A Phylogenetic and Biogeographic Analysis of the Genera of Spirorchinae (Digenea: Spirorchidae) Parasitic in Freshwater Turtles

Abstract

Cladistic analysis of the freshwater genera of Spirorchinae (Schistosomatoidea: Spirorchidae sensu Yamaguti, 1971) plus Haematotrema Stunkard, 1923, and Aphanospirorchis Platt, 1990, was completed. The Spirorchinae were considered monophyletic based on synapomorphies of the esophagus. Three lineages, Spirhapalum (Europe/Asia), Plasmiorchis+Hemiorchis (India), and Spirorchis + Henotosoma + Haematotrema + Aphanospirorchis (North America), were identified. Nelsen consensus analysis was used as the basis for recognizing 3 valid monophyletic genera: Spirhapalum, Plasmiorchis, and Spirorchis. Hapalotrematinae sensu Smith, 1972 (e.g., Hapalorhynchus/Coeuritrema), is considered the most plesiomorphic group of spirorchids. Freshwater representatives of the hapalotrematines have been reported from 7 of 12 extant turtle families, including the relatively primitive Pelomedusidae (Pleurodira) and exhibit a worldwide distribution. It is hypothesized that this group arose in the early Triassic period, prior to the breakup of Pangea. Thus, it represents a primitive lineage that was present during the diversification of turtle lineages in the mid-Mesozoic era. Spirorchinae arose later (late Cretaceous period) as a Laurasian component parasitic in the more recent pond turtles (Emydidae + Bataguridae). Species of Spirhapalum retained a relatively plesiomorphic distribution, and they are found in emydids (Europe) and batagurids (Asia). Species of Spirorchis arose and diversified with North America emydids following the separation of North America and Europe in the late Cretaceous or early Tertiary periods. Species of Plasmiorchis are hypothesized to be derived from Asian ancestors that accompanied the colonization of India by Asian batagurids during the early Tertiary period. The presence of Spirorchis species in snapping turtles (Chelydridae/North America) and of Plasmiorchis species in Indian soft-shelled turtle (Trionychidae) are considered independent colonization events.