡ Nearctic C Neotropical, Figure 1. Dorsal habitus: A, Tympanophorus greenwalti Chatzi- respectively. The two Afrotropical species of Tympano- manolis, Brunke & Schillhammer sp. nov., USNM 595140; B, Tympanophorus sp. undescribed, from Mexico. Scale bars D phorus have not been re-examined since their descriptions 2 mm. and may belong to other genera. Although currently dis- 85 junct, these distributions in Anisolinina and other lineages of Staphylinini were hypothesized by Brunke & Solodov- UK; FMNH: Field Museum of Natural History, Chicago, nikov (2013) to be formerly widespread in the northern USA; MHNG: Natural History Museum, Geneva, Swit- 115 hemisphere during the Palaeogene/early Eocene, becom- zerland; LNKD: Landessammlung fur€ Naturkunde, Karls- ing relictual after the cooling and polarization of global ruhe, Germany; NHMW: Natural History Museum of 90 climate in the late Eocene/early Oligocene. However, evi- Vienna, Vienna, Austria; USNM: National Museum of dence for an ancient Eocene age of these genera, from Natural History, Washington DC, USA. 120 either fossils or divergence estimates, is currently lacking. Evidence for boreotropical relictualism may be discov- ered amongst the diverse fossil insect faunas of subtropi- Specimen examination and documentation 95 cal to tropical Eocene North America
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