Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones).

Among the least studied harvestmen are the members of the family Caddidae sensu Shear, 1975, a group of Opiliones with massive eyes and the putative sister group of the remaining Eupnoi. Caddids were originally described as two families, Caddidae and Acropsopilionidae, but these are currently treated as subfamilies of Caddidae. These minute arachnids are rarely collected and present some interesting biogeographical patterns, including a disjunct distribution between East Asia and eastern North America, and some of the few cases of trans-Pacific genera in southern hemisphere Opiliones. We therefore obtained samples from most of the landmasses inhabited by Caddidae and undertook a phylogenetic study using nuclear and mitochondrial genes for as many samples as possible. Our results, based on a broad taxonomic sampling, surprisingly showed polyphyly of Caddidae, with the genus Caddo forming the sister group of the remaining Eupnoi, whereas the southern hemisphere genera, many of which were originally placed in Acropsopilionidae, within Dyspnoi, formed the sister clade of the remaining Dyspnoi. In addition, the more recently described genus Hesperopilio, from Western Australia and Chile, was unrelated to either Caddidae or Acropsopilionidae, despite having the supposedly diagnostic large ocularium, and instead appeared deeply nested within the Eupnoi superfamily Phalangioidea. Our results are robust to analytical treatment and to homology scheme (dynamic vs. static notions of homology), resulting in a new phylogenetic proposal for Eupnoi and Dyspnoi. Ancestral state reconstruction suggests that the ancestral Palpatores was probably a tiny harvestman with an enlarged ocularium and glandular palpal setae in its enlarged and armed palps. We take the following taxonomic actions: Acropsopilionidae is removed from synonymy under Caddidae and its family status and membership in Dyspnoi are restored. Hesperopilio Shear, 1996 is removed from Caddoidea/Caddidae and transferred to Phalangioidea, but it is not assigned to any family.