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Coleoptera: Elateroidea: Artematopodidae) from the Early Cretaceous of China and Its Systematic Position

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Coleoptera: Elateroidea: Artematopodidae) from the Early Cretaceous of China and Its Systematic Position Cretaceous Research xxx (2018) 1e7 Contents lists available at ScienceDirect Cretaceous Research journal homepage: www.elsevier.com/locate/CretRes A large artematopodid beetle (Coleoptera: Elateroidea: Artematopodidae) from the Early Cretaceous of China and its systematic position * Chenyang Cai a, b, Yanzhe Fu c, Diying Huang c, a CAS Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing 210008, China b School of Earth Sciences, University of Bristol, Life Sciences Building, Tyndall Avenue, Bristol BS8 1TQ, UK c State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Center for Excellence in Life and Paleoenvironment, Chinese Academy of Sciences, Nanjing 210008, China article info abstract Article history: Artematopodidae is small basal group of elateroid beetles. Definitive Mesozoic artematopodids are Received 9 August 2018 currently confined to the Middle Jurassic Daohugou biota. Here we report a new genus and species, Received in revised form Cretobrevipogon breviantennatus gen.et sp. nov., from the Lower Cretaceous Yixian Formation (ca. 125 Ma) 30 September 2018 in Liaoning Province, northeastern China. Cretobrevipogon exhibit many diagnostic features of the extant Accepted in revised form 10 October 2018 family Artematopodidae, such as paired carinae on prosternum and more importantly, an apical inter- Available online xxx locking tongue on the ventral side of elytron. Phylogenetic analyses indicate that Cretobrevipogon is closely related to the Middle Jurassic Sinobrevipogon. They share a presumably plesiomorphic character, Keywords: Palaeodiversity i.e. mesocoxal cavitiy closed by mesepimeron and anterolateral edge of metanepisternum. The discovery Phylogeny of a new genus from the Early Cretaceous highlights the morphological disparity and palaeodiversity of Elateroidea the family in the Mesozoic. Artematopodidae probably represent a relictual elateroid family that once Yixian Formation diversified in the middleelate Mesozoic. Jehol biota © 2018 Elsevier Ltd. All rights reserved. 1. Introduction eastern Russian, Japan; Macropogon Motschulsky, North America, Asia; and Proartematopus Crowson, Eocene Baltic amber) (Crowson, The family Artematopodidae is small group of elateroid beetles, 1973; Lawrence, 1995, 2005; 2010; Young, 2002; Wu et al., 2015). with approximately 70 described species in nine extant genera Among all elateroid beetles, adults of Artematopodidae can be (Lawrence, 2005, 2010). Recent molecular-based phylogenetic an- readily recognized by a peculiar feature, namely a tongue-like alyses strongly indicate that Artematopodidae, together with process associated with the apicoventral region of the elytron Omethidae and Telegeusidae, forms the basal-most lineage of Ela- (Lawrence, 2005). teroidea (Kundrata et al., 2014; McKenna et al., 2015; Zhang et al., Fossils belonging to Artematopodidae are poorly documented. 2018). Artematopodidae is now divided into three subfamilies: Mesozoic artematopodids have recently reported from the Mid- Electribiinae (Electribius Crowson, Mexico, Central America, Eocene dle Jurassic Daohugou beds (Inner Mongolia, northeastern Baltic amber, and Miocene Mexican amber); Allopogoninae (Allo- China). Jurassic artematopodids currently include two genera: pogonia Cockerell, California); and Artematopodinae (Artematopus Sinobrevipogon Cai et al. and Tarsomegamerus Zhang (Cai et al., Perty, Central and South America; Brevipogon Lawrence, California; 2015). Cai et al. (2015) also indicated that the unusual species, Carcinognathus Kirsch, South America; Ctesibius Champion, Mexico Forticatinius elegans Tan and Ren, from the Lower Cretaceous and Central America; Eurypogon Motschulsky, North America, Italy, Yixian Formation is closely related to members of Artematopo- didae, rather than a member of Myxophaga as originally pro- posed by Tan and Ren (2007). Peris et al. (2016) listed an undescribed artematopodid from the mid-Cretaceous amber * Corresponding author. Nanjing Institute of Geology and Palaeontology, Chinese from France. Cenozoic artematopodids are principally known Academy of Sciences, 39 East Beijing Road, Nanjing 210008, China. from the upper Eocene Baltic amber (Lawrence, 2010). Two E-mail address: [email protected] (D. Huang). https://doi.org/10.1016/j.cretres.2018.10.009 0195-6671/© 2018 Elsevier Ltd. All rights reserved. Please cite this article in press as: Cai, C., et al., A large artematopodid beetle (Coleoptera: Elateroidea: Artematopodidae) from the Early Cretaceous of China and its systematic position, Cretaceous Research (2018), https://doi.org/10.1016/j.cretres.2018.10.009 2 C. Cai et al. / Cretaceous Research xxx (2018) 1e7 genera are known from this deposit: Electribius,withfourextinct Zhang et al., 2018). All selected characters were derived from Cai species and two extant ones and the monotypic genus Proarte- et al. (2015). Exemplar taxa for phylogenetic analyses were matopus (Crowson, 1973; Lawrence, 1995; Hornschemeyer,€ 1998). selected from adults only, since larvae remains largely unknown. Wu et al. (2015) reported the youngest fossil artematopodid, Exemplars included: 1) Two fossil species: Sinobrevipogon jurassicus Electribius palaeomexicanus Wu et al., from the Miocene Mexican Cai et al. from the Middle Jurassic and C. breviantennatus from the amber, and the intriguing biogeographic implication of Electribius Early Cretaceous; 2) species belonging all nine extant artematopodid was tentatively discussed. Here we report the first definitive genera; and 3) Declinia relicta Nikitsky et al. (Decliniidae) and Nip- artematopodid fossil from the Lower Cretaceous Yixian Forma- ponocyphon nakanei Lawrence and Yoshitomi (Scirtidae) as the out- tion (Jehol biota). The systematic position of the new genus is group in the phylogenetic analyses. discussed. Maximum parsimony analysis was conducted using TNT 1.5 (Goloboff and Catalano, 2016) using New Technology search. We 2. Material and methods used two search strategies, including equal weights and implied weights. Goloboff et al. (2018) suggested that equal-weights Fossil material. The studied specimen, with part and counterpart, parsimony and model-based methods (Mk model; Lewis, 2001) has been collected from the yellowish tuff of Jingangshan Member seem to perform similarly, and implied weighting outperformed of Yixian Formation near Huangbanjigou Village (Fig. 1; Beipiao, equal weights. As suggested in Goloboff et al. (2018), better Liaoning Province, northeastern China; also refer to Cai et al., 2011). phylogenetic results can be obtained using implied weighting un- This stratum (Yixian Formation) represents a typical component of der larger concavity values of K (K ¼ 12). For the implied weighting the famous Jehol biota well known for yielding feathered dinosaurs, analysis, we tested a concavity value of 12 and found no changes to mammals, primitive birds, angiosperm plants and numerous in- the tree topology under the equally weighting analysis. Branch sects (e.g., Chang, 2003). The precise age of the Yixian Formation support values were estimated using 10,000 bootstrap replicates has been indicated as circa 125 million years ago (Swisher III et al., under implied weighting (K ¼ 12). 1999). The fossil beetle has been split through the median gap into two counterparts. Although characters of both ventral and dorsal Out-group taxa. sides are imprinted on both counterparts, one part (NIGP168714a) Declinia Nikitsky et al., data from Nikitsky et al. (1993), and better preserves characters of the dorsal side, whereas the other Lawrence et al. (1995). (NIGP168714b) displays more characters of the ventral side. The Nipponocyphon Lawrence and Yoshitomi, data from Lawrence and holotype is deposited in the Nanjing Institute of Geology and Yoshitomi (2007) and dissection of paratype of N. nakanei. Palaeontology (NIGP), Chinese Academy of Sciences (Nanjing, In-group taxa. Jiangsu Province). The specimen was examined both dry (under Allopogonia Cockerell, data from dissection of A. villosus Horn. low-angled light) and moistened with 70% alcohol (in order to Artematopus Perty, data from Crowson (1973) and dissections of improve contrast). Photographs were taken using a Zeiss Discovery several species. V20 microscope with a digital camera attached. Extended depth of Brevipogon Lawrence, data from Lawrence (2005) and dissections of field images were digitally compiled using Helicon Focus 3.10 B. confuisus (Fall). software, and arranged in Adobe Photoshop CS5. The nomenclatural Carcinognathus Kirsch, data from Crowson (1973) and dissections of acts established herein are registered under ZooBank LSID: urn:l- one species. sid:zoobank.org:pub:C87042E6-763C-4241-86DF-155707337F5D. Ctesibius Champion, data from Lawrence (1995) taken from holo- Phylogenetic analyses. We scored Cretobrevipogon breviantennatus type of C. eumolpoides Champion. sp. nov. for 30 adult characters and integrated the fossil taxon along Electribius Crowson, data from Crowson (1973), Lawrence (1995, with representatives of extant and extinct genera of Artematopodi- 2005) and Hornschemeyer€ (1998) and dissection of holotype of dae and two scirtoid outgroups (basal-most polyphagan families as E. crowsoni Lawrence. suggested by molecular-based phylogenies, McKenna et al., 2015; Fig. 1. Map of western Liaoning Province, with the solid star indicating the fossil locality at
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