Parasitic nematodes of the genus Trichinella cause significant food-borne illness and occupy a unique evolutionary position at the base of the phylum Nematoda, unlike the free-living nematode Caenorhabditis elegans. Although the forthcoming genome sequence of Trichinella spiralis can provide invaluable comparative information about nematode biology, a basic framework for understanding the history of the genus Trichinella is needed to maximize its utility. We therefore developed the first robust and comprehensive analysis of the phylogeny and biogeographic history of Trichinella using the variation in three genes (nuclear small-subunit rDNA, and second internal transcribed spacer, mitochondrial large-subunit rDNA, and cytochrome oxidase I DNA) from all 11 recognized taxa. We conclude that (i) although Trichinellidae may have diverged from their closest extant relatives during the Paleozoic, all contemporary species of Trichinella diversified within the last 20 million years through geographic colonization and pervasive host switching among foraging guilds of obligate carnivores; (ii) mammalian carnivores disseminated encapsulated forms from Eurasia to Africa during the late Miocene and Pliocene, and to the Nearctic across the Bering Land Bridge during the Pliocene and Pleistocene, when crown species ultimately diversified; (iii) the greatest risk to human health is posed by those species retaining an ancestral capacity to parasitize a wide range of hosts; and (iv) early hominids may have first acquired Trichinella on the African savannah several million years before swine domestication as their diets shifted from herbivory to facultative carnivory.biogeography ͉ mitochondrial DNA ͉ phylogeny ͉ ribosomal DNA E xceptional biological diversity among nematodes is exemplified by certain attributes of the parasite Trichinella spiralis. Organization of its mitochondrial genome more closely resembles that of coelomate metazoans than that of its presumed closest relatives, the secernentean nematodes (1). In addition, T. spiralis (Dorylaimia) shares a similar proportion (45%) of its ESTs with the nematode Caenorhabditis elegans (Rhabditina) as it does with the fruit fly Drosophila melanogaster (Arthropoda: Drosophilidae) (2). Thus, many ESTs common to T. spiralis and C. elegans are not necessarily specific to nematodes but may be conserved among diverse taxonomic groups of invertebrates. C. elegans is often thought of as a prototypical nematode because of its acceptance as a model for studying biological processes; however, genomic variation among nematodes is extensive and commensurate with their phylogenetic and ecological diversity. Therefore, the forthcoming genome sequence of T. spiralis will contribute substantially to our understanding of nematode biology and the origins of parasitism.In 1998, Blaxter et al. (3) used genetic data to delineate the phylum Nematoda into supertaxa consisting of five clades. Trichinella, a parasite of vertebrates, occupies a basal lineage (clade I) consisting of free-living Mononchida, ...