The biogeographic history of ruminant faunas determines the phylogenetic structure of their assemblages at different scales

Phylogenetic community structure may help us understand how macroecological and macroevolutionary processes shape assemblages at large geographical scales. In this paper, we test hypotheses linking the formation of large-scale assemblages, evolutionary processes and macroecology. To provide new insight into ruminant biogeography and evolution, phylogenetic community structure metrics were calculated for faunal assemblages at four hierarchical levels. Phylogenetic relatedness indices (net relatedness index and nearest taxon index) were determined for 59 ruminant assemblages at the landscape scale and scale of their respective climate domains (continuous biome stretches). Species pools at the global and biogeographic realm levels were used to construct null observation models. Signifi cantly, assemblages were selected if they were distributed across biogeographic realms and represented all the world ’ s biomes. Non-random patterns were also tested for biogeographic realms within the global ruminant species pool. By examining ruminant assemblages at diff erent scales we were able to observe that ruminant faunas show a distribution mainly limited within the boundaries of their biogeographic realms. However, the diversifi cation of some clades was found to be restricted to extremely arid domains in the Sahara and Arabia. Th e random patterns featured by other extreme climate domains could refl ect phylogenetically heterogeneous fi lling by less biome-restricted lineages outside Africa.

[1]  M. Donoghue,et al.  Historical biogeography, ecology and species richness. , 2004, Trends in ecology & evolution.

[2]  E. Vrba,et al.  Rapoport effect and biomic specialization in African mammals: revisiting the climatic variability hypothesis , 2005 .

[3]  E. Vrba Ecology in relation to speciation rates: some case histories of Miocene-Recent mammal clades , 1987, Evolutionary Ecology.

[4]  J. Richardson,et al.  Insights into the historical construction of species-rich biomes from dated plant phylogenies, neutral ecological theory and phylogenetic community structure. , 2006, The New phytologist.

[5]  E. Douzery,et al.  Molecular and morphological phylogenies of ruminantia and the alternative position of the moschidae. , 2003, Systematic biology.

[6]  Campbell O. Webb,et al.  Picante: R tools for integrating phylogenies and ecology , 2010, Bioinform..

[7]  S. Schultz Sexual Dimorphism in Gynodioecious Sidalcea hirtipes (Malvaceae). I. Seed, Fruit, and Ecophysiology , 2003, International Journal of Plant Sciences.

[8]  E. Douzery,et al.  The tribal radiation of the family Bovidae (Artiodactyla) and the evolution of the mitochondrial cytochrome b gene. , 1999, Molecular phylogenetics and evolution.

[9]  D. Ackerly Community Assembly, Niche Conservatism, and Adaptive Evolution in Changing Environments , 2003, International Journal of Plant Sciences.

[10]  A. Hassanin,et al.  Molecular phylogeny of caprines (Bovidae, Antilopinae): the question of their origin and diversification during the Miocene , 2005 .

[11]  S. Heard,et al.  The Shapes of Phylogenetic Trees of Clades, Faunas, and Local Assemblages: Exploring Spatial Pattern in Differential Diversification , 2007, The American Naturalist.

[12]  Yang Wang,et al.  Expansion of C4 ecosystems as an indicator of global ecological change in the late Miocene , 1993, Nature.

[13]  Campbell O. Webb,et al.  Exploring the Phylogenetic Structure of Ecological Communities: An Example for Rain Forest Trees , 2000, The American Naturalist.

[14]  Richard Field,et al.  ENERGY, WATER, AND BROAD‐SCALE GEOGRAPHIC PATTERNS OF SPECIES RICHNESS , 2003 .

[15]  M. Cardillo,et al.  Phylogenetic structure of mammal assemblages at large geographical scales: linking phylogenetic community ecology with macroecology , 2011, Philosophical Transactions of the Royal Society B: Biological Sciences.

[16]  E. Vrba,et al.  Plio-Pleistocene climatic change in the Turkana Basin (East Africa): evidence from large mammal faunas. , 2006, Journal of human evolution.

[17]  Jan Zima,et al.  The Atlas of European Mammals , 1999 .

[18]  A. Waldron LINEAGES THAT CHEAT DEATH: SURVIVING THE SQUEEZE ON RANGE SIZE , 2010, Evolution; international journal of organic evolution.

[19]  R. Ricklefs,et al.  A Morphological Analysis of the Structure of Communities of Lizards in Desert Habitats , 1981 .

[20]  R. Ricklefs History and Diversity: Explorations at the Intersection of Ecology and Evolution , 2007, The American Naturalist.

[21]  J. Hill,et al.  The Mammals of the Indomalayan Region: A Systematic Review , 1992 .

[22]  A. Couloux,et al.  Pattern and timing of diversification of Cetartiodactyla (Mammalia, Laurasiatheria), as revealed by a comprehensive analysis of mitochondrial genomes. , 2012, Comptes rendus biologies.

[23]  N. Gotelli Null model analysis of species co-occurrence patterns , 2000 .

[24]  J. D. Zulueta Allue Andrade J. L., 1990: Atlas Fitoclimático de España. Taxonomías. Ministerio de Agricultura, Pesca y Alimentación. Instituto Nacional de Investigaciones Agrarias. Departamento de Sistemas Forestales. Madrid. , 1990 .

[25]  E. Vrba Turnover-pulses, the Red Queen, and related topics , 1993 .

[26]  R. Ricklefs,et al.  Global concordance in diversity patterns of vascular plants and terrestrial vertebrates. , 2008, Ecology letters.

[27]  F. Woodward,et al.  Increases in terrestrial carbon storage from the Last Glacial Maximum to the present , 1990, Nature.

[28]  J. Eisenberg Mammals of the Neotropics , 1989 .

[29]  A. Bofarull,et al.  Influence of continental history on the ecological specialization and macroevolutionary processes in the mammalian assemblage of South America: Differences between small and large mammals , 2008, BMC Evolutionary Biology.

[30]  Elisabeth S. Vrba,et al.  The Fossil Record and Evolution of Bovidae: State of the Field , 2009 .

[31]  Alfred Russel Wallace,et al.  The geographical distribution of animals : with a study of the relations of living and extinct faunas as elucidating the past changes of the earth's surface / by Alfred Russel Wallace ; in two volumes ; with maps and illustrations. , 1876 .

[32]  Campbell O. Webb,et al.  Bioinformatics Applications Note Phylocom: Software for the Analysis of Phylogenetic Community Structure and Trait Evolution , 2022 .

[33]  J. Meester,et al.  The mammals of Africa. An identification manual. , 1971 .

[34]  Campbell O. Webb,et al.  Emerging patterns in the comparative analysis of phylogenetic community structure , 2009, Molecular ecology.

[35]  Walter Jetz,et al.  A framework for delineating biogeographical regions based on species distributions , 2010 .

[36]  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.

[37]  Steven W Kembel,et al.  Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests. , 2009, Ecology letters.

[38]  Elisabeth S. Vrba,et al.  Body size, biomic specialization and range size of African large mammals , 2005 .

[39]  Erik Meijaard,et al.  Phylogeny and co‐occurrence of mammal species on Southeast Asian islands , 2010 .

[40]  M. Pickford,et al.  Biostratigraphy and palaeobiogeography of East Africa and the Iberian peninsula , 1994 .

[41]  Atlas fitoclimático de España: taxonomías , 1990 .

[42]  Matthew R. Helmus,et al.  Phylogenetic Metrics of Community Similarity , 2010, The American Naturalist.

[43]  G. Eck,et al.  Patterns of abundance and diversity in late Cenozoic bovids from the Turkana and Hadar Basins, Kenya and Ethiopia , 2007 .

[44]  M. Fernández Bioclimatic discriminant capacity of terrestrial mammal faunas , 2001 .

[45]  P. Mein,et al.  Long-period astronomical forcing of mammal turnover , 2006, Nature.

[46]  T. J. Robinson,et al.  Impacts of the Cretaceous Terrestrial Revolution and KPg Extinction on Mammal Diversification , 2011, Science.

[47]  J. L. Gittleman,et al.  Global patterns in the phylogenetic structure of island mammal assemblages , 2008, Proceedings of the Royal Society B: Biological Sciences.

[48]  J. Gowlett Paleoclimate and evolution, with emphasis on human origins , 1999 .

[49]  G. Corbet Mammals of the Palaearctic Region: A Taxonomic Review , 1978 .

[50]  Maria A. Gandolfo,et al.  Phylogenetic biome conservatism on a global scale , 2009, Nature.

[51]  Campbell O. Webb,et al.  Trait Evolution, Community Assembly, and the Phylogenetic Structure of Ecological Communities , 2007, The American Naturalist.

[52]  Vegetation and climate , 1991 .

[53]  D. Jolly,et al.  Mid‐Holocene and glacial‐maximum vegetation geography of the northern continents and Africa , 2000 .

[54]  L. Dupont,et al.  Development of vegetation and continental aridity in northwestern Africa during the Late Pliocene: the pollen record of ODP site 658 , 1994 .

[55]  J. Kamilar,et al.  The phylogenetic structure of primate communities: variation within and across continents , 2010 .

[56]  Tomas Johansson,et al.  Key biosynthetic gene subfamily recruited for pheromone production prior to the extensive radiation of Lepidoptera , 2008, BMC Evolutionary Biology.

[57]  Arne Ø. Mooers,et al.  Inferring Evolutionary Process from Phylogenetic Tree Shape , 1997, The Quarterly Review of Biology.

[58]  Michael Lautenschlager,et al.  Modelling Global Vegetation Patterns and Terrestrial Carbon Storage at the Last Glacial Maximum , 1993 .

[59]  Manuel Hernández Fernández,et al.  Macroevolutionary Processes and Biomic Specialization: Testing the Resource-use Hypothesis , 2005, Evolutionary Ecology.

[60]  F. Schrenk,et al.  African biogeography, climate change, & human evolution , 1999 .

[61]  R. Baker,et al.  Squirrels: the animal answer guide , 2007 .

[62]  Jonathan Kingdon,et al.  The Kingdon field guide to African mammals , 1997 .

[63]  E. Vrba,et al.  A complete estimate of the phylogenetic relationships in Ruminantia: a dated species‐level supertree of the extant ruminants , 2005, Biological reviews of the Cambridge Philosophical Society.

[64]  Campbell O. Webb,et al.  Phylogenies and Community Ecology , 2002 .

[65]  J. Morales,et al.  Biomic Specialization and Speciation Rates in Ruminants (Cetartiodactyla, Mammalia): A Test of the Resource-Use Hypothesis at the Global Scale , 2011, PloS one.

[66]  J. Maldonado,et al.  The surprising evolutionary history of South American deer. , 2008, Molecular phylogenetics and evolution.

[67]  E. Vrba New fossils of Alcelaphini and Caprinae (Bovidae: Mammalia) from Awash, Ethiopia, and phylogenetic analysis of Alcelaphini , 1997 .

[68]  E. Vrba,et al.  Antelopes, deer, and relatives : fossil record, behavioral ecology, systematics, and conservation , 2000 .