Phylogeny of the Neotropical Hypoctonine Whip-Scorpions (Thelyphonida, Thelyphonidae), with Descriptions of Two New Genera and Species

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Abstract
Thelyphonida Blanchard, 1852, also known as vinegaroons or whip-scorpions, is a small arachnid order with 140 described species contained in a single family, Thelyphonidae Lucas, 1835. Despite being conspicuous and widely distributed across the tropics and subtropics on four continents, knowledge of the order has been slow to advance. Hypoctoninae Pocock, 1899, one of four subfamilies currently recognized and one of two represented in the New World, comprises five genera. Since its inception, Thelyphonellus Pocock, 1894 has remained the only hypoctonine genus occurring in South America, with only four species described prior to the present contribution. The first detailed morphological study and phylogenetic analysis of Thelyphonellus is presented herein. The morphological phylogenetic analysis—the first for Thelyphonida—includes all except one of the previously described species of Thelyphonellus in addition to two new species described herein; the species of Ravilops Víquez and Armas, 2005 (from the Caribbean island of Hispaniola); and the monotypic Old World genus Etienneus Heurtault, 1984 (from West Africa) scored for 45 morphological characters. A single, most parsimonious phylogenetic hypothesis revealed that Thelyphonellus is paraphyletic with respect to Ravilops. The New World Hypoctoninae comprises four clades with disjunct distributions and well supported by a combination of morphological characteristics, on the basis of which four genera, two of which are new, are recognized: Ravilops, with two species, endemic to Hispaniola; Thelyphonellus, herein restricted to Thelyphonellus amazonicus (Butler, 1872) and Thelyphonellus ruschii Weygoldt, 1979, occurring in Guyana, Suriname, French Guiana, and northern Brazil; Wounaan, gen. n., containing Wounaan vanegasae (Giupponi and Vasconcelos, 2008), comb. n. and Wounaan yarigui, sp. n. from Colombia; and Yekuana, gen. n., containing Yekuana venezolensis (Haupt, 2009), comb. n. and Yekuana wanadi, sp. n. from Venezuela. The two new species are described and illustrated. A key to the identification of the Neotropical genera of Hypoctoninae and a map plotting the known distribution of its species are also presented.
Funding Information
  • Theodore Roosevelt Postdoctoral Research Fellowship from the AMNH Richard Gilder Graduate School (RGGS) (DEB 1655050, DEB 2003382, GB-TAF-8209)
  • National Science Foundation (DEB 1655050, DEB 2003382, GB-TAF-8209)
  • National Science Foundation (DEB 1655050, DEB 2003382, GB-TAF-8209)
  • European Union SYNTHESYS+ Transnational Access (DEB 1655050, DEB 2003382, GB-TAF-8209)
  • AMNH (DEB 1655050, DEB 2003382, GB-TAF-8209)

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