Environmental conditions are dynamic, and plants respond to those dynamics on multiple time scales. Disequilibrium occurs when a response occurs more slowly than the driving environmental changes. We review evidence regarding disequilibrium in plant distributions, including their responses to paleoclimate changes, recent climate change and new species introductions. There is strong evidence that plant species distributions are often in some disequilibrium with their environmental conditions.This disequilibrium poses a challenge when projecting future species distributions using species distribution models (SDMs). Classically, SDMs assume that the set of species occurrences is an unbiased sample of the suitable environmental conditions. However, a species in disequilibrium with the environment may have higher‐than‐expected occurrence probabilities (e.g. due to extinction debts) or lower‐than‐expected occurrence probabilities (e.g. due to dispersal limitation) in different areas. If unaccounted for, this will lead to biased estimates of the environmental suitability.We review methods for avoiding such biases in SDMs, ranging from simple thinning of the occurrence dataset to complex dynamic and process‐based models. Such models require large data inputs, natural history knowledge and technical expertise, so implementing them can be challenging. Despite this, we advocate for their increased use, since process‐based models provide the best potential to account for biases in model training data and to then represent the dynamics of species occupancy as ranges shift.Synthesis. Occurrence records for a species are often in disequilibrium with climate. SDMs trained on such data will produce biased estimates of a species' niche unless this disequilibrium is addressed in the modelling. A range of tools, spanning a wide gradient of complexity and realism, can resolve this bias. 
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                            Incorporating plant phenological responses into species distribution models reduces estimates of future species loss and turnover
                        
                    
    
            Summary Anthropogenetic climate change has caused range shifts among many species. Species distribution models (SDMs) are used to predict how species ranges may change in the future. However, most SDMs rarely consider how climate‐sensitive traits, such as phenology, which affect individuals' demography and fitness, may influence species' ranges.Using > 120 000 herbarium specimens representing 360 plant species distributed across the eastern United States, we developed a novel ‘phenology‐informed’ SDM that integrates phenological responses to changing climates. We compared the ranges of each species forecast by the phenology‐informed SDM with those from conventional SDMs. We further validated the modeling approach using hindcasting.When examining the range changes of all species, our phenology‐informed SDMs forecast less species loss and turnover under climate change than conventional SDMs. These results suggest that dynamic phenological responses of species may help them adjust their ecological niches and persist in their habitats as the climate changes.Plant phenology can modulate species' responses to climate change, mitigating its negative effects on species persistence. Further application of our framework will contribute to a generalized understanding of how traits affect species distributions along environmental gradients and facilitate the use of trait‐based SDMs across spatial and taxonomic scales. 
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                            - PAR ID:
- 10576933
- Publisher / Repository:
- Wiley-Blackwell
- Date Published:
- Journal Name:
- New Phytologist
- Volume:
- 242
- Issue:
- 5
- ISSN:
- 0028-646X
- Format(s):
- Medium: X Size: p. 2338-2352
- Size(s):
- p. 2338-2352
- Sponsoring Org:
- National Science Foundation
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