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      Systematics of the Madagascar Anelosimus spiders: remarkable local richness and endemism, and dual colonization from the Americas

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          Abstract

          Abstract

          Despite the alarming rates of deforestation and forest fragmentation, Madagascar still harbors extraordinary biodiversity. However, in many arthropod groups, such as spiders, this biodiversity remains mostly unexplored and undescribed. The first subsocial Madagascan species of the theridiid spider genus Anelosimus were described in 2005 when six new species were found to coexist in the Périnet forest fragment within Andasibe-Mantadia NP. However, this discovery was based only on a few specimens and the extent of this Madagascan radiation has remained unknown. We here report on a thorough survey of >350 colonies from Périnet, and three pilot surveys into additional Madagascar forests (Ambohitantely, Ranamofana, and Montagne d’Ambre). The morphological, molecular and natural history data from these surveys facilitated a revised taxonomy and phylogenetic hypothesis of Madagascan Anelosimus . This subsocial clade currently comprises six previously known ( Anelosimus andasibe Agnarsson & Kuntner, 2005, Anelosimus may Agnarsson, 2005, Anelosimus nazariani Agnarsson & Kuntner, 2005, Anelosimus sallee Agnarsson & Kuntner, 2005, Anelosimus salut Agnarsson & Kuntner, 2005, Anelosimus vondrona Agnarsson & Kuntner, 2005) and 10 new species: Anelosimus ata sp. n., Anelosimus buffoni sp. n., Anelosimus darwini sp. n., Anelosimus hookeri sp. n., Anelosimus huxleyi sp. n., Anelosimus lamarcki sp. n., Anelosimus moramora sp. n., Anelosimus tita sp. n., Anelosimus torfi sp. n., Anelosimus wallacei sp. n.. With the exception of Anelosimus may and Anelosimus vondrona , all other species appear to be single forest endemics. While additional sampling is necessary, these data imply a much higher local richness and endemism in Madagascan forests than in any other comparable area globally. The phylogenetic results establish a sister clade relationship between the subsocial Anelosimus in Madagascar and the American ‘ eximius group’, and between the solitary Anelosimus decaryi on Madagascar and a solitary American clade. These findings imply duplicate colonizations from America, an otherwise rare biogeographical pattern, calling for more detailed investigation of Anelosimus biogeography.

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          MRBAYES: Bayesian inference of phylogenetic trees.

          The program MRBAYES performs Bayesian inference of phylogeny using a variant of Markov chain Monte Carlo. MRBAYES, including the source code, documentation, sample data files, and an executable, is available at http://brahms.biology.rochester.edu/software.html.
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            Arthropods on islands: colonization, speciation, and conservation.

            Islands have traditionally been considered to be any relatively small body of land completely surrounded by water. However, their primary biological characteristic, an extended period of isolation from a source of colonists, is common also to many situations on continents. Accordingly, theories and predictions developed for true islands have been applied to a huge array of systems, from rock pools, to single tree species in forests, to oceanic islands. Here, we examine the literature on islands in the broadest sense (i.e., whether surrounded by water or any other uninhabitable matrix) as it pertains to terrestrial arthropods. We categorize islands according to the features they share. The primary distinction between different island systems is "darwinian" islands (formed de novo) and "fragment" islands. In the former, the islands have never been in contact with the source of colonists and have abundant "empty" ecological niche space. On these islands, species numbers will initially increase through immigration, the rate depending on the degree of isolation. If isolation persists, over time species formation will result in "neo-endemics." When isolation is extreme, the ecological space will gradually be filled through speciation (rather than immigration) and adaptive radiation of neo-endemics. Fragment islands are fundamentally different. In these islands, the ecological space will initially be filled as a consequence of connection to the source of colonists prior to insularization. Species numbers will decrease following fragmentation through the process of relaxation. If these islands become more isolated, species will eventually arise through relictualization with the formation of "paleo-endemics." Given sufficient time, this process can result in generic level endemism on ancient fragment islands, a phenomenon well illustrated in Madagascar and New Zealand. Recognizing the distinction between the different kinds of islands is fundamental for understanding emerging patterns on each, in particular speciation, biodiversity (e.g., neo-endemics versus paleo-endemics), and conservation (e.g., naiveté in interactions with alien species).
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              Has Vicariance or Dispersal Been the Predominant Biogeographic Force in Madagascar? Only Time Will Tell

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                Author and article information

                Journal
                Zookeys
                Zookeys
                ZooKeys
                ZooKeys
                Pensoft Publishers
                1313-2989
                1313-2970
                2015
                22 June 2015
                : 509
                : 13-52
                Affiliations
                [1 ]Department of Biology, University of Vermont, Burlington, VT, USA
                [2 ]Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA
                [3 ]Department of Entomology, University of Madagascar, Antananarivo, Madagascar
                [4 ]Department of Biology, University of Puerto Rico, Rio Piedras, Puerto Rico, USA
                [5 ]Department of Biological Sciences, National University of Singapore, Singapore
                [6 ]Department of Biological Sciences, University of Pittsburgh, PA, USA
                [7 ]Institute of Biology, Scientific Research Centre, Slovenian Academy of Sciences and Arts, Ljubljana, Slovenia
                [8 ]Centre for Behavioural Ecology and Evolution (CBEE), College of Life Sciences, Hubei University, Wuhan, Hubei, China
                Author notes
                Corresponding author: Ingi Agnarsson ( iagnarsson@ 123456gmail.com )

                Academic editor: Cor Vink

                Article
                10.3897/zookeys.509.8897
                4493342
                cb8a916c-b492-47d8-b3bb-9004fa7c17bc
                Ingi Agnarsson, Brian B. Jencik, Giselle M. Veve, Sahondra Hanitriniaina, Diego Agostini, Seok Ping Goh, Jonathan Pruitt, 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
                : 5 November 2014
                : 5 June 2015
                Categories
                Research Article

                Animal science & Zoology
                cobweb spiders,subsocial,theridiidae,biogeography,colonization,radiation,congener coexistance,animalia,araneae

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