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Simpson, C.

Publications and source records attributed to Simpson, C..

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The measurement of species selection on evolving characters

Many processes can contribute to macroevolutionary change. This fact is the source of the wide variety of macroevolutionary change across time and taxa as well as the bane of pale-obiological research trying to understand how macroevolution works. Here, I present a general framework for understanding the variety of macroevolutionary phenomena. Based on Prices theorem, this framework provides a simple quantitative means to understand (1) the macroevolutionary processes that are possible and (2) the way those processes interact with each other. The major qualitative features of macroevolution depend first on the number of processes that co-occur and then on the magnitudes and evolutionary directions of those processes. Species selection, the major macroevolutionary process, consists of patterns of differential rates of speciation and extinction. Its macroevolutionary efficacy depends on the presences of sufficient microevolutionary change. Conversely, microevolutionary change is limited in power by the independent evolution of species, and species selection acting across populations of species can amplify or suppress microevolution. Non-trends may result if species selection sufficiently neutralizes microevolution and may yield stable macroevolutionary patterns over many millions of years.

paleontology

The case for species selection

The mere existence of speciation and extinction make macro-evolutionary processes possible. Speciation and extinction introduce discontinuities in the microevolutionary change within lineages by initiating, disrupting, and terminating the continuity of species lineages. Within a clade, speciation and extinction become potent means of macroevolution in and of themselves. This process, termed species selection, is a macroevolutionary analogue of natural selection, with species playing an analogous part akin to that played by organisms in microevolution. That said, it has proven difficult to think about levels of selection. The concept of species sorting was introduced to help our thinking on this issue by identifying two aspects inherent in hierarchical systems can confuse our attempts to understand them: uncertainty in the level that selection acts and uncertainty about if the pattern of selection is in fact caused at all. Thanks to insights from evolutionary transitions in individuality, we now know more about how to identify the level of selection and how to parse the causal structure in hierarchical evolutionary circumstances. We know that if the fitness of organisms causes the fitness of more inclusive species then they must covary. However, there is no evidence of such a covariance between fitnesses at these two levels. This covariance is just not observed; neither between cells and organisms nor between organisms and species. Rather, speciation and extinction rates appear to be completely divorced from organismal fitness. With this insight, the concept of species sorting shrinks so that it only covers the two processes of species selection and drift. I argue that we are better off focusing on understanding the processes of species selection and drift and that there is therefore no further need for the concept of species sorting.

paleontology