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      A revision of dragon millipedes I: genus Desmoxytes Chamberlin, 1923, with the description of eight new species (Diplopoda, Polydesmida, Paradoxosomatidae)

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          Abstract

          Abstract

          The dragon millipede genus Desmoxytes s.l. is split into five genera, based on morphological characters and preliminary molecular phylogenetic analyses. The present article includes a review of Desmoxytes s.s., while future articles will deal with Hylomus Cook and Loomis, 1924 and three new genera which preliminarily are referred to as the ‘ acantherpestes’, ‘ gigas’, and ‘ spiny’ groups. Diagnostic morphological characters of each group are discussed. Hylomus is resurrected as a valid genus and the following 33 species are assigned to it: H. asper (Attems, 1937), comb. n., H. cattienensis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. cervarius (Attems, 1953), comb. n., H. cornutus (Zhang & Li, 1982), comb. n., H. draco Cook & Loomis, 1924, stat. rev., H. enghoffi (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. eupterygotus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. getuhensis (Liu, Golovatch & Tian, 2014), comb. n., H. grandis (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. hostilis (Golovatch & Enghoff, 1994), comb. n., H. jeekeli (Golovatch & Enghoff, 1994), comb. n., H. lingulatus (Liu, Golovatch & Tian, 2014), comb. n., H. laticollis (Liu, Golovatch & Tian, 2016), comb. n., H. longispinus (Loksa, 1960), comb. n., H. lui (Golovatch, Li, Liu & Geoffroy, 2012), comb. n., H. minutuberculus (Zhang, 1986), comb. n., H. nodulosus (Liu, Golovatch & Tian, 2014), comb. n., H. parvulus (Liu, Golovatch & Tian, 2014), comb. n., H. phasmoides (Liu, Golovatch & Tian, 2016), comb. n., H. pilosus (Attems, 1937), comb. n., H. proximus (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. rhinoceros (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. rhinoparvus (Likhitrakarn, Golovatch & Panha, 2015), comb. n., H. scolopendroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. scutigeroides (Golovatch, Geoffroy & Mauriès, 2010), comb. n., H. similis (Liu, Golovatch & Tian, 2016), comb. n., H. simplex (Golovatch, VandenSpiegel & Semenyuk, 2016), comb. n., H. simplipodus (Liu, Golovatch & Tian, 2016), comb. n., H. specialis (Nguyen, Golovatch & Anichkin, 2005), comb. n., H. spectabilis (Attems, 1937), comb. n., H. spinitergus (Liu, Golovatch & Tian, 2016), comb. n., H. spinissimus (Golovatch, Li, Liu & Geoffroy, 2012), comb. n. and H. variabilis (Liu, Golovatch & Tian, 2016), comb. n. Desmoxytes s.s. includes the following species: D. breviverpa Srisonchai, Enghoff & Panha, 2016; D. cervina (Pocock,1895); D. delfae (Jeekel, 1964); D. des Srisonchai, Enghoff & Panha, 2016; D. pinnasquali Srisonchai, Enghoff & Panha, 2016; D. planata (Pocock, 1895); D. purpurosea Enghoff, Sutcharit & Panha, 2007; D. takensis Srisonchai, Enghoff & Panha, 2016; D. taurina (Pocock, 1895); D. terae (Jeekel, 1964), all of which are re-described based mainly on type material. Two new synonyms are proposed: Desmoxytes pterygota Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes cervina (Pocock, 1895)), Desmoxytes rubra Golovatch & Enghoff, 1994, syn. n. (= Desmoxytes delfae (Jeekel, 1964)). Six new species are described from Thailand: D. aurata Srisonchai, Enghoff & Panha, sp. n., D. corythosaurus Srisonchai, Enghoff & Panha, sp. n., D. euros Srisonchai, Enghoff & Panha, sp. n., D. flabella Srisonchai, Enghoff & Panha, sp. n., D. golovatchi Srisonchai, Enghoff & Panha, sp. n., D. octoconigera Srisonchai, Enghoff & Panha, sp. n., as well as one from Malaysia: D. perakensis Srisonchai, Enghoff & Panha, sp. n., and one from Myanmar: D. waepyanensis Srisonchai, Enghoff & Panha, sp. n. The species can mostly be easily distinguished by gonopod structure in combination with other external characters; some cases of particularly similar congeners are discussed. All species of Desmoxytes s.s. seem to be endemic to continental Southeast Asia (except the ‘tramp’ species D. planata ). Some biological observations (relationship with mites, moulting) are recorded for the first time. Complete illustrations of external morphological characters, an identification key, and distribution maps of all species are provided.

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              Discovery of a glowing millipede in California and the gradual evolution of bioluminescence in Diplopoda.

              The rediscovery of the Californian millipede Xystocheir bistipita surprisingly reveals that the species is bioluminescent. Using molecular phylogenetics, we show that X. bistipita is the evolutionary sister group of Motyxia, the only genus of New World bioluminescent millipedes. We demonstrate that bioluminescence originated in the group's most recent common ancestor and evolved by gradual, directional change through diversification. Because bioluminescence in Motyxia has been experimentally demonstrated to be aposematic, forewarning of the animal's cyanide-based toxins, these results are contrary to aposematic theory and empirical evidence that a warning pattern cannot evolve gradually in unpalatable prey. However, gradual evolution of a warning pattern is plausible if faint light emission served another function and was co-opted as an aposematic signal later in the diversification of the genus. Luminescence in Motyxia stem-group taxa may have initially evolved to cope with reactive oxygen stress triggered by a hot, dry environment and was repurposed for aposematism by high-elevation crown-group taxa colonizing new habitats with varying levels of predation. The discovery of bioluminescence in X. bistipita and its pivotal phylogenetic location provides insight into the independent and repeated evolution of bioluminescence across the tree of life.
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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2018
                29 May 2018
                : 761
                : 1-177
                Affiliations
                [1 ] Biological Sciences Program, Faculty of Science, Chulalongkorn University, Phaya Thai Road, Patumwan, Bangkok 10330, Thailand
                [2 ] Animal Systematics Research Unit, Department of Biology, Faculty of Science, Chulalongkorn University, Phayathai Road, Patumwan, Bangkok 10330, Thailand
                [3 ] Natural History Museum of Denmark, University of Copenhagen, Universitetsparken 15, DK-2100 København Ø, Denmark
                [4 ] Division of Plant Protection, Faculty of Agricultural Production, Maejo University, San Sai, Chiang Mai 50290, Thailand
                Author notes
                Corresponding authors: Somsak Panha ( somsak.pan@ 123456chula.ac.th ); Henrik Enghoff ( henghoff@ 123456snm.ku.dk )

                Academic editor: R. Mesibov

                Article
                10.3897/zookeys.761.24214
                5988806
                Ruttapon Srisonchai, Henrik Enghoff, Natdanai Likhitrakarn, Somsak Panha

                This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                Categories
                Research Article
                Diplopoda
                Paradoxosomatidae
                Polydesmida
                Systematics
                Cenozoic
                Asia

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