Demise of Marimermithida refines primary routes of transition to parasitism in roundworms

Nematodes (roundworms) are ubiquitous animals commonly dominating in ecological communities and networks, with many parasites and pathogen vectors of great economic and medical significance. Nematode parasites are remarkably diverse in life strategies and adaptations at a great range of hosts and dimension scales, from whales to protozoan cells. Their life history is intricate and requires understanding to study the genomic, structural and ecological bases of successful transitions to parasitism. Based on analyses of rDNA for a representative sampling of host-associated and free-living groups, we dismiss the last higher-rank nematode taxon uniting solely parasitic forms (Marimermithida) and show that primarily marine parasitism emerged independently and repeatedly within only few free-living lineages. We re-evaluate the significance of some traditionally important phenotypic characters and report the phenomenon of dramatic adaptation to parasitism on very short evolutionary timescales. A cross-phylum character interpretation vindicates that non-intestinal (in-tissue or cavitary) host capture was likely a primary route of transition to truly exploitive parasitism (vs. intestinal commensalism) in roundworms, and extant nematode parasitoids (larval parasites) infesting the host body cavity or internal organs realise this primary lifestyle. Parasitism may have evolved in nematodes as part of innate pre-adaptations to crossing environmental boarders, and such transitions have been accomplished multiple times successfully in the phylum history.