Biology of the phylum nematomorpha.

Compared with most animal phyla, the Nematomorpha, also known as hair worms, is a relatively understudied metazoan phylum. Although nematomorphs make up only 1 of 3 animal phyla specializing solely on a parasitic life style, little attention has been focused on this enigmatic group scientifically. The phylum contains two main groups. The nectonematids are parasites of marine invertebrates such as hermit crabs. The gordiids are parasites of terrestrial arthropods, such as mantids, beetles, and crickets. Members of both of these groups are free-living as adults in marine and freshwaters respectively. In recent years, large strides have been made to understand this group more fully. New information has come from collection efforts, new approaches in organismal biology, modern techniques in microscopy and molecular biology. This review will focus on the advances made in four main areas of research: (1) morphology, (2) taxonomy and systematics, (3) life cycle and ecology and (4) host behavioural alterations. Recent research focus on the structure of both nectonematids and gordiids has added new insights on the morphology of adult worms and juveniles. The nervous system of gordiids is now well described, including the documentation of sensory cells. In addition, the availability of material from the juvenile of several species of gordiids has made it possible to document the development of the parasitic stage. New collections and reinvestigations of museum specimens have allowed for a critical reevaluation of the validity of established genera and species. However, traditional taxonomic work on this group continues to be hampered by two impeding factors: first is the lack of species-specific characters; and second is the problem of intraspecific variation, which has likely led to the description of numerous synonyms. Modern molecular techniques have been used recently to support independently the broad relationships among gordiids. During the turn of the millennium, the study of the life cycle and general ecology of gordiids enjoyed a revival. The pivotal outcome of this research was the domestication of a common American gordiid species, Paragordius varius. This species was the first of this phylum to be laboratory-reared. Through this research, the life cycle of several distantly related gordiid species was investigated. Other work showed that gordiids persist in the environment in the cyst stage by moving through different hosts by paratenesis. These cysts have been shown to retain infectivity for up to a year. These factors have likely contributed to the finding that gordiid cysts are one of the most common metazoans in some aquatic environments. Finally, recent work has focused on elucidating the mechanism of how gordiids make the transition from terrestrially based definitive hosts to a free-living aquatic environment. It has been shown that hosts are manipulated by the parasites to enter water. Using this study system, and using histology and proteomic tools, the method of manipulation used by these parasites is being further investigated. This manipulation, and the reaction of the cricket to this manipulation, has been postulated to benefit both the parasite and the host. Although large strides have been made within the last 10 years in the understanding of nematomorphs, we make the case that a lot of basic information remains to be uncovered. Although seemingly a daunting task, the recent advances in information and techniques lay a solid foundation for the future study of this unique group of parasites.