Integrating the metapopulation and habitat paradigms for understanding broad-scale declines of species

Caughley (1994) argued that researchers working on threatened populations tended to follow the "small population paradigm" or the "declining population paradigm," and that greater integration of these paradigms was needed. Here I suggest that two related paradigms exist at the broader spatial scale, namely the metapopulation paradigm and habitat paradigm, and that these two paradigms also need to be integrated if we are to provide sound management advice. This integration is not trivial, and I outline five problems that need to be addressed: (1) habitat variables may not measure habitat quality, so site-specific data on vital rates are needed to resolve the effects of habitat quality and metapopulation dynamics; (2) measurements of vital rates may be confounded by movements, (3) vital rates may be density dependent, (4) vital rates may be affected by genotype; and (5) vital rates cannot be measured in unoccupied patches. I reviewed papers published in Conservation Biology from 1994 to 2003 and found 41 studies that analyzed data from 10 or more sites to understand the factors limiting species' distributions. Five of the analyses presented were purely within the metapopulation paradigm, 14 were purely within the habitat paradigm, 17 involved elements of both paradigms, and 7 were theoretically ambiguous (2 papers presented 2 distinct analyses and were counted twice). This suggests that many researchers appreciate the need to integrate the paradigms. Only one study, however, used data on vital rates to resolve the effects of habitat quality and metapopulation dynamics (problem 1), and this study did not address problems 2-5. I conclude that more intensive research incorporating site-specific data on vital rates and movement is needed to complement the numerous analyses of distributional data being produced.