gen3sis: the general engine for eco-evolutionary simulations on the origins of biodiversity

Understanding the origins of biodiversity has been an aspiration since the days of early naturalists. The immense complexity of ecological, evolutionary and spatial processes, however, has made this goal elusive to this day. Computer models serve progress in many scientific fields, but in the fields of macroecology and macroevolution, eco-evolutionary models are comparatively less developed. We present a general, spatially-explicit, eco-evolutionary engine with a modular implementation that enables the modelling of multiple macroecological and macroevolutionary processes and feedbacks across representative spatio-temporally dynamic landscapes. Modelled processes can include environmental filtering, biotic interactions, dispersal, speciation and evolution of ecological traits. Commonly observed biodiversity patterns, such as α, β and γ diversity, species ranges, ecological traits and phylogenies, emerge as simulations proceed. As a case study, we examined alternative hypotheses expected to have shaped the latitudinal diversity gradient (LDG) during the Earth’s Cenozoic era. We found that a carrying capacity linked with energy was the only model variant that could simultaneously produce a realistic LDG, species range size frequencies, and phylogenetic tree balance. The model engine is open source and available as an R-package, enabling future exploration of various landscapes and biological processes, while outputs can be linked with a variety of empirical biodiversity patterns. This work represents a step towards a numeric and mechanistic understanding of the physical and biological processes that shape Earth’s biodiversity.

[1]  Franck Pommereau,et al.  Discrete-event models for conservation assessment of integrated ecosystems , 2020, Ecol. Informatics.

[2]  Roger Bivand,et al.  Progress in the R ecosystem for representing and handling spatial data , 2020, Journal of Geographical Systems.

[3]  G. Dauby,et al.  Tectonics, climate and the diversification of the tropical African terrestrial flora and fauna , 2020, Biological reviews of the Cambridge Philosophical Society.

[4]  F. Delsuc,et al.  The role of the Neotropics as a source of world tetrapod biodiversity , 2020, Global Ecology and Biogeography.

[5]  A. Culina,et al.  Low availability of code in ecology: A call for urgent action , 2020, PLoS biology.

[6]  K. Nisancioglu,et al.  Global Cenozoic Paleobathymetry with a focus on the Northern Hemisphere Oceanic Gateways , 2020 .

[7]  A. S. Meseguer,et al.  Ancient tropical extinctions at high latitudes contributed to the latitudinal diversity gradient * , 2020, Evolution; international journal of organic evolution.

[8]  A. Tanentzap,et al.  Angiosperm speciation cools down in the tropics , 2020, Ecology letters.

[9]  M. Di Febbraro,et al.  A 450 million years long latitudinal gradient in age‐dependent extinction , 2019, Ecology letters.

[10]  P. Valdes,et al.  Extinction intensity during Ordovician and Cenozoic glaciations explained by cooling and palaeogeography , 2019, Nature Geoscience.

[11]  E. Bertuzzo,et al.  Mapping landscape connectivity as a driver of species richness under tectonic and climatic forcing , 2019, Earth Surface Dynamics.

[12]  J. Okie,et al.  The carrying capacity for species richness , 2019, Global Ecology and Biogeography.

[13]  J. Singarayer,et al.  Spatio-temporal climate change contributes to latitudinal diversity gradients , 2019, Nature Ecology & Evolution.

[14]  D. Bonte,et al.  A simple spatially explicit neutral model explains the range size distribution of reef fishes , 2019, Global Ecology and Biogeography.

[15]  K. Wiegand,et al.  Assessing predicted isolation effects from the general dynamic model of island biogeography with an eco‐evolutionary model for plants , 2019, Journal of Biogeography.

[16]  K. Wiegand,et al.  Interactions between ecological, evolutionary and environmental processes unveil complex dynamics of insular plant diversity , 2019, Journal of Biogeography.

[17]  K. Connellan Who am I and why? , 2019, Dreaming, Healing and Imaginative Arts Practice.

[18]  Enrico Bertuzzo,et al.  Evolving biodiversity patterns in changing river networks. , 2019, Journal of theoretical biology.

[19]  M. Cardillo,et al.  Reconstructing the Geography of Speciation from Contemporary Biodiversity Data , 2019, The American Naturalist.

[20]  Chengshan Wang,et al.  The evolution of latitudinal temperature gradients from the latest Cretaceous through the Present , 2019, Earth-Science Reviews.

[21]  Mikael Pontarp,et al.  Inferring community assembly processes from macroscopic patterns using dynamic eco‐evolutionary models and Approximate Bayesian Computation (ABC) , 2019, Methods in Ecology and Evolution.

[22]  T. Stadler,et al.  Closing the gap between palaeontological and neontological speciation and extinction rate estimates , 2018, Nature Communications.

[23]  M. Pärtel,et al.  Plant diversity in Oceanic archipelagos: realistic patterns emulated by an agent‐based computer simulation , 2018, Ecography.

[24]  E. Paradis,et al.  Evaluating alternative explanations for an association of extinction risk and evolutionary uniqueness in multiple insular lineages , 2018, Evolution; international journal of organic evolution.

[25]  Philip B. Holden,et al.  Modeling the ecology and evolution of biodiversity: Biogeographical cradles, museums, and graves , 2018, Science.

[26]  Daiqin Li,et al.  Formation of rivers and mountains drives diversification of primitively segmented spiders in continental East Asia , 2018, Journal of Biogeography.

[27]  M. Huber,et al.  Synchronous tropical and polar temperature evolution in the Eocene , 2018, Nature.

[28]  G. Glauser,et al.  Growth-competition-herbivore resistance trade-offs and the responses of alpine plant communities to climate change , 2018 .

[29]  J. Okie,et al.  The more-individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity-diversity relationship. , 2018, Ecology letters.

[30]  R. Milo,et al.  The biomass distribution on Earth , 2018, Proceedings of the National Academy of Sciences.

[31]  Owen L. Petchey,et al.  Ecological opportunity and predator–prey interactions: linking eco-evolutionary processes and diversification in adaptive radiations , 2018, Proceedings of the Royal Society B: Biological Sciences.

[32]  P. Valdes,et al.  Eocene greenhouse climate revealed by coupled clumped isotope-Mg/Ca thermometry , 2018, Proceedings of the National Academy of Sciences.

[33]  Carsten Rahbek,et al.  Process, Mechanism, and Modeling in Macroecology. , 2017, Trends in ecology & evolution.

[34]  Matthew W. Pennell,et al.  Speciation gradients and the distribution of biodiversity , 2017, Nature.

[35]  Patrice Descombes,et al.  Spatial imprints of plate tectonics on extant richness of terrestrial vertebrates , 2017 .

[36]  G. Mittelbach,et al.  "Latitudinal Gradients in Species Diversity": Reflections on Pianka's 1966 Article and a Look Forward. , 2017, The American naturalist.

[37]  J. Wiens,et al.  The origin of species richness patterns along environmental gradients: uniting explanations based on time, diversification rate and carrying capacity , 2017 .

[38]  F. Mazel,et al.  Global determinants of zoogeographical boundaries , 2017, Nature Ecology &Evolution.

[39]  Jacob van Etten,et al.  R package gdistance: distances and routes on geographical grids (version 1.1-4) , 2012 .

[40]  Gregory J. L. Tourte,et al.  The DeepMIP contribution to PMIP4 , 2017 .

[41]  F. Hartig,et al.  Mechanistic simulation models in macroecology and biogeography: state‐of‐art and prospects , 2017 .

[42]  N. Salamin,et al.  Niche width impacts vertebrate diversification , 2016 .

[43]  Jorge Hidalgo,et al.  Eco-evolutionary Model of Rapid Phenotypic Diversification in Species-Rich Communities , 2016, PLoS Comput. Biol..

[44]  M. K. Borregaard,et al.  The general dynamic model: towards a unified theory of island biogeography? , 2016 .

[45]  Constantinos Tsirogiannis,et al.  PhyloMeasures: a package for computing phylogenetic biodiversity measures and their statistical moments , 2016 .

[46]  D. Tittensor,et al.  A neutral‐metabolic theory of latitudinal biodiversity , 2016 .

[47]  Carlos J. Melián,et al.  Plate tectonics drive tropical reef biodiversity dynamics , 2016, Nature Communications.

[48]  Jeet Sukumaran,et al.  Machine Learning Biogeographic Processes from Biotic Patterns: A New Trait-Dependent Dispersal and Diversification Model with Model Choice By Simulation-Trained Discriminant Analysis. , 2016, Systematic biology.

[49]  L. Pellissier Stability and the competition-dispersal trade-off as drivers of speciation and biodiversity gradients , 2015, Front. Ecol. Evol..

[50]  J. Wiens,et al.  Is diversification rate related to climatic niche width , 2015 .

[51]  Amaury Lambert,et al.  Phylogenies support out-of-equilibrium models of biodiversity. , 2015, Ecology letters.

[52]  M. Steel,et al.  Age-Dependent Speciation Can Explain the Shape of Empirical Phylogenies , 2015, Systematic biology.

[53]  Nathan J B Kraft,et al.  Functional trait space and the latitudinal diversity gradient , 2014, Proceedings of the National Academy of Sciences.

[54]  H. Dijkstra,et al.  The role of ocean gateways on cooling climate on long time scales , 2014 .

[55]  J. Stegen,et al.  When should species richness be energy limited, and how would we know? , 2014, Ecology letters.

[56]  H. Morlon Phylogenetic approaches for studying diversification. , 2014, Ecology letters.

[57]  A. Duputié,et al.  An empiricist's guide to theoretical predictions on the evolution of dispersal , 2013, Interface Focus.

[58]  R. A. Pyron,et al.  Large-scale phylogenetic analyses reveal the causes of high tropical amphibian diversity , 2013, Proceedings of the Royal Society B: Biological Sciences.

[59]  Ignacio Quintero,et al.  Rates of projected climate change dramatically exceed past rates of climatic niche evolution among vertebrate species. , 2013, Ecology letters.

[60]  Stuart L. Pimm,et al.  Global patterns of terrestrial vertebrate diversity and conservation , 2013, Proceedings of the National Academy of Sciences.

[61]  Susanne A. Fritz,et al.  An Update of Wallace’s Zoogeographic Regions of the World , 2013, Science.

[62]  M. Benton Origins of Biodiversity , 2013, PLoS biology.

[63]  Juliano Sarmento Cabral,et al.  Linking ecological niche, community ecology and biogeography: insights from a mechanistic niche model , 2012 .

[64]  J. Annan,et al.  A new global reconstruction of temperature changes at the Last Glacial Maximum , 2012 .

[65]  H. Balslev,et al.  Cenozoic imprints on the phylogenetic structure of palm species assemblages worldwide , 2012, Proceedings of the National Academy of Sciences.

[66]  O. Seehausen,et al.  Thermal adaptation and ecological speciation , 2011, Molecular ecology.

[67]  Steven F Railsback,et al.  Pattern-oriented modelling: a ‘multi-scope’ for predictive systems ecology , 2012, Philosophical Transactions of the Royal Society B: Biological Sciences.

[68]  Hélène Morlon,et al.  Reconciling molecular phylogenies with the fossil record , 2011, Proceedings of the National Academy of Sciences.

[69]  L. Ivany,et al.  Warm, not super-hot, temperatures in the early Eocene subtropics , 2011 .

[70]  M. C. Urban The evolution of species interactions across natural landscapes. , 2011, Ecology Letters.

[71]  Tanja Stadler,et al.  Mammalian phylogeny reveals recent diversification rate shifts , 2011, Proceedings of the National Academy of Sciences.

[72]  Nathan G. Swenson,et al.  Deterministic tropical tree community turnover: evidence from patterns of functional beta diversity along an elevational gradient , 2011, Proceedings of the Royal Society B: Biological Sciences.

[73]  A. Phillimore,et al.  The shape and temporal dynamics of phylogenetic trees arising from geographic speciation. , 2010, Systematic biology.

[74]  Damaris Zurell,et al.  The virtual ecologist approach: simulating data and observers , 2010 .

[75]  B. Enquist,et al.  Advancing the metabolic theory of biodiversity. , 2009, Ecology letters.

[76]  Walter Jetz,et al.  Patterns and causes of species richness: a general simulation model for macroecology. , 2009, Ecology letters.

[77]  G. Mittelbach,et al.  Is There a Latitudinal Gradient in the Importance of Biotic Interactions , 2009 .

[78]  Richard Field,et al.  Spatial species‐richness gradients across scales: a meta‐analysis , 2009 .

[79]  C. Graham,et al.  Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time. , 2008, Ecology letters.

[80]  O. Ronce How Does It Feel to Be Like a Rolling Stone? Ten Questions About Dispersal Evolution , 2007 .

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

[82]  James H. Brown,et al.  Kinetic effects of temperature on rates of genetic divergence and speciation. , 2006, Proceedings of the National Academy of Sciences of the United States of America.

[83]  Owen L. Petchey,et al.  Functional diversity: back to basics and looking forward. , 2006, Ecology letters.

[84]  Uta Berger,et al.  Pattern-Oriented Modeling of Agent-Based Complex Systems: Lessons from Ecology , 2005, Science.

[85]  C. Graham,et al.  Niche Conservatism: Integrating Evolution, Ecology, and Conservation Biology , 2005 .

[86]  J. Diniz‐Filho,et al.  An evolutionary tolerance model explaining spatial patterns in species richness under environmental gradients and geometric constraints , 2005 .

[87]  James H. Brown,et al.  The rate of DNA evolution: effects of body size and temperature on the molecular clock. , 2005, Proceedings of the National Academy of Sciences of the United States of America.

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

[89]  R. Ricklefs,et al.  Beta diversity of angiosperms in temperate floras of eastern Asia and eastern North America , 2004 .

[90]  P. Stephens,et al.  Explaining Species Richness from Continents to Communities: The Time‐for‐Speciation Effect in Emydid Turtles , 2002, The American Naturalist.

[91]  L. Sloan,et al.  Trends, Rhythms, and Aberrations in Global Climate 65 Ma to Present , 2001, Science.

[92]  G. Graves,et al.  Multiscale assessment of patterns of avian species richness , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[93]  D. Aldous Stochastic models and descriptive statistics for phylogenetic trees, from Yule to today , 2001 .

[94]  K. Gaston Global patterns in biodiversity , 2000, Nature.

[95]  J. Lawton Are there general laws in ecology , 1999 .

[96]  James S. Clark,et al.  Why Trees Migrate So Fast: Confronting Theory with Dispersal Biology and the Paleorecord , 1998, The American Naturalist.

[97]  T. F. Hansen STABILIZING SELECTION AND THE COMPARATIVE ANALYSIS OF ADAPTATION , 1997, Evolution; international journal of organic evolution.

[98]  K. Gaston,et al.  Species-range-size distributions: patterns, mechanisms and implications. , 1996, Trends in ecology & evolution.

[99]  K. Rohde Latitudinal gradients in species diversity: the search for the primary cause , 1992 .

[100]  R. Lewin Biologists Disagree Over Bold Signature of Nature: Adaptation to the different annual climatic fluctuations at low and high latitudes is suggested as an explanation for a famous, unexplained pattern of nature. , 1989, Science.

[101]  S. Stanley Population size, extinction, and speciation: the fission effect in Neogene Bivalvia , 1986, Paleobiology.

[102]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[103]  R. Macarthur,et al.  The Limiting Similarity, Convergence, and Divergence of Coexisting Species , 1967, The American Naturalist.

[104]  E. Pianka Convexity, Desert Lizards, and Spatial Heterogeneity , 1966 .

[105]  R. Macarthur PATTERNS OF SPECIES DIVERSITY , 1965 .

[106]  R. Macarthur,et al.  AN EQUILIBRIUM THEORY OF INSULAR ZOOGEOGRAPHY , 1963 .

[107]  A. Gray,et al.  I. THE ORIGIN OF SPECIES BY MEANS OF NATURAL SELECTION , 1963 .

[108]  A. G. Fischer LATITUDINAL VARIATIONS IN ORGANIC DIVERSITY , 1960 .

[109]  C. Pigott Genetics and the Origin of Species , 1959, Nature.

[110]  D. Rosauer,et al.  Are global hotspots of endemic richness shaped by plate tectonics , 2018 .

[111]  Katja Gruenewald,et al.  Species Diversity In Space And Time , 2016 .

[112]  C. Scotese,et al.  Phanerozoic Paleoclimate: An Atlas of Lithologic Indicators of Climate , 2013 .

[113]  Peter Dalgaard,et al.  R Development Core Team (2010): R: A language and environment for statistical computing , 2010 .

[114]  J. Emlen,et al.  ON THE RELATIONSHIP BETWEEN ABUNDANCE AND DISTRIBUTION OF SPECIES , 2008 .

[115]  H. Shaffer,et al.  Annual review of ecology, evolution, and systematics , 2003 .

[116]  T. Ohta THE NEARLY NEUTRAL THEORY OF MOLECULAR EVOLUTION , 1992 .

[117]  R. Levins The strategy of model building in population biology , 1966 .

[118]  E LECTRONIC SUPPLEMENTARY MATERIAL : 1 ECOLOGICAL CONSTRAINTS COUPLED WITH DEEP-TIME HABITAT DYNAMICS 2 PREDICT THE LATITUDINAL DIVERSITY GRADIENT IN REEF FISHES , 2022 .