Landscape Evolution as a Diversification Driver in Freshwater Fishes

The exceptional concentration of vertebrate diversity in continental freshwaters has been termed the “freshwater fish paradox,” with > 15,000 fish species representing more than 20% of all vertebrate species compressed into tiny fractions of the Earth’s land surface area (<0.5%) or total aquatic habitat volume (<0.001%). This study asks if the fish species richness of the world’s river basins is explainable in terms of river captures using topographic metrics as proxies. The River Capture Hypothesis posits that drainage-network rearrangements have accelerated biotic diversification through their combined effects on dispersal, speciation, and extinction. Yet rates of river capture are poorly constrained at the basin scale worldwide. Here we assess correlations between fish species density (data for 14,953 obligate freshwater fish species) and basin-wide metrics of landscape evolution (data for 3,119 river basins), including: topography (elevation, average relief, slope, drainage area) and climate (average rainfall and air temperature). We assess the results in the context of both static landscapes (e.g., species-area and habitat heterogeneity relationships) and transient landscapes (e.g., river capture, tectonic activity, landscape disequilibrium). We also relax assumptions of functional neutrality of basins (tropical vs. extratropical, tectonically stable vs. active terrains). We found a disproportionate number of freshwater species in large, lowland river basins of tropical South America, Africa, and Southeast Asia, under predictable conditions of large geographic area, tropical climate, low topographic relief, and high habitat volume (i.e., high rainfall rates). However, our results show that these conditions are only necessary, but not fully sufficient, to explain the basins with the highest diversity. Basins with highest diversity are all located on tectonically stable regions, places where river capture is predicted to be most conducive to the formation of high fish species richness over evolutionary timescales. Our results are consistent with predictions of several landscape evolution models, including the River Capture Hypothesis, Mega Capture Hypothesis, and Intermediate Capture Rate Hypothesis, and support conclusions of numerical modeling studies indicating landscape transience as a mechanistic driver of net diversification in riverine and riparian organisms with widespread continental distributions.

Transmissivity and groundwater flow exert a strong influence on drainage density

The extent to which groundwater flow affects drainage density and erosion has long been debated but is still uncertain. Here, I present a new hybrid analytical and numerical model that simulates groundwater flow, overland flow, hillslope erosion and stream incision. The model is used to explore the relation between groundwater flow and the incision and persistence of streams for a set of parameters that represent average humid climate conditions. The results show that transmissivity and groundwater flow exert a strong control on drainage density. High transmissivity results in low drainage density and high incision rates (and vice versa), with drainage density varying roughly linearly with transmissivity. The model evolves by a process that is defined here as groundwater capture, whereby streams with a higher rate of incision draw the water table below neighbouring streams, which subsequently run dry and stop incising. This process is less efficient in models with low transmissivity due to the association between low transmissivity and high water table gradients. A comparison of different parameters shows that drainage density is most sensitive to transmissivity, followed by parameters that govern the initial slope and base level. The results agree with field data that show a negative correlation between transmissivity and drainage density. These results imply that permeability and transmissivity exert a strong control on drainage density, stream incision and landscape evolution. Thus, models of landscape evolution may need to explicitly include groundwater flow.

Supplemental Material: Tectonic controls on Quaternary landscape evolution in the Ventura basin, southern California, USA, quantified using cosmogenic isotopes and topographic analyses

Supplemental text, 6 figures, and 10 data tables to support the methods and results from the main manuscript.

Supplemental Material: Tectonic controls on Quaternary landscape evolution in the Ventura basin, southern California, USA, quantified using cosmogenic isotopes and topographic analyses

Supplemental text, 6 figures, and 10 data tables to support the methods and results from the main manuscript.