Erosion through ancient geologic structures as a mechanism for freshwater fish speciation in a post-orogenic mountain range

The legacy of tectonic deformation affects geomorphic and biological dynamics, even in post-orogenic mountain ranges. As ancient geologic structures originally created through tectonic deformation are exhumed through erosion, rocks with different chemical and physical properties are exposed at the surface of the landscape. We propose that this process not only influences landscape dynamics but is also a mechanism for speciation in freshwater fish.  As rivers erode through layers of different kinds of rock, the spatial distribution of rocks at the surface of the landscape changes. For fish with habitat specificity linked to rock type, erosion can progressively expose either favorable or unfavorable rock types, creating either barriers to or corridors for dispersal. The underlying structural geology will dictate which of those scenarios occurs. We present two case-studies that illustrate each scenario from the southeast United States, a freshwater biodiversity hotspot. First, we show that populations of the Greenfin Darter (Nothonotus chlorobranchius) are genetically isolated within tributaries flowing over the metamorphic rocks making up the thrust sheets of the Blue Ridge geologic province. In contrast, they are not found in rivers flowing over sedimentary rock of the Valley and Ridge. We show that over time, more sedimentary rock has been exposed, which has progressively isolated N. chlorobranchius populations from one another. In this case, river incision is introducing more barriers (sedimentary rock) into the landscape, leading to lineage diversification (i.e., speciation). In the second case-study, we explore the diversification of the Vermilion Darter complex that includes the federally endangered Vermilion Darter (Etheostoma chermocki) and the closely related Warrior Darter (E. bellator). Unique lineages of this species complex are restricted to tributaries flowing over carbonate rocks in the Black Warrior River. In contrast to the N. chlorobranchius case-study, here river incision is progressively expanding habitat by exposing more carbonate rock, driving dispersal-mediated allopatric speciation within the Vermilion Darter complex. Our results suggest that in bedrock-dominated rivers found throughout much of the Appalachian Mountains, erosion through ancient geologic structures can drive the diversification of freshwater fish, highlighting links between tectonic deformation, surface processes, and biological evolution in an ancient mountain range.