Abstract
Roots are essential to the strategies plants use to survive in variable environments, yet we know little of how they vary within species. Experimental conditions demonstrate that intraspecific plant root traits respond strongly to variation in the environment; however, it is unclear when these responses can be characterized as evolution in response to selective pressures of climate change over many generations. Sky Islands are model, natural climate relict ecosystems to examine climate-change driven evolution. Utilizing a common garden with replicate genotypes of Populus angustifolia (Narrowleaf cottonwood) from six Sky Island (SI) populations and nine adjacent Mountain Chain (MC) populations across three genetic provenances, we hypothesized that SI root traits have diverged due to historical isolation in warmer, drier climates. When grown in common conditions, populations originating on SI’s showed convergent evolution across three distinct genetic provenances, which was characterized by 44.16% decreased total root length, 42.64% decreased average root volume, 43.31% decreased root surface area, and significantly less root trait variation, relative to adjacent mountain chains. Convergent evolution of root traits from trees originating on SI’s is correlated with changes in mean annual precipitation and potential evapotranspiration in the field over the past ~ 125 years. These results demonstrate a consistent pattern in root trait evolution at the landscape scale and the role of climate on the evolution of root traits in a genetic and geographic context relevant to climate change.
Similar content being viewed by others
Climate-driven convergent evolution in riparian ecosystems on sky islands
A systematic review to identify target genes that modulate root system architecture in response to abiotic stress
Root traits explain plant species distributions along climatic gradients yet challenge the nature of ecological trade-offs
Data availability
The data that support the findings of this study are available in figshare at https//doi.org/. The long-term climate data, including mean annual temperature and mean annual precipitation from 1895 to 2022, were derived from the Oregon State University PRISM Weather Data, available in the public domain: https//doi.org/10.6084/m9.figshare.29374034. The modern climate data, including potential evapotranspiration, mean annual temperature, and mean annual precipitation from 2021, were derived from the following resources, available in the public domain: ClimateNA https://climatena.ca/mapversion (for mean annual temperature and mean annual precipitation); CGIAR-CSI Global Aridity Index and Potential Evapotranspiration (ET0) Database v3 (for potential evapotranspiration).
References
Garnier, E. et al. Consistency of species ranking based on functional leaf traits. New Phytol. 152, 69–83 (2001).
McGill, B., Enquist, B., Weiher, E. & Westoby, M. Rebuilding community ecology from functional traits. Trends in Ecol. And Evol. 21, 178–185 (2006).
Laughlin, D. C. et al. Root traits explain plant species distributions along climatic gradients yet challenge the nature of ecological trade-offs. Nat. Ecol. Evolut. 5, 1123–1134 (2021).
Violle, C. et al. The return of the variance: intraspecific variability in community ecology. Trends Ecol. Evol. 27, 244–252 (2012).
Siefert, A. et al. A global meta-analysis of the relative extent of intraspecific trait variation in plant communities. Ecol. Lett. 18, 1406–1419 (2015).
Read, Q. D., Henning, J. A. & Sanders, N. J. Intraspecific variation in traits reduces ability of trait-based models to predict community structure. J. Veg. Sci. 28, 1070–1081 (2017).
Des Roches, S. et al. The ecological importance of intraspecific variation. Nat. Ecol. Evol. 2, 57–64 (2017).
Gazol, A., Fajardo, A. & Camarero, J. J. Contributions of intraspecific variation to drought tolerance in trees. Curr. For. Rep. 9, 461–472 (2023).
Kooyers, N. J., Greenlee, A. B., Colicchio, J. M., Oh, M. & Blackman, B. K. Replicate altitudinal clines reveal that evolutionary flexibility underlies adaptation to drought stress in annual Mimulus guttatus. New Phytol. 206, 152–165 (2015).
Norby, R. J., Ledford, J., Reilly, C. D., Miller, N. E. & O’Neill, E. G. Fine-root production dominates response of a deciduous forest to atmospheric CO2 enrichment. Proc. Natl. Acad. Sci. 101, 9689–9693 (2004).
Pregitzer, K. S., Burton, A. J., King, J. S. & Zak, D. R. Soil respiration, root biomass, and root turnover following long-term exposure of northern forests to elevated atmospheric CO2 and tropospheric O3. New Phytol. 180, 153–161 (2008).
Iversen, C. M., Ledford, J. & Norby, R. J. CO2 enrichment increases carbon and nitrogen input from fine roots in a deciduous forest. New Phytol. 179, 837–847 (2008).
Pritchard, S. G. et al. Fine root dynamics in a loblolly pine forest are influenced by free-air-CO2 -enrichment: a six-year-minirhizotron study. Glob. Change Biol. 14, 588–602 (2008).
Iversen, C. M. Digging deeper: fine-root responses to rising atmospheric CO2 concentration in forested ecosystems. New Phytol. 186, 346–357 (2010).
Norby, R. J. & Zak, D. R. Ecological lessons from free-air CO2 enrichment (FACE) experiments. Annu. Rev. Ecol. Evol. Syst. 42, 181–203 (2011).
Nie, M., Lu, M., Bell, J., Raut, S. & Pendall, E. Altered root traits due to elevated CO2: a meta-analysis. Glob. Ecol. Biogeogr. 22, 1095–1105 (2013).
Mueller, K. E. et al. Root responses to elevated CO2, warming and irrigation in a semi-arid grassland: Integrating biomass, length and life span in a 5-year field experiment. J. Ecol. 106, 2176–2189 (2018).
Zhou, L. et al. Responses of biomass allocation to multi-factor global change: A global synthesis. Agric. Ecosyst. Environ. 304, 107115 (2020).
Wang, J. et al. Fine-root functional trait responses to experimental warming: a global meta-analysis. New Phytol. 230, 1856–1867 (2021).
Brunner, I., Herzog, C., Dawes, M. A., Arend, M. & Sperisen, C. How tree roots respond to drought. Front. Plant Sci. 6, 547 (2015).
Zhou, G. et al. Drought-induced changes in root biomass largely result from altered root morphological traits: Evidence from a synthesis of global field trials. Plant Cell Environ. 41, 2589–2599 (2018).
Poorter, H. et al. Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control. New Phytol. 193, 30–50 (2012).
Zhang, B. et al. Plants alter their vertical root distribution rather than biomass allocation in response to changing precipitation. Ecology 100, e02828 (2019).
Wang, P., Huang, K. & Hu, S. Distinct fine-root responses to precipitation changes in herbaceous and woody plants: a meta-analysis. New Phytol. 225, 1491–1499 (2020).
Endler, J. A. Geographic Variation, Speciation, and Clines (Princeton University Press, 1977).
Storfer, A. Gene flow and endangered species translocations: a topic revisited. Biol. Conserv. 87, 173–180 (1999).
Foster, S. A., McKinnon, G. E., Steane, D. A., Potts, B. M. & Vaillancourt, R. E. Parallel evolution of dwarf ecotypes in the forest tree Eucalyptus globulus. New Phytol. 175, 370–380 (2007).
Read, Q. D., Moorhead, L. C., Swenson, N. G., Bailey, J. K. & Sanders, N. J. Convergent effects of elevation on functional leaf traits within and among species. Funct. Ecol. 28, 37–45 (2014).
Zadworny, M., McCormack, M. L., Mucha, J., Reich, P. B. & Oleksyn, J. Scots pine fine roots adjust along a 2000-km latitudinal climatic gradient. New Phytol. 212, 389–399 (2016).
Barber N.H., Jackson, D.W. Natural selection in action in Eucalyptus. Nature (1957).
Kremer, A., Potts, B. M. & Delzon, S. Genetic divergence in forest trees: understanding the consequences of climate change. Funct. Ecol. 28, 22–36 (2014).
Keller, S. R., Chhatre, V. E. & Fitzpatrick, M. C. Influence of range position on locally adaptive gene–environment associations in Populus flowering time genes. J. Hered. 109, 47–58 (2018).
Pfennigwerth, A. A., Bailey, J. K. & Schweitzer, J. A. Trait variation along elevation gradients in a dominant woody shrub is population-specific and driven by plasticity. AoB Plants 9, plx027 (2017).
Van Nuland, M. E., Ware, I. M., Bailey, J. K. & Schweitzer, J. A. Ecosystem feedbacks contribute to geographic variation in plant–soil eco-evolutionary dynamics across a fertility gradient. Funct. Ecol. 33, 95–106 (2019).
Bayliss, S. L. J., Mueller, L. O., Ware, I. M., Schweitzer, J. A. & Bailey, J. K. Plant genetic variation drives geographic differences in atmosphere–plant–ecosystem feedbacks. Plant-Environ. Interact. 1, 166–180 (2020).
Fischer, D. G. et al. Do high-tannin leaves require more roots?. Oecologia 149, 668–675 (2006).
Jung, J. K. H. & McCouch, S. Getting to the roots of it: Genetic and hormonal control of root architecture. Front. Plant Sci. 4, 186 (2013).
Lynch, J. P. Harnessing root architecture to address global challenges. Plant J. 109, 415–431 (2022).
Nakahata, R., Azuma, W. A., Tanabe, T., Kawai, K. & Hiura, T. Genotypic variations appear in fine root morphological traits of Cryptomeria japonica trees grown in a common garden. Ecol. Res. 39, 717–729 (2024).
Perret, D. L., Evans, M. E. K. & Sax, D. F. A species’ response to spatial climatic variation does not predict its response to climate change. Proc. Natl. Acad. Sci. 121, 2304404124 (2024).
Hendry, A. P. A critique for eco-evolutionary dynamics. Funct. Ecol. 33, 84–94 (2019).
Lowry, D. B., Popovic, D., Brennan, D. J. & Holeski, L. M. Mechanisms of a locally adaptive shift in allocation among growth, reproduction, and herbivore resistance in Mimulus guttatus. Evolution 73, 1168–1181 (2019).
Love, S. J., Schweitzer, J. A. & Bailey, J. K. Climate-driven convergent evolution in riparian ecosystems on Sky Islands. Sci. Rep. 13, 2817 (2023).
Love, S. J., Schweitzer, J. A., Woolbright, S. A. & Bailey, J. K. Sky islands are a global tool for predicting the ecological and evolutionary consequences of climate change. Annu. Rev. Ecol. Evol. Syst. 54, 219–236 (2023).
Evans, L. M., Allan, G. J., Meneses, N., Max, T. L. & Whitham, T. G. Herbivore host-associated genetic differentiation depends on the scale of plant genetic variation examined. Evol. Ecol. 27, 65–81 (2013).
DeBano, L. H. et al. Biodiversity and management of the Madrean Archipelago: The Sky Islands of southwestern United States and northwestern Mexico: 1994 September 19–23; Tucson, AZ. Gen Tech Rep RM-GTR-264. Fort Collins, CO: US Dep Agric For Serv, Rocky Mt For Range Exp Stn. 669 (1995).
Ware, I. M. et al. Climate-driven reduction of genetic variation in plant phenology alters soil communities and nutrient pools. Glob. Change Biol. 25, 1514–1528 (2019).
PRISM Climate Group, Oregon State University, https://prism.oregonstate.edu, data created 4 Feb 2014, accessed 16 Jan 2024.
Zomer, R. J., Bossio, D. A., Trabucco, A., Yuanjie, L., Gupta, D. C, & Singh, V.P. Trees and water: Smallholder agroforestry on irrigated lands in Northern India. Colombo. International Water Management Institute. 45 (2007).
Zomer, R. J., Trabucco, A., Bossio, D. A., van Straaten, O. & Verchot, L. V. Climate change mitigation: a spatial analysis of global land suitability for clean development mechanism afforestation and reforestation. Agric. Ecosyst. Environ. 126, 67–80 (2008).
Roach, D. A. & Wulff, R. D. Maternal effects in plants. Annu. Rev. Ecol. Evol. Syst. 18, 209–235 (1987).
Regent Instruments Inc. WinRHIZO Pro: Analysis of washed roots and arabidopsis seedlings 2009b (1991).
Bardgett, R. D., Mommer, L. & De Vries, F. T. Going underground: root traits as drivers of ecosystem processes. Trends Ecol. Evol. 29, 692–699 (2014).
Freschet, G. T. et al. A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements. New Phytol. 232, 973–1122 (2021).
Wang, H., Inukai, Y. & Yamauchi, A. Root development and nutrient uptake. Crit. Rev. Plant Sci. 25, 279–301 (2006).
McCormack, M. L., Adams, T. S., Smithwick, E. A. H. & Eissenstat, D. M. Predicting fine root lifespan from plant functional traits in temperate trees. New Phytol. 195, 823–831 (2012).
Atkinson, D 2000 Root characteristics: why and what to measure. In Root Methods. 1–29 (Springer, 2000).
Kong, D. et al. Leading dimensions in absorptive root trait variation across 96 subtropical forest species. New Phytol. 203, 863–872 (2014).
Chen, W. et al. Root morphology and mycorrhizal symbioses together shape nutrient foraging strategies of temperate trees. Proc. Natl. Acad. Sci. 113, 8741–8746 (2016).
Bergmann, J. et al. The fungal collaboration gradient dominates the root economics space in plants. Sci. Adv. 6, 3756 (2020).
Wang, G. et al. Shifts in ectomycorrhizal exploration types complement root traits in nutrient foraging of alpine coniferous forests along an elevation gradient. Plant Soil (2025).
R Development Core Team. R: a language and environment for statistical computing, v.4.4.1. Vienna. R foundation for Statistical Computing (2024).
Kattge, J. et al. TRY – a global database of plant traits. Glob. Change Biol. 17, 2905–2935 (2011).
Brito-Morales, I. et al. Climate velocity can inform conservation in a warming world. Trends Ecol. Evol. 33, 441–457 (2018).
Shishkova, S. et al. Determinate primary root growth as an adaptation to aridity in Cactaceae: towards an understanding of the evolution and genetic control of the trait. Ann. Bot. 112, 239–252 (2013).
Calleja-Cabrera, J., Boter, M., Oñate-Sánchez, L. & Pernas, M. Root growth adaptation to climate change in crops. Front. Plant Sci. 11, 544 (2020).
Acknowledgements
We would like to thank Jeff Martin and greenhouse staff as well as Emma Kruse for supporting the research and writing process. We would also like to give special thanks to Sarah Love for assisting in the identification and collection of cuttings from the field.
Funding
LEP was supported by the University of Tennessee – Oak Ridge Innovation Institute through the Graduate Advancement, Training and Education (GATE) Fellowship. LMY was supported by the Center for Bioenergy Innovation (CBI), which is a U.S. Department of Energy Bioenergy Research Center supported by the Office of Biological and Environmental Research in the DOE Office of Science. This manuscript has been authored in part by UT-Battelle, LLC that manages Oak Ridge National Laboratory under contract DE-AC05-00OR22725 with the US Department of Energy (DOE). The publisher acknowledges the US government license to provide public access under the DOE Public Access Plan (https://energy.gov/downloads/doe-public-access-plan).
Author information
Authors and Affiliations
Contributions
LEP, JKB, and JAS planned and designed the research. LMY informed data collection methodology and LEP performed data collection and experiments. LEP analyzed data and wrote the manuscript, while LMY, JKB, and JAS provided essential feedback and suggestions.
Corresponding author
Ethics declarations
Competing interests
The authors declare no competing interests.
Additional information
Publisher’s note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Supplementary Information
Below is the link to the electronic supplementary material.
Supplementary Material 1
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
Reprints and permissions
About this article
Cite this article
Politano, L.E., York, L.M., Bailey, J.K. et al. Climate change drives convergent evolution of root traits on Sky Island climate relicts.
Sci Rep (2026). https://doi.org/10.1038/s41598-025-31134-7
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41598-025-31134-7
Keywords
- Convergent evolution
- Local adaptation
Populus
- Root functional traits
- Sky Islands
Source: Ecology - nature.com
