A framework for detecting natural selection on traits above the species level

To what extent can natural selection act on groupings above the species level? Despite extensive theoretical discussion and growing practical concerns over increased rates of global ecological turnover, the question has largely evaded empirical resolution. A flexible and robust hypothesis-testing framework for detecting the phenomenon could facilitate significant progress in resolving this issue. We introduce a permutation-based approach, implemented in the R package perspectev, which provides an explicit test of whether empirical patterns of correlation between upper level trait values and survivorship are reducible to correlations manifested at lower levels. The package is applicable to virtually any nested set of upper- and lower level groupings, a wide variety of upper level traits, and both historical and contemporary occurrence data. We apply this approach to five paleontological data sets that represent different magnitudes of extinction and differ in taxonomic breadth, geologic...