Abstract
Resource availability is a primary factor predicting population performance. Synchrony between resource availability and consumer requirements plays a critical role in reproduction, and mismatches in the timing of resource availability and consumer requirements can have negative implications for reproductive success. Our objective was to determine when mass of juvenile and yearling white-tailed deer (Odocoileus virginianus) in semiarid rangelands is most negatively affected by limited forage availability. Thus, we determined the biological period when rainfall, a primary driver of resource availability, was most predictive of juvenile and yearling mass. Over 12 years, we captured 1,123 juveniles and yearlings across five distinct populations. We linked georeferenced capture records to rainfall data from biological seasons hypothesized to affect juvenile and yearling mass. We found that rainfall during the early growing season exhibited the strongest effect on mass. The resource pulse associated with early growing season rainfall is likely used to fuel fetal development during the critical final trimester of gestation. As environmental change continues to exacerbate the potential for mismatches in resource availability and consumer requirements to occur, it is important to identify when limited forage availability may most negatively affect reproduction to inform species conservation and support long-term sustainability.
Data availability
The data used in this manuscript is provided via Zenodo at the following link: https://doi.org/10.5281/zenodo.12611417.
References
Blomberg, E. J., Sedinger, J. S., Gibson, D., Coates, P. S. & Casazza, M. L. Carryover Effects and Climatic Conditions Influence the Postfledging Survival of Greater Sage-grouse. Ecol. Evol. 4, 4488–4499 (2014).
Lopp, J. & Sammul, M. The Impact of Timing of Resource Availability on Clonal Propagation of Species with Different Growth Forms. Folia Geobot. 52, 411–422 (2017).
Mazzamuto, M. V. et al. Timing of Resource Availability Drives Divergent Social Systems and Home Range Dynamics in Ecologically Similar Tree Squirrels. Front. Ecol. Evol. 8, 174 (2020).
Durant, J. M., Hjermann, D. Ø., Ottersen, G. & Stenseth, N. C. Climate and the Match or Mismatch Between Predator Requirements and Resource Availability. Clim. Res. 33, 271–283 (2007).
Neumann, W. et al. Divergence in Parturition Timing and Vegetation Onset in a Large Herbivore—Differences Along a Latitudinal Gradient. Biol. Lett. 16, 20200044 (2020).
Peláez, M., Gaillard, J. M., Bollmann, K., Heurich, M. & Rehnus, M. Large-scale Variation in Birth Timing and Synchrony of a Large Herbivore Along the Latitudinal and Altitudinal Gradients. J. Anim. Ecol. 89, 1906–1917 (2020).
Visser, M. E. & Gienapp, P. Evolutionary and Demographic Consequences of Phenological Mismatches. Nat. Ecol. Evol. 3, 879–885 (2019).
Kerby, J. T., Wilmers, C. C., & Post, E. Climate Change Phenology, and the Nature of Consumer–resource Interactions: Advancing the Match/mismatch Hypothesis. in Trait-Mediated Indirect Interactions: Ecological and Evolutionary Perspectives (eds. Ohgushi, T., Schmitz, O. & Holt, R. D.) 508–525. (Cambridge University Press, 2012).
Post, E. & Forchhammer, M. C. Climate Change Reduces Reproductive Success of an Arctic Herbivore through Trophic Mismatch. Philos. Trans. R Soc. B Biol. Sci. 363, 2367–2373 (2008).
Rehnus, M., Peláez, M. & Bollmann, K. Advancing Plant Phenology Causes an Increasing Trophic Mismatch in an Income Breeder Across a Wide Elevational Range. Ecosphere 11, 03144 (2020).
Youngflesh, C. et al. Demographic Consequences of Phenological Asynchrony for North American Songbirds. Proc. Natl. Acad. Sci. 120, 2221961120 (2023).
Lincoln, G. A Brief History of Circannual Time. J. Neuroendocrinol. 31, e12694 (2019).
Tyler, N. J. C., Gregorini, P., Parker, K. L. & Hazlerigg, D. G. Animal Responses to Environmental Variation: Physiological Mechanisms in Ecological Models of Performance in Deer (Cervidae). Anim. Prod. Sci. 60, 1248–1270 (2020).
Apollonio, M. & Chirichella, R. Deer and Climate Change: Impacts and Perspectives (Animal Production Science, 2023).
Williams, C. T. et al. Seasonal Reproductive Tactics: Annual Timing and the Capital-to-Income Breeder Continuum. Philos. Trans. R Soc. B Biol. Sci. 372, 20160250 (2017).
Zerbe, P. et al. Reproductive Seasonality in Captive Wild Ruminants: Implications for Biogeographical Adaptation, Photoperiodic Control, and Life History. Biol. Rev. 87, 965–990 (2012).
Gaillard, J. M. et al. How Does Climate Change Influence Demographic Processes of Widespread Species? Lessons from the Comparative Analysis of Contrasted Populations of Roe Deer. Ecol. Lett. 16, 48–57 (2013).
Hill, A. P., Nolan, C. J., Hemes, K. S., Cambron, T. W. & Field, C. B. Low-Elevation Conifers in California’s Sierra Nevada Are Out of Equilibrium with Climate. PNAS Nexus. 2, 004 (2023).
Parmesan, C. Ecological and Evolutionary Responses to Recent Climate Change. Annu. Rev. Ecol. Evol. Syst. 37, 637–669 (2006).
Marshal, J. P., Krausman, P. R., & Bleich, V. C. Rainfall, Temperature, and Forage Dynamics Affect Nutritional Quality of Desert Mule Deer Forage. Rangel. Ecol. Manag. 58, 360–365 (2005).
Plaisir, C. A., King, W. J., Forsyth, D. M. & Festa-Bianchet, M. Effects of Rainfall, Forage Biomass, and Population Density, on Survival and Growth of Juvenile Kangaroos. J. Mammal. 103, 491–502 (2022).
Walter, J. et al. Increased Rainfall Variability Reduces Biomass and Forage Quality of Temperate Grassland Largely Independent of Mowing Frequency. Agric. Ecosyst. Environ. 148, 1–10 (2012).
Ortega, A. C., Aikens, E. O., Merkle, J. A., Monteith, K. L. & Kauffman, M. J. Migrating Mule Deer Compensate en Route for Phenological Mismatches. Nat. Commun. 14, 2008. (2023).
Aikens, E. O. et al. Migration Distance and Maternal Resource Allocation Determine Timing of Birth in a Large Herbivore. Ecology 102, 03334 (2021).
Rachlow, J. L. & Bowyer, R. T. Variability in Maternal Behavior by Dall’s Sheep: Environmental Tracking or Adaptive Strategy? J. Mammal. 75, 328–337 (1994).
Plard, F. et al. Mismatch Between Birth Date and Vegetation Phenology Slows the Demography of Roe Deer. PLoS Biol. 12, 1001828 (2014).
Renaud, L. A., Festa-Bianchet, M. & Pelletier, F. Testing the Match–Mismatch Hypothesis in Bighorn Sheep in the Context of Climate Change. Glob Change Biol. 28, 21–32 (2021).
Servanty, S., Gaillard, J. M., Toïgo, C., Brandt, S. & Baubet, E. Pulsed Resources and Climate-induced Variation in the Reproductive Traits of Wild Boar under High Hunting Pressure. J. Anim. Ecol. 78, 1278–1290 (2009).
Tveraa, T., Stien, A., Bårdsen, B., & Fauchald, P. Population Densities, Vegetation Green-Up, and Plant Productivity: Impacts on Reproductive Success and Juvenile Body Mass in Reindeer. PloS One. 8, 56450 (2013).
Cook, J. G. et al. Effects of Summer-Autumn Nutrition and Parturition Date on Reproduction and Survival of Elk. Wildl. Monogr. 155, 1–61 (2004).
Strickland, B. K., Demarais, S. & Gerard, P. D. Variation in Mass and Lactation Among Cohorts of White-tailed Deer Odocoileus virginianus. Wildl. Biol. 14, 263–271 (2008).
DeYoung, C. A. et al. Density Dependence in Deer Populations: Relevance for Management in Variable Environments (Hewitt Wildlife Science, 2007).
Folks, D. J. et al. Drought but not Population Density Influences Dietary Niche Breadth in White-tailed Deer in a Semiarid Environment. Ecosphere 5, 1–15 (2014).
Fulbright, T. E. et al. Forb Standing Crop Response to Grazing and Precipitation. Rangel. Ecol. Manag. 79, 175–185 (2021).
Kharouba, H. M. & Wolkovich, E. M. Lack of Evidence for the Match-Mismatch Hypothesis Across Terrestrial Trophic Interactions. Ecol. Lett. 26, 955–964 (2023).
IPCC. IPCC, 2023: Climate Change 2023: Synthesis Report. Contribution of Working Groups I, II and III to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change [Core Writing Team, H. Lee and J. Romero (Eds.)]. https://www.ipcc.ch/report/ar6/syr/https://doi.org/10.59327/IPCC/AR6-9789291691647 (IPCC, 2023).
Gaillard, J. M., Festa-Bianchet, M. & Yoccoz, N. G. Population Dynamics of Large Herbivores: Variable Recruitment with Constant Adult Survival. Trends Ecol. Evol. 13, 58–63 (1998).
Hossain, M. L. & Beierkuhnlein, C. Enhanced Aboveground Biomass by Increased Precipitation in a Central European Grassland. Ecol. Process. 7, 1–13 (2018).
La Pierre, K. J., Blumenthal, D. M., Brown, C. S., Klein, J. A. & Smith, M. D. Drivers of Variation in Aboveground Net Primary Productivity and Plant Community Composition Differ Across a Broad Precipitation Gradient. Ecosystems 19, 521–533 (2016).
Ginnett, T. F. & Young, E. B. Stochastic Recruitment in White-tailed Deer Along an Environmental Gradient. J. Wildl. Manag. 64, 713–720 (2000).
Michel, E. S., Jenks, J. A., Kaskie, K. D., Klaver, R. W. & Jensen, W. F. Weather and Landscape Factors Affect White-tailed Deer Neonate Survival at Ecologically Important Life Stages in the Northern Great Plains. PLoS One. 13, 0195247 (2018).
Garel, M. et al. Population Abundance and Early Spring Conditions Determine Variation in Body Mass of Juvenile Chamois. J. Mammal. 92, 1112–1117 (2011).
Gaillard, J. M. et al. Early Survival in Roe Deer: Causes and Consequences of Cohort Variation in Two Contrasted Populations. Oecologia 112, 502–513 (1997).
Andersen, R., Gaillard, J. M., Linnell, J. D. C. & Duncan, P. Factors Affecting Maternal Care in an Income Breeder, the European Roe Deer. J. Anim. Ecol. 69, 672–682 (2000).
Michel, E. S., Demarais, S., Strickland, B. K., Belant, J. L. & Castle, L. E. Body Mass Influences Maternal Allocation More Than Parity Status for a Long-lived Cervid Mother. J. Mammal. 100, 1459–1465 (2019).
Thorne, E. T., Dean, R. E. & Hepworth, W. G. Nutrition During Gestation in Relation to Successful Reproduction in Elk. J. Wildl. Manag. 40, 330–335 (1976).
Aubin, G. R. et al. Survival of White-tailed Deer Fawns on Marine Corps Base Quantico. J. Wildl. Manag. 86, 22180 (2022).
Heffelfinger, L. J. et al. Timing of Precipitation in an Arid Environment: Effects on Population Performance of a Large Herbivore. Ecol. Evol. 8, 3354–3366 (2018).
Hediger, J. A. et al. Physiological Carry-over Effects of Variable Precipitation are Mediated by Reproductive Status in a Long-lived Ungulate. Conserv. Physiol. 12, (2024).
Douhard, M. et al. Variation in Adult Body Mass of Roe Deer: Early Environmental Conditions Influence Early and Late Body Growth of Females. Ecology 94, 1805–1814 (2013).
Michel, E. S. et al. Improved Nutrition Cues Switch from Efficiency to Luxury Phenotypes for a Long-lived Ungulate. Ecol. Evol. 6, 7276–7285 (2016).
Monteith, K. L., Schmitz, L. E., Jenks, J. A., Delger, J. A. & Bowyer, R. T. Growth of Male White-Tailed Deer: Consequences of Maternal Effects. J. Mammal. 90, 651–660 (2009).
Pettorelli, N. et al. Variations in Adult Body Mass in Roe Deer: The Effects of Population Density at Birth and of Habitat Quality. Proc. R Soc. Lond. B Biol. Sci. 269, 747–753 (2002).
Solberg, E. J., Loison, A., Gaillard, J. M. & Heim, M. Lasting Effects of Conditions at Birth on Moose Body Mass. Ecography 27, 677–687 (2004).
Hewison, A. M. et al. Reproductive Constraints, Not Environmental Conditions, Shape the Ontogeny of Sex-Specific Mass–Size Allometry in Roe Deer. Oikos 120, 1217–1226 (2011).
Foley, A. M. et al. Fine-scale Phenotypic Variation of a Large Herbivore in a Pulsed Environment. Ecosphere 15, e4921 (2024).
Plard, F. et al. The Influence of Birth Date via Body Mass on Individual Fitness in a Long-lived Mammal. Ecology 96, 1516–1528 (2015).
Moen, A. N. Wildlife Ecology: An Analytical Approach (W.H. Freeman, 1973).
PRISM Group at Oregon State University. https://prism.oregonstate.edu/.
Crider, B. L. et al. Influence of White-tailed Deer Population Density on Vegetation Standing Crop in a Semiarid Environment. J. Wildl. Manag. 79, 413–424 (2015).
DeYoung, C. A. et al. Linking White-Tailed Deer Density, Nutrition, and Vegetation in a Stochastic Environment. Wildl. Monogr. 202, 1–63 (2019).
Gann, W. J. et al. Vegetation Response to White-tailed Deer Density and Enhanced Nutrition. Wildl. Monogr. 202, 1–63 (2019).
Gann, K. R. et al. Deer Density Effects on White-tailed Deer Diets and Foraging Behavior Under Natural Nutrition. In Linking White-tailed Deer Density, Nutrition, and Vegetation in a Stochastic Environment 202 19–26 (2019).
Cook, R. S., White, M., Trainer, D. O. & Glazener, W. C. Mortality of Young White-Tailed Deer Fawns in South Texas. J. Wildl. Manag. 35, 47–56 (1971).
Krausman, P. R., Hervert, J. J. & Ordway, L. L. Capturing Deer and Mountain Sheep with a Net-Gun. Wildl. Soc. Bull. 13, 71–73 (1985).
Foley, A. M. et al. Accuracies and Biases of Ageing White-tailed Deer in Semiarid Environments. Wildl. Res. 49, 237–249 (2021).
Hirth, D. H. Mother-Young Behavior in White-tailed Deer, Odocoileus virginianus. Southwest. Nat. 30, 297–302 (1985).
Therrien, J. F., Côté, S. D., Festa-Bianchet, M. & Ouellet, J. P. Maternal Care in White-tailed Deer: Trade-off Between Maintenance and Reproduction Under Food Restriction. Anim. Behav. 75, 235–243 (2008).
Verme, L. J. Reproductive Patterns of White-tailed Deer Related to Nutritional Plane. J. Wildl. Manag. 33, 881–887 (1969).
TPWD. The Rut in White-tailed Deer. https://tpwd.texas.gov/huntwild/hunt/planning/rut_whitetailed_deer/.
Ortega-S, J. A., Lukefahr, S. D. & Bryant, F. C. Optimum Stocking Rate, Monitoring, and Flexibility: Key Components of Successful Grazing Management Programs. Rangelands 35, 22–27 (2013).
Akaike, H. Maximum Likelihood Identification of Gaussian Autoregressive Moving Average Models. Biometrika 60, 255–265 (1973).
Burnham, K. P. & Anderson, D. R. Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach (Springer, 2002).
Bolker, B. M. et al. Generalized Linear Mixed Models: A Practical Guide for Ecology and Evolution. Trends Ecol. Evol. 24, 127–135 (2009).
Bates, D. M. lme4: Mixed-effects Modeling with R. (2010).
Acknowledgements
We thank the East Foundation and the Patton Center for Deer Research for their funding and support during this research. Additional support was provided by the Hixon Fellowship, the Barrientos Scholarship, and the Houston Safari Club Dan L. Duncan Scholarship. J. A. Ortega-Sanchez, K. Gann, M. Rice, and S. Rankins coordinated capture activities during 2011–2020, assisted by numerous student volunteers from Texas A&M University-Kingsville, Southwest Texas Junior College, Stephen F. Austin State University, Sul Ross State University, Tarleton State University, Texas A&M University, Texas State University, and Texas Tech University. A. Montalvo, A. Woodard, J. Haynes, A. Lopez, R. Douglas, S. Vasquez, and M. Robinson of the East Foundation were instrumental in field logistics and activities. A. Tanner and H. Abernathy reviewed earlier drafts of the paper. All clipart was obtained from the University of Maryland Center for Environmental Science Integration and Application Network. B. Green assisted in editing conceptual figures. This is #25-114 of the Caesar Kleberg Wildlife Research Institute and #096 of the East Foundation.
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The authors gratefully acknowledge the financial support provided by the East Foundation and the Patton Center for Deer Research.
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M.L.H. and M.J.C. conceived the ideas of this manuscript. All authors collected the data. M.L.H. developed the analytical framework and analyzed the data. M.L.H. developed and designed all figures (with illustrations provided by the University of Maryland Center for Environmental Science Integration and Application Network and Breanna Green). M.L.H. led the writing of the original manuscript. All authors contributed critically to all drafts and gave final approval for publication.
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Hopper, M.L., Spencer, B.D., DeYoung, R.W. et al. Timing of rainfall influences juvenile and yearling mass of a long-lived herbivore in a semiarid environment.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-40861-4
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DOI: https://doi.org/10.1038/s41598-026-40861-4
Keywords
- Climate change
- Drought
- Environmental stochasticity
Odocoileus virginianus
- Phenological mismatch
- White-tailed deer
Source: Ecology - nature.com

