This paper reports the results of a 25 year study on an obligate quadritrophic system involving a midge, Cricotopus nostocicola (Diptera: Chironomidae), its cyanobacterial food source, Nostoc parmeliodes (Nostocales), a new species of nematode parasite of the midge, Gastromermis anisotis sp. n. .Mermithidae), and a virus infecting the nematode. Occurring in a mountain stream in California, the components of this system are adapted for survival at temperatures just above freezing (4-8 ± C). Developmental studies of the nematode include mating, oviposition, egg hatch, host penetration, parasitic development, free-living habits, spermatogenesis, platelet structure, and virus infection. Rates of parasitism varied between 5-42% over the 25 year period. The nematode was considered to be the most signi cant biological control agent of Cricotopus populations in the stream.The present study, which has been conducted over a 25 year period in a mountain stream in northern California, provides a detailed account of a new species of Mermithidae (Nematoda) that is one component of a unique, obligate quadritrophic system involving Nostoc thalli, herbivorous midges, a parasitoid nematode and an infectious virus. The chironomid/Nostoc/mermithid association was rst noted by Brock (1960) who studied the relationship between the midge and its Nostoc food source, concluding that it represented a case of mutualism. We describe the mermithid nematode, provide new information on its biology and developmentand discuss the system as a unique quadritrophic food web involving three levels of parasitism.