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, geological timing and geographic extent. Using simulations, we demonstrate that this method is a robust means of detecting irreducibility in the relationship between upper level traits and survivorship, and outline circumstances in which the method is less effective. We also find evidence consistent with previous findings of selection above the species level for geographic range size in North American K-Pg molluscs and show that this phenomenon was evident for the same molluscan genera globally. Ultimately, we conclude that at certain points in history, some higher level taxonomic groups have survived differentially with respect to geographic range size in a manner that is not explained by the same trait at the species level, and we show that evidence for this phenomenon varies across taxa and extinction events. We release our method as a flexible and easy-to-use R package that will allow others to help determine the relative frequency of this macroevolutionary phenomenon, both in the fossil record and in estimates of contemporary extinction risk.
Hoehn, KB; Harnik, PG; Roth, VL
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