Historical biogeography among species of Varestrongylus lungworms (Nematoda: Protostrongylidae) in ungulates: episodic expansion and host colonization linking Eurasia and North America.

Varestrongylus lungworms (Nematoda: Protostrongylidae) include 10 nominal species that parasitize wild and domesticated artiodactyles. Eight species are endemic to the western Palearctic and Eurasia, whereas two are limited in distribution to the Nearctic. Complex host associations, primarily among Cervidae and Bovidae (Caprinae), and biogeography were explored based on direct comparisons of parasite and host phylogenies to reveal the historical development of this fauna. Diversification among Varestrongylus species has an intricate history extending over the Pliocene and Quaternary involving episodic processes for geographic and host colonization: (1) Varestrongylus has origins in Eurasia with independent expansion events into bordering ecozones; (2) cervids are ancestral hosts; (3) the caprine-associated V. pneumonicus is basal and a result of an independent host colonization event; (4) secondary diversification, linked to sequential and independent host colonization events, occurred within cervids (V. sagittatus + V. tuvae; V. alpenae; and V. capreoli, V. alces + V. eleguneniensis); (5) at least two additional host colonization events into caprines occurred, followed or not by diversification (V. qinghaiensis + V. longispiculatus; V. capricola, respectively); (6) two independent events of geographic expansion into North America from Eurasia with cervids in the late Pliocene and early Pleistocene are postulated (V. alpenae, V. eleguneniensis). Comparisons based on phylogenetic hypotheses derived from comparative morphology and molecular inference for these nematodes are consistent with the postulated history for coevolutionary and biogeographic history. Episodes of geographic and host colonization, often in relation to rapid shifts in climate and habitat perturbation, have dominated the history of diversification of Varestrongylus.