Morphological Selection and the Evaluation of Potential Tradeoffs Between Escape from Predators and the Climbing of Waterfalls in the Hawaiian Stream Goby Sicyopterus stimpsoni

Abstract

Environmental pressures may vary over the geographic range of a species, exposing subpopulations to divergent functional demands. How does exposure to competing demands shape the morphology of species and influence the divergence of populations? We explored these questions by performing selection experiments on juveniles of the Hawaiian goby Sicyopterus stimpsoni, an amphidromous fish that exhibits morphological differences across portions of its geographic range where different environmental pressures predominate. Juvenile S. stimpsoni face two primary and potentially opposing selective pressures on body shape as they return from the ocean to freshwater streams on islands: (1) avoiding predators in the lower reaches of a stream; and (2) climbing waterfalls to reach the habitats occupied by adults. These pressures differ in importance across the Hawaiian Islands. On the youngest island, Hawai'i, waterfalls are close to shore, thereby minimizing exposure to predators and placing a premium on climbing performance. In contrast, on the oldest major island, Kaua'i, waterfalls have eroded further inland, lengthening the exposure of juveniles to predators before migrating juveniles begin climbing. Both juvenile and adult fish show differences in body shape between these islands that would be predicted to improve evasion of predators by fish from Kaua'i (e.g., taller bodies that improve thrust) and climbing performance for fish from Hawai'i (e.g., narrower bodies that reduce drag), matching the prevailing environmental demand on each island. To evaluate how competing selection pressures and functional tradeoffs contribute to the divergence in body shape observed in S. stimpsoni, we compared selection imposed on juvenile body shape by (1) predation by the native fish Eleotris sandwicensis versus (2) climbing an artificial waterfall (∼100 body lengths). Some variables showed opposing patterns of selection that matched predictions: for example, survivors of predation had lower fineness ratios than did control fish (i.e., greater body depth for a given length), whereas successful climbers had higher fineness ratios (reducing drag) than did fish that failed. However, most morphological variables showed significant selection in only one treatment rather than opposing selection across both. Thus, functional tradeoffs between evasion of predators and minimizing drag during climbing might influence divergence in body shape across subpopulations, but even when selection is an important contributing mechanism, directly opposite patterns of selection across environmental demands are not required to generate morphological divergence.