Clade Age and Species Richness Are Decoupled Across the Eukaryotic Tree of Life
Daniel L. Rabosky1,2#*, Graham J. Slater3#, Michael E. Alfaro3#
1 Department of Integrative Biology and Museum of Vertebrate Zoology, University of California, Berkeley, California, United States of America, 2 Department of Ecology and Evolutionary Biology, University of Michigan, Ann Arbor, Michigan, United States of America, 3 Department of Ecology and Evolutionary Biology, University of California, Los Angeles, California, United States of America
Abstract
Explaining the dramatic variation in species richness across the tree of life remains a key challenge in evolutionary biology. At the largest phylogenetic scales, the extreme heterogeneity in species richness observed among different groups of organisms is almost certainly a function of many complex and interdependent factors. However, the most fundamental expectation in macroevolutionary studies is simply that species richness in extant clades should be correlated with clade age: all things being equal, older clades will have had more time for diversity to accumulate than younger clades. Here, we test the relationship between stem clade age and species richness across 1,397 major clades of multicellular eukaryotes that collectively account for more than 1.2 million described species. We find no evidence that clade age predicts species richness at this scale. We demonstrate that this decoupling of age and richness is unlikely to result from variation in net diversification rates among clades. At the largest phylogenetic scales, contemporary patterns of species richness are inconsistent with unbounded diversity increase through time. These results imply that a fundamentally different interpretative paradigm may be needed in the study of phylogenetic diversity patterns in many groups of organisms.
Author Summary
Species richness varies by many orders of magnitude across the evolutionary "tree of life." Some groups, like beetles and flowering plants, contain nearly incomprehensible species diversity, but the overwhelming majority of groups contain far fewer species. Many processes presumably contribute to this variation in diversity, but the most general explanatory variable is the evolutionary age of each group: older groups will simply have had more time for diversity to accumulate than younger groups. We tested whether evolutionary age explains differences in species richness by compiling diversity and age estimates for nearly 1,400 groups of multicellular organisms. Surprisingly, we find no evidence that old groups have more species than young groups. This result appears to hold across the entire tree of life, for taxa as diverse as ferns, fungi, and flies. We demonstrate that this pattern is highly unlikely under simple but widely used evolutionary models that allow diversity to increase through time without bounds. Paleontologists have long contended that diversity-dependent processes have regulated species richness through time, and our results suggest that such processes have left a footprint on the living biota that can even be seen without data from the fossil record.
Citation: Rabosky DL, Slater GJ, Alfaro ME (2012) Clade Age and Species Richness Are Decoupled Across the Eukaryotic Tree of Life. PLoS Biol 10(8): e1001381. doi:10.1371/journal.pbio.1001381
Academic Editor: Georgina M. Mace, University College London, United Kingdom
Received: October 28, 2011; Accepted: July 12, 2012; Published: August 28, 2012
Copyright: © 2012 Rabosky et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: This research was supported by NSF-DEB-081-4277, by NSF-DEB-091-8748, and by the Miller Institute for Basic Research in Science at the University of California. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
Competing interests: The authors have declared that no competing interests exist.
Abbreviations: AIC, Akaike Information Criterion; MCMC, Markov Chain Monte Carlo; MEDUSA, Modeling Evolutionary Diversification Using Stepwise AIC; PGLS, phylogenetic generalized least squares; SES, standardized effect size
* E-mail: drabosky@umich.edu
# These authors contributed equally to this work.
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