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      A genus-level taxonomic review of primitively segmented spiders (Mesothelae, Liphistiidae)

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          Abstract

          Abstract

          The spider suborder Mesothelae , containing a single extant family Liphistiidae , represents a species-poor and ancient lineage. These are conspicuous spiders that primitively retain a segmented abdomen and appendage-like spinnerets. While their classification history is nearly devoid of phylogenetic hypotheses, we here revise liphistiid genus level taxonomy based on original sampling throughout their Asian range, and on the evidence from a novel molecular phylogeny. By combining morphological and natural history evidence with phylogenetic relationships in the companion paper, we provide strong support for the monophyly of Liphistiidae , and the two subfamilies Liphistiinae and Heptathelinae . While the former only contains Liphistius Schiödte, 1849, a genus distributed in Indonesia (Sumatra), Laos, Malaysia, Myanmar, Thailand, we recognize and diagnose seven heptatheline genera, all but three removed from the synonymy of Heptathela : i) Ganthela Xu & Kuntner, gen. n. with the type species Ganthela yundingensis Xu, sp. n. is known from Fujian and Jiangxi, China; ii) a rediagnosed Heptathela Kishida, 1923 is confined to the Japanese islands (Kyushu and Okinawa); iii) Qiongthela Xu & Kuntner, gen. n. with the type species Qiongthela baishensis Xu, sp. n. is distributed disjunctly in Hainan, China and Vietnam; iv) Ryuthela Haupt, 1983 is confined to the Ryukyu archipelago (Japan); v) Sinothela Haupt, 2003 inhabits Chinese areas north of Yangtze; vi) Songthela Ono, 2000 inhabits southwest China and northern Vietnam; and vii) Vinathela Ono, 2000 ( Abcathela Ono, 2000, syn. n.; Nanthela Haupt, 2003, syn. n.) is known from southeast China and Vietnam.

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          Evolution of arthropod silks.

          Silks belong to the class of molecules called structural proteins. The ability to produce silk proteins has evolved multiple times in the arthropods, and silk secreting glands have evolved via two different pathways. The comparative data and phylogenetic analyses in this review suggest that the silk-secreting systems of spiders and insects are homologous and linked to the crural gland (origin of systemic pathway to silk production) and cuticular secretions (origin of surficial pathway to silk production) of an onychophoran-like ancestor. The evolution of silk secreting organs via a surficial pathway is possible in adult and larval hexapods, regardless of their developmental mode. Silk secretion via a systemic pathway is possible in either adult or larval hexapods, but only larval insects have dedicated silk producing glands. Spiders, however, have evolved silk producing systems via both systemic pathway and surficial pathways, and a single individual retains both throughout its lifespan. Early in the evolution of spiders, silk glands were undifferentiated, suggesting that the number of silk secreting glands of any individual was related to the spider's energetic need to produce large quantities of protein. However, the complex silk-producing systems that characterize the aerial web-building spiders and the diverse types of proteins they produce suggest that their silks reflect the diverse and increasing number of ways in which spiders use them. Because the muscular and innervated spinnerets and spigots of spiders allow them to control fiber functional properties, silk proteins represent an avenue through which animal behavior may directly affect the molecular properties of a protein.
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            The Role of Behavior in the Evolution of Spiders, Silks, and Webs

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              Early Events in the Evolution of Spider Silk Genes

              Silk spinning is essential to spider ecology and has had a key role in the expansive diversification of spiders. Silk is composed primarily of proteins called spidroins, which are encoded by a multi-gene family. Spidroins have been studied extensively in the derived clade, Orbiculariae (orb-weavers), from the suborder Araneomorphae (‘true spiders’). Orbicularians produce a suite of different silks, and underlying this repertoire is a history of duplication and spidroin gene divergence. A second class of silk proteins, Egg Case Proteins (ECPs), is known only from the orbicularian species, Lactrodectus hesperus (Western black widow). In L. hesperus, ECPs bond with tubuliform spidroins to form egg case silk fibers. Because most of the phylogenetic diversity of spiders has not been sampled for their silk genes, there is limited understanding of spidroin gene family history and the prevalence of ECPs. Silk genes have not been reported from the suborder Mesothelae (segmented spiders), which diverged from all other spiders >380 million years ago, and sampling from Mygalomorphae (tarantulas, trapdoor spiders) and basal araneomorph lineages is sparse. In comparison to orbicularians, mesotheles and mygalomorphs have a simpler silk biology and thus are hypothesized to have less diversity of silk genes. Here, we present cDNAs synthesized from the silk glands of six mygalomorph species, a mesothele, and a non-orbicularian araneomorph, and uncover a surprisingly rich silk gene diversity. In particular, we find ECP homologs in the mesothele, suggesting that ECPs were present in the common ancestor of extant spiders, and originally were not specialized to complex with tubuliform spidroins. Furthermore, gene-tree/species-tree reconciliation analysis reveals that numerous spidroin gene duplications occurred after the split between Mesothelae and Opisthothelae (Mygalomorphae plus Araneomorphae). We use the spidroin gene tree to reconstruct the evolution of amino acid compositions of spidroins that perform different ecological functions.
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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2015
                21 March 2015
                : 488
                : 121-151
                Affiliations
                [1 ]Centre for Behavioural Ecology and Evolution (CBEE), and Hubei Collaborative Innovation Center for Green Transformation of Bio-Resources, College of Life Sciences, Hubei University, Wuhan, 430062, China
                [2 ]Department of Biological Sciences, National University of Singapore, Singapore
                [3 ]Evolutionary Zoology Laboratory, Biological Institute ZRC SAZU, Ljubljana, Slovenia
                [4 ]Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C, USA
                [5 ]Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba-shi, Ibaraki-ken, 305-0005, Japan
                Author notes
                Corresponding author: Daiqin Li ( dbslidq@ 123456nus.edu.sg ); Matjaž Kuntner ( kuntner@ 123456gmail.com )

                Academic editor: J. Miller

                Article
                10.3897/zookeys.488.8726
                4389128
                0b8d2c93-d70b-4cec-99eb-6f3235f79787
                Xin Xu, Fengxiang Liu, Jian Chen, Hirotsugu Ono, Daiqin Li, Matjaž Kuntner

                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.

                History
                : 10 October 2014
                : 14 January 2015
                Categories
                Research Article

                Animal science & Zoology
                east asia,southeast asia,biogeography,classification,trapdoor spiders,living fossils,animalia,araneae,liphistiidae

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