Intraspecific Phylogeography of the Stonecat Madtom, Noturus flavus
Two morphologically distinct forms of the Stonecat Madtom, Noturus flavus, are found in lowland and Eastern Highland drainages of North America. Evolutionary diversity of aquatic organisms in these regions has been attributed to divergence during the Pliocene and Pleistocene epochs followed by dispersal into previously glaciated regions. We surveyed mitochondrial DNA sequence variation across the range of this widely distributed species to determine the evolutionary relationship between the two forms, and to investigate the possible roles of pre-Pleistocene vicariance and post-Pleistocene dispersal on phylogeographic patterns within the species. Analysis revealed four distinct clades, two within the lowland form and two within the highland form, with no geographic overlap. A deep phylogenetic split between the two forms implies historical fragmentation during the Pliocene. Shallow divergences and limited geographic structuring in the lowland form imply recent fragmentation in two glacial refugia during the Pleistocene, followed by dispersal across much of the modern range of the species. The highland form is common to the Cumberland and Tennessee rivers and clades correspond to each drainage, apparently owing to their separation during the Pleistocene. Phylogeographic patterns echo those of other eastern North American stream fishes, with the persistence of deep phylogenetic divergence predicted by the pre-Pleistocene Vicariance Hypothesis and widespread distribution of closely related haplotypes in glaciated regions consistent with the Leading Edge Model of population expansion.Abstract

Collecting localities for Noturus flavus. Numbers correspond to Table 1. Inset map shows southern limit of Pleistocene glacial advance and limit of Driftless Region (compound line) and Central Highlands regions (shaded). (A) Eastern Highlands, (B) Ozark Highlands, (C) Ouachita Highlands.

Minimum spanning network of Noturus flavus mtDNA control region sequence haplotypes. Haplotypes are designated by letters and empty circles denote hypothetical intermediates. Each line connecting individual haplotypes represents one mutational step. Numbers on lines denote the minimum number of mutational steps between clades.

Neighbor-joining distance tree for the mtDNA control region and tRNAPhe gene of 20 haplotypes of Noturus flavus (A–T) and outgroup taxa N. insignis and N. lachneri. Relationships were estimated using pairwise p-distances. Bootstrap values above 60% for the neighbor joining analysis are located above the nodes while those for the maximum parsimony analysis are below the nodes. Clade names correspond to those of the minimum spanning network. Group I corresponds to clades collected in lowland rivers and Group II corresponds to clades collected in rivers of the Eastern Highlands.

Frequency distributions of pairwise nucleotide differences between haplotypes of Noturus flavus. (A) All haplotypes, (B) Group I including haplotypes A through K, (C) Group II including haplotypes L through T. A solid line represents observed data, and dashed lines represent 95% confidence intervals. Values for the raggedness statistic and the sum of squared deviations statistic are shown with P values.
Contributor Notes
Associate Editor: J. M. Quattro.