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HIGHER LEVEL SYSTEMATICS AND EVOLUTION OF THE SPIDER FAMILY OONOPIDAE (ARANEAE) AND THE GENUS OPOPAEA IN MADAGASCAR Open Access

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ABSTRACTThe family of Oonopidae is one of the most diverse and yet poorly known families of haplogyne spiders. Knowledge about oonopids has expanded tremendously since the Planetary Biodiversity Inventory (PBI) project. This dissertation is part of the efforts to document the diversity of oonopids in Madagascar and also to test some hypotheses about the higher level relationships of Oonopidae. This thesis is organized in four chapters: In Chapter 1, the Opopaea fauna of Madagascar is documented for the first time. The genus Opopaea Simon 1891 is one of the most diverse and widespread genus among goblin spiders. There are 27 species of Opopaea on the island of which 26 are newly described here and 25 are apparently endemic to Madagascar: Opopaea andranomay n. sp., O. ankarafantsika n. sp., O. ankarana n. sp., O. antsalova n. sp., O. andringitra n. sp., O. antsiranana n. sp., O. bemarivo n. sp., O. bemaraha n. sp., O. berenty n. sp., O. betioky n. sp., O. itampolo n. sp., O. kirindy n. sp., O. manderano n. sp., O. mahafaly n. sp., O. manongarivo n. sp., O. namoroka n. sp., O. sandranantitra n. sp., O. torotorofotsy n. sp., O. tsimaloto n. sp., O. tsimbazaza n. sp., O. tsimembo n. sp., O. tsinjoriaky n. sp., O. tsingy n. sp., O. vohibazaha n. sp., O. foulpointe n. sp. and O. maroantsetra n. sp. (shared with Kenya and the Comoros Islands), and O. concolor (Blackwall, 1859), a cosmopolitan species. All species are described and illustrated. An identification key to the species and maps of their distribution in Madagascar are also provided.Chapter 2 proposes the first character based phylogenetic hypothesis for Opopaea to test the monophyly of this genus, to examine its relationships with other goblin spider genera such as Epectris Simon 1893, Camptoscaphiella Caporiacco 1934, Brignolia Dumitrescu and Georgescu 1983, Cavisternum Baehr et al. 2010, Prethopalpus Baehr et al. 2012, Malagiella Ubick and Griswold 2011, Tolegnaro Alvarez et al. 2012 and Ischnothyreus Simon 1891which have evolved a heavily sclerotized palps like Opopaea, and to assess the placement of Opopaea within Oonopidae. Opopaea was recovered as a well-supported monophyletic genus sister to Epectris. Opopaea, Epectris and Brignolia form a clade sister to rest of the oonopid genera with sclerotized palps. Likewise, Ischnothyreus, Camptoscaphiella and Malagiella are supported as monophyletic group in which Camptoscaphiella and Malagiella are sister taxa.Chapter 3 addresses three central problems regarding the higher level systematics of Oonopidae: the monophyly and exact placement of Oonopidae among haplogyne spiders, the validity of the two hypotheses of sub-familial classification within Oonopidae, and the monophyly and phylogenetic relationships of the various oonopid genera and groups. Within Haplogynae, Oonopidae was recovered and well supported monophyletic family within the super-family Dysderoidea. Oonopidae sister relationship with the family Orsolobidae was confirmed by the present analysis as well as the monophyly of the super-family Dysderoidea sister to members of the family Caponiidae. Members of the families Scytodidae, Ochyroceratidae and Telemidae were recovered as a monophyletic group, in which Usophila pacifica and Ochyrocera. sp were sister taxa. Tetrablemma cambridgei was recovered as their sister taxa. Similarly, Artema doriai, Kibramoa guapa and Diguetia. sp were recovered monophyletic, in which Kibramoa guapa and Artema doriai were sister taxa. Kukulcania hibernalis is the most basal taxa, sister to all haplogynes. According to our analyses, there are apparently six major clades and eleven groups of genera within Oonopidae. Moreover, the two proposed sub-familial division based on the presence of dorsal scutum and tarsal organ structure were partially supported by our analyses which suggest that there is an increased cuticle sclerotization in Oonopids that starts with the carapace and passes through several intermediate states in which males are always in a more advanced stages compared to females rather than a clear shift from molle to scutate state.

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