Life-stage-specific physiology defines invasion extent of a riverine fish

Many ecologists have called for mechanism-based investigations to identify the underlying controls on species distributions. Understanding these controls can be especially useful to construct robust predictions of how a species range may change in response to climate change or the extent to which a non-native species may spread in novel environments. Here, we link spatially intensive observations with mechanistic models to illustrate how physiology determines the upstream extent of the aquatic ectotherm smallmouth bass (Micropterus dolomieu) in two headwater rivers. Our results demonstrate that as temperatures become increasingly cold across a downstream to upstream gradient, food consumption in age 0 bass becomes increasingly constrained, and as a result, these fish become growth limited. Sufficient first summer growth of age 0 bass is essential for overwinter survival because young bass must persist from energy reserves accumulated during the summer, and those reserves are determined by body size. Our field data reveal the upstream extent of adult bass reproduction corresponds to a point in the downstream/upstream gradient where cold temperatures impair growth opportunities in young bass. This pattern was repeated in both study streams and explained why bass positioned nests twice as far upstream in the warm compared to the cold stream in the same basin. Placement of spawning nests by adult bass is likely subject to strong evolutionary selection in temperate systems: if bass spawn too far upstream, their young are unlikely to grow large enough to survive the winter. Consumption and growth in older bass (age 3-4) was far less sensitive to temperature. Based on these data, we suggest that temperature-sensitive age 0 bass constrain the upstream distribution limits of bass within temperate streams. In this study, we investigated how temperature-dependent physiology changed through the life history of a species and, in doing so, identified a climate-sensitive life-history stage that likely sets the distributional limits of all other life-history stages. We anticipate the framework developed here could be employed to identify how similar stage-specific environmental sensitivity determines distribution in many other ectothermic species.