Relationship of habitat variability to population size in a stream fish.

The relationship between habitat variability and population size in fragmented habitats is poorly understood, yet might have important evolutionary consequences. For instance, fragmentation could (1) shift habitat characteristics, and by extension, selective regimes, in a consistent direction as populations and the fragments they occupy are reduced in size (directional hypothesis): or (2) increase variability in habitats among similarly sized populations as fragment size decreases (variable hypothesis). We investigated these alternatives based on multiyear habitat, demographic, and genetic data from 19 fragmented populations of a stream fish varying in census size (N) and effective number of breeders (N(b)). Mean habitat parameters were significantly related to N and N(b), but the forms of the relationships varied, and there was no evidence of consistent directional differences in habitat parameters from small to large population size. Small populations exhibited a wider range of variances in habitat parameters than large populations, and to a lesser extent, small populations also had greater variability in mean habitat parameters, possibly signaling more diverse selective regimes. These results suggest that many different environments are associated with small population size in nature, counter to the frequently cited assumption that small populations tend to occur only in marginal environments. In addition to well-documented demographic and genetic stochasticity operating within small populations, our work raises the possibility that small populations exhibit more variable and potentially less predictable evolutionary responses to future environmental change.