Molecular systematics and historical biogeography of the Nocomis biguttatus species group (Teleostei: Cyprinidae): nuclear and mitochondrial introgression and a cryptic Ozark species.

The Nocomis biguttatus species group ranges widely across North America from the Red River in Oklahoma and Arkansas north to Minnesota and east-west from Wyoming to Ontario. The group includes three traditionally recognized allopatric species: the wide-ranging N. biguttatus and two geographically more restricted species, N. asper from the western Ozarks (Arkansas River system) and two disjunct locations in the Red River system, and N. effusus from the Green, Cumberland, and lower Tennessee rivers. Separate analyses of the mitochondrial cytb gene and two nuclear genes (S7 intron 1 and a portion of the gene for growth hormone, GH), each resolved a cryptic species previously treated as N. biguttatus from the southern Ozarks (White River). Relationships among the four species were unresolved because of conflicts between cytb and S7 and a lack of resolution for GH. A previously indicated N. biguttatus-N. effusus sister-relationship appears to reflect past hybridization and mtDNA capture by N. effusus. Nocomis biguttatus includes four primary cytb clades with unresolved inter-relationships. A Northern Ozarks-Great Plains-Upper Midwest Clade and an Ohio River-Eastern Great Lakes Clade presumably represent late Quaternary dispersal from glacial refugia in, respectively, the northern Ozarks and an unglaciated portion of the Ohio River system. Other clades include one from the Meramec River and a Black River-St. Francis River Clade. There was evidence in N. effusus for a phylogeographic break between the lower Tennessee River and the Green-Cumberland basins. Geographic structure is weak in N. asper, indicating relatively recent contact between now disjunct populations in the Arkansas and Red river basins. The Blue River population of N. asper appears to reflect late Pleistocene or Holocene hybridization and genetic swamping of a resident native population of N. biguttatus by an invading population of N. asper. This postulates past occurrence of N. biguttatus far south of its present range.

Rapid diversification in the North American minnow genus Nocomis.

Diversification rates among stream fishes are likely governed by geographical factors that impact connectivity among and between stream systems. Herein we investigate rates of diversification within species of River Chubs, Nocomis, a monophyletic group of stream fishes distributed throughout the drainage basins of the Mississippi River, Atlantic Slope and Gulf Coast. We used nucleotide sequences of two protein-coding nuclear genes (IRBP and rhodopsin) and one mitochondrial gene (cytochrome b) to generate a phylogenetic hypothesis of population-level relationships within and among species. Results of molecular analyses suggest that morphological and meristic treatments of Nocomis have underestimated species diversity within the group. Tree-based Bayesian methods and diversification statistics were implemented to model the rate of evolutionary change along lineages and estimate divergence dates among phylogroups. Significantly elevated rates of cladogenesis are observed among coastally distributed populations 2-3 million years before present. This period was dominated by dramatic sea level fluctuations that suggest a period of climatic instability. Climatic instability and other factors may have driven the burst of rapid diversification observed in Nocomis. Results generated in this study reinforce faunistic and geologic arguments for the hypothesized existence of extinct rivers, such as White's River and the Appalachian River.

Evolution of herbivory in a carnivorous clade of minnows (teleostei: cyprinidae): effects on gut size and digestive physiology.

We constructed a phylogeny for 10 minnow species (family Cyprinidae) previously revealed to be members of sister genera with different dietary affinities and used the phylogeny to examine whether the evolution of digestive tract size and physiology is correlated with the evolution of diet in these fishes. We studied a total of 11 taxa: four herbivorous species in the genus Campostoma and six largely carnivorous species in the genus Nocomis, including two populations of Nocomis leptocephalus, the carnivorous Chattahoochee River drainage population and the herbivorous Altamaha River drainage population. Thus, we were able to compare digestive tract size and physiology among sister genera (Campostoma and Nocomis) and among sister taxa (N. leptocephalus Chattahoochee and N. leptocephalus Altamaha) in dietary and phylogenetic contexts. The herbivorous taxa had longer digestive tracts and higher activity of the carbohydrases amylase and laminarinase in their guts, whereas the carnivorous species had higher chitinase activity. Phylogenetic independent-contrast analysis suggested that the evolution of amylase and chitinase activities was correlated with the evolution of diet in these species, whereas trypsin and lipase activities showed no pattern associated with diet or phylogenetic history. Concentrations of short-chain fatty acids were low in all taxa, indicating that these fishes rely largely on endogenous digestive mechanisms to subsist on their respective diets. Subtle differences in tooth shape were observed between species in the two genera. Overall, our results suggest that dietary specialization can be observed on the level of anatomy and physiology of the digestive tracts of fishes but that such differences are most appropriately viewed in comparisons of closely related species with different diets.