Abstract

Survivorship biases can generate remarkable apparent rate heterogeneities through time in otherwise homogeneous birth‐death models of phylogenies. They are a potential explanation for many striking patterns seen in the fossil record and molecular phylogenies. One such bias is the “push of the past”: clades that survived a substantial length of time are likely to have experienced a high rate of early diversification. This creates the illusion of a secular rate slow‐down through time that is, rather, a reversion to the mean. An extra effect increasing early rates of lineage generation is also seen in large clades. These biases are important but relatively neglected influences on many aspects of diversification patterns in the fossil record and elsewhere, such as diversification spikes after mass extinctions and at the origins of clades; they also influence rates of fossilization, changes in rates of phenotypic evolution and even molecular clocks. These inevitable features of surviving and/or large clades should thus not be generalized to the diversification process as a whole without additional study of small and extinct clades, and raise questions about many of the traditional explanations of the patterns seen in the fossil record.

Highlights

  • How do the effects we outline here intersect with them? We have shown above the expected sizes of both the push of the past” (POTPa) and large clade effect (LCE), which are themselves rate heterogeneities that arise from homogeneous models when conditioned on either/or survival and clade size

  • We have explored the patterns of diversification that can be generated by a retrospective view of a purely homogeneous model of diversification

  • Patterns of diversification through time have been much discussed in the literature (e.g., Hopkins and Smith 2015), with a common pattern being seen that diversification rates are high at the beginning of major evolutionary radiations, in both raw diversity counts and lineages through time plots

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Summary

Mathematical Analysis

We extend on the approach of Nee et al (1994b) by explicitly conditioning the birth-death model on the number of extant species in the crown group, and considering the full distribution of clade abundances over time rather than just the central expectation. Slowdowns seen in rates of diversification that begin with a wide range of values and quickly decline (largely being over by the time of the establishment of the crown group), with reconstructed rates in the stem lineage being significantly higher than in the generated plesions (i.e., extinct branches), are attributable to the POTPa. For the case of the birds discussed below, one would expect (in the fossil record) an observed initial diversification rate of about 1.25, that is about 20 times faster than the background rate.

The Push of the Past and the Fossil Record
Mass Extinctions
Remaining species
Rate Heterogeneity
Summary
Supporting Information
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