Phylogenetic placement and microthrix pattern of Paranybelinia otobothrioides Dollfus, 1966 (Trypanorhyncha) from krill Nyctiphanes simplex Hansen, 1911

Plerocerci of the monotypic Paranybelinia otobothrioides were found parasitizing the subtropical neritic krill Nyctiphanes simplex in the Gulf of California, Mexico. The plerocerci were recovered from two microhabitats of the intermediate host, typically embedded inside the digestive gland (hepatopancreas) or rarely in the hemocoel. The morphology of the simple, single-layered blastocyst surrounding the entire scolex is unique within the Trypanorhyncha by having four large funnel-like pori or openings possibly with feeding and/or excretory function. One of the openings is located anteriorly and three at the posterior end. Scolex surface ultrastructure shows hamulate and lineate spinitriches covering the bothrial surface, capilliform filitriches at the anterior scolex end and on the scolex peduncle, and short papilliform filitriches on the long appendix. This pattern resembles that of species of the Tentaculariidae; but differs in that the hamulate spinitriches, which appear lineate at the bothrial margins, densely cover the entire distal bothrial surface. Tegumental grooves are present on the posterior bothrial margin, lacking spinitriches. Paranybelinia otobothrioides and Pseudonybelinia odontacantha share the following unique combination of characters: two bothria with free lateral and posterior bothrial margins, homeoacanthous homeomorphous armature, tegumental grooves, the distribution of the hamulate spinitriches, and the absence of prebulbar organs. Both genera infect euphausiids as intermediate hosts. Sequence data of the partial ssrDNA gene place Pa. otobothrioides sister to the family Tentaculariidae, and the Kimura two-parameters (K2P) distance between Pa. otobothrioides and species of the family Tentaculariidae ranged from 0.027 to 0.039 (44-62 nucleotide differences). These data suggest both species be recognized in a family, the Paranybeliniidae, distinct from, albeit as sister taxon to, the Tentaculariidae. High prevalence of infection (<14%) and ontogenetic changes of Pa. otobothrioides support N. simplex as a required intermediate host and suggest a zooplanktophagous elasmobranch as final host in the Gulf of California.