Abstract

Explaining the uneven distribution of species richness across the branches of the tree of life has been a major challenge for evolutionary biologists. Advances in phylogenetic reconstruction, allowing the generation of large, well-sampled, phylogenetic trees have provided an opportunity to contrast competing hypotheses. Here, we present a new time-calibrated phylogeny of seed plant families using Bayesian methods and 26 fossil calibrations. While there are various published phylogenetic trees for plants which have a greater density of species sampling, we are still a long way from generating a complete phylogeny for all ~300,000+ plants. Our phylogeny samples all seed plant families and is a useful tool for comparative analyses. We use this new phylogenetic hypothesis to contrast two alternative explanations for differences in species richness among higher taxa: time for speciation versus ecological limits. We calculated net diversification rate for each clade in the phylogeny and assessed the relationship between clade age and species richness. We then fit models of speciation and extinction to individual branches in the tree to identify major rate-shifts. Our data suggest that the majority of lineages are diversifying very slowly while a few lineages, distributed throughout the tree, are diversifying rapidly. Diversification is unrelated to clade age, no matter the age range of the clades being examined, contrary to both the assumption of an unbounded lineage increase through time, and the paradigm of fixed ecological limits. These findings are consistent with the idea that ecology plays a role in diversification, but rather than imposing a fixed limit, it may have variable effects on per lineage diversification rates through time.

Highlights

  • It is well recognised that species richness is unevenly distributed across the tree of life and the origin of this variation is the subject of significant debate [1,2,3,4,5]

  • Total diversification, and diversification rate estimation are reported for all 849 clades of the phylogeny (S4 Table)

  • The magnitude of extinction influenced the absolute estimates of diversification rate, changes to the rank order of clades were generally modest, with the rank of 8 of the top 10 fastest diversifying clades being conserved between the two models

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Summary

Introduction

It is well recognised that species richness is unevenly distributed across the tree of life and the origin of this variation is the subject of significant debate [1,2,3,4,5]. Ecological Limit vs Time for Speciation species richness between sister clades is often large, leading to highly imbalanced phylogenetic topologies [6, 7]. Despite the long history of research on these patterns and an increasingly accurate understanding of the evolutionary history for many taxonomic groups, the mechanisms underlying differences in species richness remain largely undetermined, with possible explanations spanning biological, historical, geographical and neutral processes [8]. A better understanding of the causes of imbalance in species richness across lineages could provide insight into the mechanisms governing the evolution and proliferation of life. We reconstruct a complete phylogenetic tree for all seed plant families, and use this tree to evaluate two alternative explanations for variation in species richness among clades: ‘time for speciation’ versus ‘ecological limits’ [4]

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