THE SYSTEMATICS OF THE SPIDER GENUS JOSA (ARANEAE: ANYPHAENIDAE: AMAUROBIOIDINAE): A TOTAL EVIDENCE APPROACH

The subfamily Amaurobioidinae currently comprises 22 genera, most of which have restricted distributions in the Andean-Patagonian regions of southern South America. In contrast to this general biogeographic pattern, Josa reaches its greatest diversity in the Tropical Andes, with some species extending into the Yungas of Argentina and Bolivia and into isolated montane systems in Brazil. Here, we present a comprehensive systematic revision of Josa based on a total-evidence phylogenetic framework integrating morphological and molecular sequence data. This revision includes redescriptions of all previously known species when necessary and describes 75 new species, increasing Josa to 94 species and establishing it as the most diverse genus within Amaurobioidinae. Females are distinguished by spermathecae continuous with the anteriorly connected copulatory ducts, which are coiled at least once, and by sclerotized, usually posteriorly projecting extensions of the epigyne. We also propose several nomenclatural changes: Josa insignis Banks, 1909 is removed from synonymy with Josa nigrifrons (Simon, 1897), redescribed, and its male is described for the first time; Gayenna vittata (Keyserling, 1881) is transferred to Josa and redescribed from the female holotype; Josa lojensis (Berland, 1913) is synonymized with Josa personata (Simon, 1897); and Josa maura (Simon, 1897) is considered a nomen dubium because the holotype is an immature female. Complementary descriptions are also provided for Josa personata and Josa bryantae (Caporiacco, 1955), including the first descriptions of their males. In addition, we describe the following 75 new species: Josa winayhuaina, Josa napo, Josa nigra, Josa faivovichi, Josa belmira, Josa celica, Josa michaliki, Josa cisnerosi, Josa minca, Josa adami, Josa inca, Josa laplata, Josa sanfrancisco, Josa laquipampa, Josa ayacucho, Josa quito, Josa acjanacoensis, Josa paisa, Josa pinzoni, Josa paramo, Josa hannahae, Josa tolimensis, Josa bruscai, Josa santosi, Josa mucuyensis, Josa pallida, Josa banos, Josa machupicchuensis, Josa elaguila, Josa betoma, Josa nevada, Josa sanpedro, Josa kruegeri, Josa sorkini, Josa cornuta, Josa kochalkai, Josa magdalenensis, Josa cundinamarquensis, Josa sciosciae, Josa tungurahua, Josa sibundoy, Josa guacharos, Josa ligiae, Josa huamanga, Josa chingaza, Josa toledoensis, Josa perijaensis, Josa merida, Josa azulita, Josa guasca, Josa boyaca, Josa iguaque, Josa dianae, Josa siberia, Josa guayabal, Josa paipa, Josa cota, Josa sabogali, Josa denticulata, Josa junin, Josa bongara, Josa argentina, Josa tuberculata, Josa magalhaesi, Josa abeli, Josa laminaris, Josa huari, Josa peruviensis, Josa sierwaldae, Josa libertad, Josa ancash, Josa silvae, Josa cajamarca, Josa duembolaris, and Josa ecuadorensis. Based on the morphological diversity of the genus, particularly in sexual characters, we recognize eight species groups and test their monophyly using four molecular markers and a morphological dataset comprising 560 characters. Analyses under parsimony and maximum likelihood, including a total-evidence maximum-likelihood analysis of 324 terminals, support the monophyly of most proposed species groups, several of which were also recovered in the remaining analyses. Finally, we provide an overview of morphological diversity within Josa and discuss major patterns in its geographic distribution.