Density-Dependent Cladogenesis in Birds

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Abstract
A characteristic signature of adaptive radiation is a slowing of the rate of speciation toward the present. On the basis of molecular phylogenies, studies of single clades have frequently found evidence for a slowdown in diversification rate and have interpreted this as evidence for density dependent speciation. However, we demonstrated via simulation that large clades are expected to show stronger slowdowns than small clades, even if the probability of speciation and extinction remains constant through time. This is a consequence of exponential growth: clades, which, by chance, diversify at above the average rate early in their history, will tend to be large. They will also tend to regress back to the average diversification rate later on, and therefore show a slowdown. We conducted a meta-analysis of the distribution of speciation events through time, focusing on sequence-based phylogenies for 45 clades of birds. Thirteen of the 23 clades (57%) that include more than 20 species show significant slowdowns. The high frequency of slowdowns observed in large clades is even more extreme than expected under a purely stochastic constant-rate model, but is consistent with the adaptive radiation model. Taken together, our data strongly support a model of density-dependent speciation in birds, whereby speciation slows as ecological opportunities and geographical space place limits on clade growth. It is probable that the number of species that a given region can support is limited; however, it is unclear whether the limit is approached sufficiently in nature such that the rate at which new species form slows down. Using the pattern of phylogenetic branching, a technique that estimates evolutionary relationships based on molecular data, we demonstrate that in large clades of birds, there is a decrease in the per-lineage probability of speciation as the number of species in the clade increase. We also show that this pattern can arise even if speciation and extinction occur randomly through time. This is because large clades are likely, by chance, to have rapidly speciated early in their history, and will relax back to the average speciation rate later on. We account for this effect, and we still find evidence that, as a clade grows to large size, the per-lineage probability of speciation declines. These results strongly suggest that speciation rates are slowed as environments fill up with competitors.