A comparative morphology of trichobothrial bases in araneoid spiders and its significance for the phylogeny and system of the superfamily Araneoidea (Arachnida, Araneae).

Bothrial morphology was studied by SEM in 137 araneoid genera representing all 22 currently recognized extant families and all 42 conventional subfamilies of the Araneoidea. The ancestral type in the superfamily Araneoidea is a 'hooded' bothrium with a single well-developed transverse ridge, dividing its proximal and distal plates ('-type'); the advanced type is a solid dome-like bothrium without vestiges of the ridge ('-type'); there are several intermediate types reflecting various pathways and stages of the ridge reduction (united here as '-type'). The parallel trends in bothrial evolution, recognized as continuous series from the ancestral type up to the advanced one through some intermediate stages, are distinguished in each of the seven main phylogenetic lineages of the superfamily: 'tetragnathoids', 'araneoids', 'cyatholipoids', and 'theridioids' possess a complete set of the three types, while 'malkariods', 'symphytognathoids'. and 'linyphioids' lack the advanced, dome-like type ('-type'). Only three taxa have been proposed earlier as the sister group of the superfamily Araneoidea: Nicodamoidea, Deinopoidea, and Leptonetoidea; morphology of bothria, as well as other cuticular microstructures, clearly supports the araneoid-nicodamoid relationship hypothesis, purely 'molecular' to date. Bothrial morphology provides the additional arguments for several taxonomic acts, e.g., for the reranking the Agnarsson's (2004) 'clade 35' (, , , and ) up to the Theonoeinae Simon, 1894, , and for the revalidation the micropholcommatid Butler, 1932, and zygiellid Wunderlich, 2004,