Estimating diversification rates from phylogenetic information.

Patterns of species richness reflect the balance between speciation and extinction over the evolutionary history of life. These processes are influenced by the size and geographical complexity of regions, conditions of the environment, and attributes of individuals and species. Diversity within clades also depends on age and thus the time available for accumulating species. Estimating rates of diversification is key to understanding how these factors have shaped patterns of species richness. Several approaches to calculating both relative and absolute rates of speciation and extinction within clades are based on phylogenetic reconstructions of evolutionary relationships. As the size and quality of phylogenies increases, these approaches will find broader application. However, phylogeny reconstruction fosters a perceptual bias of continual increase in species richness, and the analysis of primarily large clades produces a data selection bias. Recognizing these biases will encourage the development of more realistic models of diversification and the regulation of species richness.

[1]  Sean Nee,et al.  Birth-Death Models in Macroevolution , 2006 .

[2]  E. Gilesleighjr The average lifetime of a population in a varying environment , 1981 .

[3]  R. Ricklefs,et al.  Evolutionary diversification of clades of squamate reptiles , 2007, Journal of evolutionary biology.

[4]  DIVERGENT TIMING AND PATTERNS OF SPECIES ACCUMULATION IN LOWLAND AND HIGHLAND NEOTROPICAL BIRDS , 2006, Evolution; international journal of organic evolution.

[5]  P H Harvey,et al.  Macroevolutionary inferences from primate phylogeny , 1995, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[6]  Eric Durand,et al.  apTreeshape: statistical analysis of phylogenetic tree shape , 2006, Bioinform..

[7]  A. Grafen The phylogenetic regression. , 1989, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[8]  A. Phillimore,et al.  Ecology Predicts Large‐Scale Patterns of Phylogenetic Diversification in Birds , 2006, The American Naturalist.

[9]  T. Clutton‐Brock,et al.  The Latitudinal Gradient in Recent Speciation and Extinction Rates of Birds and Mammals , 2007 .

[10]  E. Abouheif A method for testing the assumption of phylogenetic independence in comparative data , 1999 .

[11]  rme,et al.  TESTING FOR LATITUDINAL BIAS IN DIVERSIFICATION RATES : AN EXAMPLE USING NEW WORLD BIRDS , 2005 .

[12]  A. Vogler,et al.  Revealing the factors that promote speciation , 1998 .

[13]  E. Paradis,et al.  Analysis of comparative data using generalized estimating equations. , 2002, Journal of theoretical biology.

[14]  J. Hey USING PHYLOGENETIC TREES TO STUDY SPECIATION AND EXTINCTION , 1992, Evolution; international journal of organic evolution.

[15]  Brian R. Moore,et al.  SYMMETREE: whole-tree analysis of differential diversification rates , 2005, Bioinform..

[16]  Andy Purvis,et al.  Power of eight tree shape statistics to detect nonrandom diversification: a comparison by simulation of two models of cladogenesis. , 2002, Systematic biology.

[17]  Milton Rueda,et al.  Cenozoic Plant Diversity in the Neotropics , 2006, Science.

[18]  M. Chase,et al.  Environmental causes for plant biodiversity gradients. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[19]  H. Comes,et al.  Quaternary diversification in European alpine plants: pattern and process. , 2004, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[20]  Sister-group analysis in identifying correlates of diversification , 2004, Evolutionary Ecology.

[21]  E. Paradis STATISTICAL ANALYSIS OF DIVERSIFICATION WITH SPECIES TRAITS , 2005, Evolution; international journal of organic evolution.

[22]  M. Pagel,et al.  Large Punctuational Contribution of Speciation to Evolutionary Divergence at the Molecular Level , 2006, Science.

[23]  K. Chan,et al.  Whole-tree methods for detecting differential diversification rates. , 2002, Systematic biology.

[24]  Sean Nee,et al.  PHYLOGENIES WITHOUT FOSSILS , 1994, Evolution; international journal of organic evolution.

[25]  D. Schluter,et al.  Speciation, extinction and diversity , 2001 .

[26]  M. Sanderson,et al.  ABSOLUTE DIVERSIFICATION RATES IN ANGIOSPERM CLADES , 2001, Evolution; international journal of organic evolution.

[27]  John Wakeley,et al.  Estimating Divergence Times from Molecular Data on Phylogenetic and Population Genetic Timescales , 2002 .

[28]  Daniel L Rabosky,et al.  LIKELIHOOD METHODS FOR DETECTING TEMPORAL SHIFTS IN DIVERSIFICATION RATES , 2006, Evolution; international journal of organic evolution.

[29]  M. Sanderson,et al.  Age and rate of diversification of the Hawaiian silversword alliance (Compositae). , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[30]  D. Weisrock,et al.  Rapid lineage accumulation in a non-adaptive radiation: phylogenetic analysis of diversification rates in eastern North American woodland salamanders (Plethodontidae: Plethodon) , 2006, Proceedings of the Royal Society B: Biological Sciences.

[31]  Kate E. Jones,et al.  The delayed rise of present-day mammals , 1990, Nature.

[32]  CONTRASTING PATTERNS OF RADIATION IN AFRICAN AND AUSTRALIAN RESTIONACEAE , 2003, Evolution; international journal of organic evolution.

[33]  M. Cardillo Latitude and rates of diversification in birds and butterflies , 1999, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[34]  Steinar Engen,et al.  The Spatial Scale of Population Fluctuations and Quasi‐Extinction Risk , 2002, The American Naturalist.

[35]  Craig Moritz,et al.  Tropical rainforests: past, present and future. , 2005 .

[36]  R. Ricklefs Global diversification rates of passerine birds , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[37]  J. Alroy Successive approximations of diversity curves: Ten more years in the library , 2000 .

[38]  Helmut Hillebrand,et al.  On the Generality of the Latitudinal Diversity Gradient , 2004, The American Naturalist.

[39]  E. Paradis Assessing temporal variations in diversification rates from phylogenies: estimation and hypothesis testing , 1997, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[40]  R. Ricklefs,et al.  Phylogenetic perspectives on patterns of regional and local species richness. , 2005 .

[41]  Andrew Meade,et al.  Detecting the node-density artifact in phylogeny reconstruction. , 2006, Systematic biology.

[42]  Sudhir Kumar,et al.  Molecular clocks: four decades of evolution , 2005, Nature Reviews Genetics.

[43]  Daniel J. Howard,et al.  Endless Forms: Species and Speciation , 1998 .

[44]  P H Harvey,et al.  Tempo and mode of evolution revealed from molecular phylogenies. , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[45]  D. Janzen,et al.  Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[46]  R. Ricklefs The unified neutral theory of biodiversity: do the numbers add up? , 2006, Ecology.

[47]  M. Donoghue Key innovations, convergence, and success: macroevolutionary lessons from plant phylogeny , 2005, Paleobiology.

[48]  R. Lande Risks of Population Extinction from Demographic and Environmental Stochasticity and Random Catastrophes , 1993, The American Naturalist.

[49]  S. Hubbell,et al.  The unified neutral theory of biodiversity and biogeography at age ten. , 2011, Trends in ecology & evolution.

[50]  A. Vogler,et al.  The effect of habitat type on speciation rates and range movements in aquatic beetles: inferences from species‐level phylogenies , 2001, Molecular ecology.

[51]  J. Ahlquist Phylogeny and classification of birds , 1985 .

[52]  Nancy Knowlton,et al.  Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography. , 2007, Ecology letters.

[53]  Stephen Jay Gould,et al.  Stochastic Models of Phylogeny and the Evolution of Diversity , 1973, The Journal of Geology.

[54]  D. Penny,et al.  The modern molecular clock , 2003, Nature Reviews Genetics.

[55]  K. Queiroz**,et al.  The General Lineage Concept of Species, Species Criteria, and the Process of Speciation , 1998 .

[56]  S. Nee,et al.  INFERRING SPECIATION RATES FROM PHYLOGENIES , 2001, Evolution; international journal of organic evolution.

[57]  S. Renner Relaxed molecular clocks for dating historical plant dispersal events. , 2005, Trends in plant science.

[58]  TESTING FOR EQUAL RATES OF CLADOGENESIS IN DIVERSE TAXA , 2003, Evolution; international journal of organic evolution.

[59]  Oliver G. Pybus,et al.  Testing macro–evolutionary models using incomplete molecular phylogenies , 2000, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[60]  H. Linder,et al.  Taxon sampling effects in molecular clock dating: an example from the African Restionaceae. , 2005, Molecular phylogenetics and evolution.

[61]  E. Paradis Analysis of diversification: combining phylogenetic and taxonomic data , 2003, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[62]  G. J. Rodgers,et al.  Speciation and extinction in a simple model of evolution , 1996 .

[63]  A. P. Allen,et al.  Assessing latitudinal gradients in speciation rates and biodiversity at the global scale. , 2006, Ecology letters.

[64]  S. Heard,et al.  PATTERNS IN PHYLOGENETIC TREE BALANCE WITH VARIABLE AND EVOLVING SPECIATION RATES , 1996, Evolution; international journal of organic evolution.

[65]  F. Ayala,et al.  A methodological bias toward overestimation of molecular evolutionary time scales , 2002, Proceedings of the National Academy of Sciences of the United States of America.

[66]  Lindell Bromham,et al.  Molecular dating when rates vary. , 2005, Trends in ecology & evolution.

[67]  E. Paradis Detecting Shifts in Diversification Rates without Fossils , 1998, The American Naturalist.

[68]  R M May,et al.  The reconstructed evolutionary process. , 1994, Philosophical transactions of the Royal Society of London. Series B, Biological sciences.

[69]  C. Guyer,et al.  Testing Whether Certain Traits have Caused Amplified Diversification: An Improved Method Based on a Model of Random Speciation and Extinction , 1993, The American Naturalist.

[70]  J. W. Valentine,et al.  Morphological complexity increase in metazoans , 1994, Paleobiology.

[71]  Alexei J Drummond,et al.  Time dependency of molecular rate estimates and systematic overestimation of recent divergence times. , 2005, Molecular biology and evolution.

[72]  R. Ricklefs Global variation in the diversification rate of passerine birds. , 2006, Ecology.