The Origins of the Latitudinal Diversity Gradient: Revisiting the Tropical Conservatism Hypothesis
Corresponding Author
John J. Wiens
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA
Correspondence:
John J. Wiens ([email protected])
Search for more papers by this authorMichael J. Donoghue
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA
Department of Ecology and Evolutionary Biology, Yale University, New Haven, Connecticut, USA
Search for more papers by this authorCorresponding Author
John J. Wiens
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA
Correspondence:
John J. Wiens ([email protected])
Search for more papers by this authorMichael J. Donoghue
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA
Department of Ecology and Evolutionary Biology, Yale University, New Haven, Connecticut, USA
Search for more papers by this authorFunding: The authors received no specific funding for this research.
ABSTRACT
Aim
Understanding the origins of species richness patterns (especially high tropical richness) is a long-standing challenge at the intersection of biogeography, ecology, and evolutionary biology. One hypothesis that can potentially explain the latitudinal richness gradient is the tropical conservatism hypothesis (TCH). The TCH proposes that there are presently more species in tropical regions because many clades originated in the tropics and have only colonised the temperate zones more recently, leaving less time for speciation to build up temperate richness, and with niche conservatism limiting temperate colonisation by tropical clades. Here, we review the empirical evidence for the TCH. We first define this hypothesis, outline its major predictions, and describe its relationship to similar hypotheses. We then perform a systematic review to quantitatively evaluate the support for (and against) its major predictions. Finally, we describe several areas for future research.
Location
Global.
Time Period
Present to ~750 million years ago.
Major Taxa Studied
All (especially plants and animals).
Methods
We perform a systematic review of the evidence for the TCH over the last ~20 years.
Results
Most predictions of the TCH were supported in a significant majority of the studies that examined them. Further, a significant majority of relevant studies rejected the role of higher tropical diversification rates in driving the latitudinal diversity gradient (contrary to the diversification-rate and out-of-the-tropics hypotheses). Surprisingly, the importance of diversification rates did not depend on the ages of the clades studied.
Main Conclusions
Our results generally support the TCH, but also highlight several important issues moving forward. Most studies tested very few predictions of the TCH, and the pivotal role of colonisation time was often untested. Many studies analysed phylogenetic diversity measures, but their relevance for explaining richness patterns remains disturbingly unclear. Finally, we discuss several unresolved questions about the TCH and the origins of richness patterns.
Conflicts of Interest
The authors declare no conflicts of interest.
Open Research
Data Availability Statement
The data and code used are available as Datasets S1–S5, which are freely available on figshare at: https://doi.org/10.6084/m9.figshare.25505086.
Supporting Information
Filename | Description |
---|---|
jbi15172-sup-0001-DatasetS1.docxWord 2007 document , 113 KB |
Supplementary Material, including Tables S1, S2, and S3 and Appendix S1. |
Please note: The publisher is not responsible for the content or functionality of any supporting information supplied by the authors. Any queries (other than missing content) should be directed to the corresponding author for the article.
References
- Algar, A. C., J. T. Kerr, and D. J. Currie. 2009. “Evolutionary Constraints on Regional Faunas: Whom, but Not How Many.” Ecology Letters 12: 57–65.
- Allen, A. P., and J. F. Gillooly. 2006. “Assessing Latitudinal Gradients in Speciation Rates and Biodiversity at the Global Scale.” Ecology Letters 9: 947–954.
- Althaus, K. N., M. S. González-Elizondo, A. González-Rodríguez, H. Rodríguez Correa, and A. L. Hipp. 2025. “Phylogenetic Niche Conservatism Drives Floristic Assembly Across Mexico's Temperate-Tropical Divide.” Global Ecology and Biogeography 34: e70017. https://doi.org/10.1111/geb.70017.
- Barros, M. J., J. A. F. Diniz-Filho, and L. B. Freitas. 2018. “Ecological Drivers of Plant Genetic Diversity at the Southern Edge of Geographical Distributions: Forestal Vines in a Temperate Region.” Genetics and Molecular Biology 41: 318–326.
- Beerling, D. J., and F. I. Woodward. 2001. Vegetation and the Terrestrial Carbon Cycle: Modeling the First 400 Million Years. Cambridge University Press.
10.1017/CBO9780511541940 Google Scholar
- Blomberg, S. P., T. Garland, and A. R. Ives. 2003. “Testing for Phylogenetic Signal in Comparative Data: Behavioral Traits Are More Labile.” Evolution 57: 717–745.
- Brodie, J. F., and P. D. Mannion. 2023. “The Hierarchy of Factors Predicting the Latitudinal Diversity Gradient.” Trends in Ecology & Evolution 38: 15–23.
- Brown, J. H. 2014. “Why Are There So Many Species in the Tropics?” Journal of Biogeography 41: 8–22.
- Brown, J. H., and M. V. Lomolino. 1998. Biogeography. 2nd ed. Sinauer Associates.
- Carta, A., L. Peruzzi, and S. Ramirez-Barahona. 2022. “A Global Phylogenetic Regionalization of Vascular Plants Reveals a Deep Split Between Gondwanan and Laurasian Biotas.” New Phytologist 233: 1494–1504.
- Crisp, M. D., and L. G. Cook. 2012. “Phylogenetic Niche Conservatism: What Are the Underlying Evolutionary and Ecological Causes?” New Phytologist 196: 681–694.
- Donoghue, M. J., and E. J. Edwards. 2014. “Biome Shifts and Niche Evolution in Plants.” Annual Review of Ecology, Evolution, and Systematics 45: 547–572.
- Economo, E. P., N. Narula, N. R. Friedman, M. D. Weiser, and B. Guénard. 2018. “Macroecology and Macroevolution of the Latitudinal Diversity Gradient in Ants.” Nature Communications 9: 1778.
- Etienne, R. S., J. Sarmento Cabral, O. Hagen, et al. 2019. “A Minimal Model for the Latitudinal Diversity Gradient Suggests a Dominant Role for Ecological Limits.” American Naturalist 194: E122–E133.
- Farrell, B. D., C. Mitter, and D. J. Futuyma. 1992. “Diversification at the Plant–Insect Interface.” Bioscience 42: 34–42.
10.2307/1311626 Google Scholar
- Fernández, L. D., C. V. W. Seppey, D. Singer, et al. 2022. “Niche Conservatism Drives the Elevational Diversity Gradient in Major Groups of Free-Living Soil Unicellular Eukaryotes.” Microbial Ecology 83: 459–469.
- Fine, P. V. A. 2015. “Ecological and Evolutionary Drivers of Geographic Variation in Species Diversity.” Annual Review of Ecology, Evolution, and Systematics 46: 369–392.
- Fine, P. V. A., and R. H. Ree. 2006. “Evidence for a Time-Integrated Species-Area Effect on the Latitudinal Gradient in Tree Diversity.” American Naturalist 168: 796–804.
- Fischer, A. G. 1960. “Latitudinal Variation in Organic Diversity.” Evolution 14: 64–81.
- Futuyma, D. J. 1998. Evolutionary Biology. 3rd ed. Sinauer Associates.
- Galván, S., S. Varela, and S. Gamboa. 2025. “Classic Hypotheses of Area, Time, and Climatic Stability Fall Short in Explaining High Tropical Species Richness.” Journal of Biogeography 52: e15126.
- Harmon, L. J., and S. Harrison. 2015. “Species Diversity Is Dynamic and Unbounded at Local and Continental Scales.” American Naturalist 185: 584–593.
- Hawkins, B. A. 2001. “Ecology's Oldest Pattern?” Trends in Ecology & Evolution 16: 470.
- Helmus, M. R., T. J. Bland, C. K. Williams, and A. R. Ives. 2007. “Phylogenetic Measures of Biodiversity.” American Naturalist 169: E69–E83.
- Hernández-Hernández, T., and J. J. Wiens. 2020. “Why Are There So Many Flowering Plants? A Multi-Scale Analysis of Plant Diversification.” American Naturalist 195: 948–963.
10.1086/708273 Google Scholar
- Holt, B. G., J.-P. Lessard, M. K. Borregaard, et al. 2013. “An Update of Wallace's Zoogeographic Regions of the World.” Science 339: 74–78.
- Jablonski, D., K. Roy, and J. W. Valentine. 2006. “Out of the Tropics: Evolutionary Dynamics of the Latitudinal Diversity Gradient.” Science 314: 102–106.
- Jansson, R., G. Rodriguez-Castaneda, and L. E. Harding. 2013. “What Can Multiple Phylogenies Say About the Latitudinal Diversity Gradient? A New Look at the Tropical Conservatism, Out of the Tropics, and Diversification Rate Hypotheses.” Evolution 67: 1741–1755.
- Kerkhoff, A. J., P. E. Moriarity, and M. D. Weiser. 2014. “The Latitudinal Species Richness Gradient in New World Woody Angiosperms Is Consistent With the Tropical Conservatism Hypothesis.” Proceedings of the National Academy of Sciences of the United States of America 111: 8125–8130.
- Koehler, K., A. Center, and J. Cavender-Bares. 2012. “Evidence for a Freezing Tolerance–Growth Rate Trade-Off in the Live Oaks (Quercus Series Virentes) Across the Tropical–Temperate Divide.” New Phytologist 193: 730–744.
- Landis, M. J., N. J. Matzke, B. R. Moore, and J. P. Huelsenbeck. 2013. “Bayesian Analysis of Biogeography When the Number of Areas Is Large.” Systematic Biology 62: 789–804.
- Latham, R. E., and R. E. Ricklefs. 1993. “Global Patterns of Tree Species Richness in Moist Forests: Energy-Diversity Theory Does Not Account for Variation in Species Richness.” Oikos 67: 325–333.
- Li, H., and J. J. Wiens. 2019. “Time Explains Regional Richness Patterns Within Clades More Often Than Diversification Rates or Area.” American Naturalist 193: 514–529.
- Li, P., and J. J. Wiens. 2022. “What Drives Diversification? Range Expansion Tops Climate, Life History, Habitat, and Size in Lizards and Snakes.” Journal of Biogeography 49: 237–247.
- Looney, B. P., M. Ryberg, F. Hampe, M. Sánchez-Garcia, and B. P. Matheny. 2016. “Into and Out of the Tropics: Global Diversification Patterns in a Hyperdiverse Clade of Ectomycorrhizal Fungi.” Molecular Ecology 25: 630–647.
- Lu, L. M., L. F. Mao, T. Yang, et al. 2018. “Evolutionary History of the Angiosperm Flora of China.” Nature 554: 234–238.
- Machac, A. 2020. “The Dynamics of Bird Diversity in the New World.” Systematic Biology 69: 1180–1199.
- Mazel, F., T. J. Davies, L. Gallien, et al. 2016. “Influence of Tree Shape and Evolutionary Time-Scale on Phylogenetic Diversity Metrics.” Ecography 39: 913–1024.
- Meseguer, A. S., and F. L. Condamine. 2020. “Ancient Tropical Extinctions at High Latitudes Contributed to the Latitudinal Diversity Gradient.” Evolution 74: 1966–1987.
- Miller, E. C., K. T. Hayashi, D. Song, and J. J. Wiens. 2018. “Explaining the Ocean's Richest Biodiversity Hotspot and Global Patterns of Fish Diversity.” Proceedings of the Royal Society of London B: Biological Sciences 285: 20181314.
- Miller, E. C., and C. Román-Palacios. 2021. “Evolutionary Time Best Explains the Latitudinal Diversity Gradient of Living Freshwater Fish Diversity.” Global Ecology and Biogeography 30: 749–763.
- Miller, E. C., and J. J. Wiens. 2017. “Extinction and Time Help Drive the Marine-Terrestrial Biodiversity Gradient: Is the Ocean a Deathtrap?” Ecology Letters 20: 911–921.
- Mittelbach, G. G., D. W. Schemske, H. V. Cornell, et al. 2007. “Evolution and the Latitudinal Diversity Gradient: Speciation, Extinction and Biogeography.” Ecology Letters 10: 315–331.
- Moen, D. S., E. Cabrera-Guzmán, I. W. Caviedes-Solis, E. González-Bernal, and A. R. Hanna. 2022. “Phylogenetic Analysis of Adaptation in Comparative Physiology and Biomechanics: Overview and a Case Study of Thermal Physiology in Treefrogs.” Journal of Experimental Biology 225: jeb243292.
- Moreira, M. O., J. J. Wiens, C. Fonseca, and D. Rojas. 2024. “Climatic Niche Breadth, Niche Position, and Speciation in Lizards and Snakes.” Journal of Biogeography 51: 969–981.
- Morley, R. J. 2000. Origin and Evolution of Tropical Rain Forests. Wiley.
- O'Dea, R. E., M. Lagisz, M. D. Jennions, et al. 2021. “Preferred Reporting Items for Systematic Reviews and Meta-Analyses in Ecology and Evolutionary Biology: A PRISMA Extension.” Biological Reviews 96: 1695–1722.
- Oliveira, B. F., A. Machac, G. C. Costa, et al. 2016. “Species and Functional Diversity Accumulate Differently in Mammals.” Global Ecology and Biogeography 25: 1119–1130.
- Pagel, M. 1999. “Inferring the Historical Patterns of Biological Evolution.” Nature 401: 877–884.
- Pianka, E. R. 1966. “Latitudinal Gradients in Species Diversity: A Review of Concepts.” American Naturalist 100: 33–46.
- Pontarp, M., L. Bunnefeld, J. Sarmento Cabral, et al. 2019. “The Latitudinal Diversity Gradient: Novel Understanding Through Mechanistic Eco-Evolutionary Models.” Trends in Ecology & Evolution 34: 211–223.
- Pontarp, M., and J. J. Wiens. 2017. “The Origin of Species Richness Patterns Along Environmental Gradients: Uniting Explanations Based on Time, Diversification Rate, and Carrying Capacity.” Journal of Biogeography 44: 722–735.
- Pyron, R. A., and J. J. Wiens. 2013. “Large-Scale Phylogenetic Analyses Reveal the Causes of High Tropical Amphibian Diversity.” Proceedings of the Royal Society of London B: Biological Sciences 280: 20131622.
- Qian, H., and S. Chen. 2016. “Ecological Determinants of Mean Family Age of Angiosperm Trees in Forest Communities in China.” Scientific Reports 6, no. 1: 28662.
- Qian, H., Y. Zhang, R. E. Ricklefs, and X. Wang. 2022. “Relationship of Minimum Winter Temperature and Temperature Seasonality to the Northern Range Limit and Species Richness of Trees in North America.” Journal of Geographical Sciences 32: 280–290.
- R Core Team. 2023. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. https://www.R-project.org/.
- Rabosky, D. L. 2009. “Ecological Limits and Diversification Rate: Alternative Paradigms to Explain the Variation in Species Richness Among Clades and Regions.” Ecology Letters 12: 735–743.
- Ree, R. H., B. R. Moore, C. Webb, and M. J. Donoghue. 2005. “A Likelihood Framework for Inferring the Evolution of Geographic Range on Phylogenetic Trees.” Evolution 59: 2299–2311.
- Ree, R. H., and S. A. Smith. 2008. “Maximum Likelihood Inference of Geographic Range Evolution by Dispersal, Local Extinction, and Cladogenesis.” Systematic Biology 57: 4–14.
- Ricklefs, R. E. 1987. “Community Diversity: Relative Roles of Local and Regional Processes.” Science 235: 167–171.
- Ricklefs, R. E., and S. S. Renner. 1994. “Species Richness Within Families of Flowering Plants.” Evolution 48: 1619–1636.
- Ricklefs, R. E., and D. Schluter. 1993. “ Species Diversity: Regional and Historical Influences.” In Species Diversity in Ecological Communities: Historical and Geographical Perspectives, edited by R. E. Ricklefs and D. Schluter, 350–363. University of Chicago Press.
- Romdal, T. S., M. B. Araújo, and C. Rahbek. 2013. “Life on a Tropical Planet: Niche Conservatism and the Global Diversity Gradient.” Global Ecology and Biogeography 22: 344–350.
- Saupe, E. E. 2023. “Explanations for Latitudinal Diversity Gradients Must Invoke Rate Variation.” Proceedings of the National Academy of Sciences of the United States of America 120: e2306220120.
- Schluter, D. 2016. “Speciation, Ecological Opportunity, and Latitude: (American Society of Naturalists Address).” American Naturalist 187: 1–18.
- Segovia, R. A., R. T. Pennington, T. R. Baker, et al. 2020. “Freezing and Water Availability Structure the Evolutionary Diversity of Trees Across the Americas.” Science Advances 6: eaaz5373.
- Stephens, P. R., M. J. Farrell, T. J. Davies, et al. 2025. “Global Diversity Patterns Are Explained by Diversification Rates and Dispersal at Ancient, Not Shallow, Timescales.” Systematic Biology: syaf018.
- Stephens, P. R., and J. J. Wiens. 2003. “Explaining Species Richness From Continents to Communities: The Time-For-Speciation Effect in Emydid Turtles.” American Naturalist 161: 112–128.
- Tolmos, M. L., H. Kreft, J. Ramirez, R. Ospina, and D. Craven. 2022. “Water and Energy Availability Mediate Biodiversity Patterns Along an Elevational Gradient in the Tropical Andes.” Journal of Biogeography 49: 712–726.
- Treseder, K. K., M. R. Maltz, B. A. Hawkins, N. Fierer, J. E. Stajich, and K. L. McGuire. 2014. “Evolutionary Histories of Soil Fungi Are Reflected in Their Large-Scale Biogeography.” Ecology Letters 17: 1086–1093.
- Tucker, C. M., M. W. Cadotte, S. B. Carvalho, et al. 2017. “A Guide to Phylogenetic Metrics for Conservation, Community Ecology and Macroecology.” Biological Reviews 92: 698–715.
- Vamosi, J. C., and S. M. Vamosi. 2010. “Key Innovations Within a Geographical Context in Flowering Plants: Towards Resolving Darwin's Abominable Mystery.” Ecology Letters 13: 1270–1279.
- Vieira, C., F. Steen, S. D'hondt, et al. 2021. “Global Biogeography and Diversification of a Group of Brown Seaweeds (Phaeophyceae) Driven by Clade-Specific Evolutionary Processes.” Journal of Biogeography 48: 703–715.
- Wang, Z., J. Fang, Z. Tang, and X. Lin. 2011. “Patterns, Determinants and Models of Woody Plant Diversity in China.” Proceedings of the Royal Society of London B: Biological Sciences 278: 2122–2132.
- Webb, C. O. 2000. “Exploring the Phylogenetic Structure of Ecological Communities: An Example for Rain Forest Trees.” American Naturalist 156: 145–155.
- Wiens, J. J. 2024. “Speciation Across Life and the Origins of Biodiversity Patterns.” Evolutionary Journal of the Linnean Society 3: kzae025.
10.1093/evolinnean/kzae025 Google Scholar
- Wiens, J. J., D. D. Ackerly, A. P. Allen, et al. 2010. “Niche Conservatism as an Emerging Principle in Ecology and Conservation Biology.” Ecology Letters 13: 1310–1324.
- Wiens, J. J., and M. J. Donoghue. 2004. “Historical Biogeography, Ecology, and Species Richness.” Trends in Ecology & Evolution 19: 639–644.
- Wiens, J. J., C. H. Graham, D. S. Moen, S. A. Smith, and T. W. Reeder. 2006. “Evolutionary and Ecological Causes of the Latitudinal Diversity Gradient in Hylid Frogs: Treefrog Trees Unearth the Roots of High Tropical Diversity.” American Naturalist 168: 579–596.
- Ziegler, A. M., G. Eshel, P. M. Rees, T. A. Rothfus, D. B. Rowley, and D. Sunderlin. 2003. “Tracing the Tropics Across Land and Sea: Permian to Present.” Lethaia 36: 227–254.