Molecular systematics of new world gopher, bull, and pinesnakes (Pituophis: Colubridae), a transcontinental species complex

Pituophis melanoleucus (gopher, bull, and pine-snakes) is among the most widely distributed polytypic species complexes in North America, with most authors recognizing from a single transcontinental species (the melanoleucus complex, composed of 15 subspecies) to four (monotypic and polytypic) species. We used mitochondrial gene sequences from the two middle American species, P. deppei and P. lineaticollis, and from 13 subspecies from most of the range of the melanoleucus complex to test various phylogenetic hypotheses for Pituophis. Maximum parsimony and maximum likelihood methods identified the same major clades within Pituophis and indicated that two segments of the melanoleucus complex, the lodingi-melanoleucus-mugitus eastern pinesnake clade and the affinis-annectens-bimaris-catenifer-deserticola-sayi-ruthveni vertebralis clade from central and western United States and northern Mexico, represent divergent, allopatric lineages with no known intergradation zone. We recognize each of these two groupings as a different species. Our data also indicate that some ruthveni are more closely related to sayi than to other ruthveni. Nonetheless, ruthveni is an allopatric taxon diagnosable from its closest relatives by a combination of morphometric characters, and because it is likely that at least some of these traits are independent and genetically inherited, we interpret this as evidence that ruthveni has attained the status of independent evolutionary lineage, despite the fact that it retains strong genetic affinities with sayi. The endemic Baja Californian gopher snakes (bimaris and vertebralis) are considered by some taxonomists as a different species, P. vertebralis, but we discovered that these serpents belong to two different clades and hence we do not agree with the recognition of P. vertebralis as presently defined. In summary, we believe that three distinct species are included in the melanoleucus complex, Pituophis melanoleucus (sensu stricto), P. catenifer, and P. ruthveni, and that their recognition better represents the evolutionary diversity within this species complex, (C) 2000 Academic Press.