Morphological phylogeny of cobweb spiders and their relatives (Araneae, Araneoidea, Theridiidae)

This paper offers the first cladistic analysis of a wide selection of theridiid genera based on morphological data. The analysis treats 53 theridiid taxa representing 32 genera (Achaearanea, Anelosimus, Ameridion, Argyrodes, Ariamnes, Carniella, Cerocida, Chrysso, Coleosoma, Dipoena, Emertonella, Enoplognatha, Episinus, Euryopis, Faiditus, Kochiura, Latrodectus, Neospintharus, Nesticodes, Pholcomma, Phoroncidia, Rhomphaea, Robertus, Selkirkiella, Spintharus, Steatoda, Stemmops, Theridion, Theridula, Thymoites, Thwaitesia, Tidarren) and eight outgroup taxa representing the families Nesticidae (Eidmanella and Nesticus), Synotaxidae (Synotaxus, two species), Pimoidae (Pimoa), Linyphiidae (Linyphia), Tetragnathidae (Tetragnatha) and Araneidae (Argiope). The parsimony analysis of 242 morphological and behavioural characters found a single, most parsimonious tree. The monophyly of theridiids and their sister relationship with nesticids is strongly supported. The recent resurrection of Ariamnes and Rhomphaea from Argyrodes made the latter paraphyletic. However, Ariamnes and Rhomphaea are characterized by an array of characters, and Argyrodes still contains dramatically distinct clades for which names are available: Faiditus (removed from synonymy - RS) and Neospintharus (RS). These revalidations provide a classification with greater information content and utility. These three genera, along with Ariamnes, Rhomphaea and Spheropistha, comprise the subfamily Argyrodinae. The monophyly and composition of the subfamilies Hadrotarsinae, Spintharinae, Pholcommatinae, Latrodectinae and Theridiinae are discussed. Theridion is paraphyletic and in need of revision. Anelosimus as currently circumscribed is paraphyletic, a problem resolved by revalidating Selkirkiella (RS) and Kochiura (RS). Numerous new combinations are established. The results suggest the monophyletic origin of both kleptoparasitism and araneophagy in the lineage leading to Argyrodinae, negating hypotheses that either arose from the other. Sociality evolved multiple times within the family, accounting for as much as one fourth of the origins of social behaviour among all spiders. No losses of sociality are implied. The hypothesis of maternal care as the pathway to sociality receives support. Evolution of theridiid webs is complex, with multiple modifications and loss of the basic theridiid cobweb. (C) 2004 The Linnean Society of London.