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| Schematic demonstrating how metapopulation dynamics can lead to range limits. A metapopulation approach to range dynamics hypothesizes that range limits are determined by colonization (c, blue lines) and extinction (m, red lines) probabilities as they vary along an environmental gradient 9. a, c and m for Acer saccharum vary with temperature, resulting in a latitudinal gradient. More generally, we describe c and m as functions of the environment: c(E) and m(E). The theory predicts that the species will be present at equilibrium (shaded region) where c(E) > m(E); therefore the temperature at which c(E) = m(E) determines the latitude of the range limits. b, A simulated 2.5 °C of warming produces a northward shift in the suitable range (grey and blue shading). However, over short time scales, slow extinction probabilities will cause the species to persist where m(E) > c(E) (red shading) until all occupied populations have gone extinct. A similar lag in colonization ability will result in the species being absent in many areas where c(E) > m(E) (blue shading). Thin lines show 90% credible intervals around mean c and m probabilities. 

| Schematic demonstrating how metapopulation dynamics can lead to range limits. A metapopulation approach to range dynamics hypothesizes that range limits are determined by colonization (c, blue lines) and extinction (m, red lines) probabilities as they vary along an environmental gradient 9. a, c and m for Acer saccharum vary with temperature, resulting in a latitudinal gradient. More generally, we describe c and m as functions of the environment: c(E) and m(E). The theory predicts that the species will be present at equilibrium (shaded region) where c(E) > m(E); therefore the temperature at which c(E) = m(E) determines the latitude of the range limits. b, A simulated 2.5 °C of warming produces a northward shift in the suitable range (grey and blue shading). However, over short time scales, slow extinction probabilities will cause the species to persist where m(E) > c(E) (red shading) until all occupied populations have gone extinct. A similar lag in colonization ability will result in the species being absent in many areas where c(E) > m(E) (blue shading). Thin lines show 90% credible intervals around mean c and m probabilities. 

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Global climate change is already having an impact on species ranges. For species with slow demography and limited dispersal, such as trees, lags between climate change and distribution shifts are likely to increase in the future. Such lags can be of critical importance to management and biodiversity of forests, because they can result in ‘extinctio...

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... Behind the fundamental thermal niche, deaths exceed births, known as 'extinction debt' (Tilman et al., 1994). Ahead of the niche, due to dispersal limitation, the lag between the location of the niche and the density of the population increases (Hurford et al., 2019), as does the 'colonization credit', the future potential range of the species (Talluto et al., 2017) (Figure 1a). For example, extinction debt and colonization credit are widespread and affect bird species in about half of the area of the entire US (Haddou et al., 2022). ...
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... If species' cold-edge range limits are strongly constrained by soil, then potential range shifts and biodiversity change under climate change would be much smaller than is often assumed. Recent studies have found that ongoing plant migration lags behind climate warming [15][16][17][18], a pattern potentially explained by nonclimatic constraints. As such, it is imperative to evaluate whether and how soil and other non-climate factors influence plant distributions along climatic gradients. ...
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... Negative CTI changes correspond to an increase in cold-affinity species (i.e., borealization) and/or a decrease of warm-affinity species (i.e., detropicalization). The reason why tropicalization may prevail (or not) over deborealization has been recently debated [32][33][34] . Assuming an ecophysiological equilibrium between habitat suitability and the occurrence of species, we can expect that species track temperature change equally 33,35 , hence tropicalization would be similar to deborealization. ...
... Assuming an ecophysiological equilibrium between habitat suitability and the occurrence of species, we can expect that species track temperature change equally 33,35 , hence tropicalization would be similar to deborealization. However, in species with slow demography and limited dispersal, lags between climate change and distribution shifts can result in 'extinction debts' 36 , where populations temporarily persist under unsuitable conditions, and 'colonization credits', where suitable locations are not occupied 32 . This latter pattern has been observed for instance in some studies on trees 32 and demersal fish 33 , whilst other studies found equally responsive shifts at both range boundaries in marine ectotherms 35 , and prevalence of the tropicalization in demersal fish 15 . ...
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... Importantly, colonization and extinction rates are heterogeneous in space, shifting the focus of the model from the regional scale to the local scale. Other processes of interest can also be easily incorporated; for example, competition with other species can be incorporated in the extinction term 114 , dispersal rates can influence colonization 115 , and habitat/niche dimensions can be made a part of the colonization and extinction functions 116,117 . ...
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... Differences in plant species richness and composition between post-agricultural secondary forests and both mature forests and forests regenerating after natural disturbance may persist for decades, or even centuries (Boucher et al., 2001;Flinn & Marks, 2007;De Frenne et al., 2011;Naaf & Kolk, 2015;Muscarella et al., 2016;Kolk et al., 2017;Rozendaal et al., 2019;Brubaker & Cosentino, 2021;Brunet et al., 2021). This 'colonisation credit' (Talluto et al., 2017) may be a major barrier to post-agricultural forests achieving their full potential in ecological integrity, ecosystem functioning and resilience (Conti & Díaz, 2013;Gamfeldt et al., 2013;van der Plas et al., 2016;Ratcliffe et al., 2017). Understanding the processes which constrain species accumulation is an important step in identifying management interventions to enhance the biodiversity gain and ecosystem function of post-agricultural forest successions (Bartha et al., 2003;Török & Helm, 2017). ...
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... Potential plant migration could be strongly impacted if sites that become climatically suitable have unsuitable soil (Lafleur et al., 2010;Brown & Vellend, 2014;Carteron et al., 2020;Greiser et al., 2020;Benning & Moeller, 2021a,b). If so, apparent migration lags (Fei et al., 2017;Talluto et al., 2017;Boisvert-Marsh et al., 2019) might be due in part to overestimates of how far suitable growing conditions have shifted. Potential future species richness in poleward sites might also be overestimated by climate-only models. ...
... Specifically, current soils in tundra and boreal forests, which are generally shallow, acidic and even frozen (permafrost) are likely to be unsuitable for many southern plant species (Lafleur et al., 2010). Our results also indicate that the observed migration lags in some plant species under climate change (Fei et al., 2017;Talluto et al., 2017;Rumpf et al., 2018;Boisvert-Marsh et al., 2019;Rubenstein et al., 2023) might be caused, at least partly, by unsuitable soil environments beyond leading range edges. While climate change and associated vegetation responses might alter some soil properties to make them more suitable for 'southern' species in the coming decades (e.g. the melting permafrost at tundra or reduced acidity following conifer-to-hardwood transitions), other soil properties, such as depth and texture, are likely to represent persistent barriers over much longer time frames. ...
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Questions Near‐ground temperatures can vary substantially over relatively short distances, enabling species with different temperature preferences and geographical distributions to co‐exist within a small area. In a forest landscape, the near‐ground temperatures may change due to management activities that alter forest density. As a result of such management activities, current species distributions and performances might not only be affected by current microclimates, but also by past conditions due to time‐lagged responses. Location Sweden. Methods We examined the effects of past and current microclimates on the distributions and performances of two northern, cold‐favoured, and two southern, warm‐favoured, plant species in 53 managed forest sites. Each pair was represented by one vascular plant and one bryophyte species. We used temperature logger data and predictions from microclimate models based on changes in basal area to relate patterns of occurrence, abundance, and reproduction to current and past microclimate. Results The two northern species were generally favoured by microclimates that were currently cold, characterised by later snowmelt and low accumulated heat over the growing season. In contrast, the two southern species were generally favoured by currently warm microclimates, characterised by high accumulated heat over the growing season. Species generally had higher abundance in sites with a preferred microclimate both in the past and present, and lower abundance than expected from current conditions, if the past microclimate had changed from warm to cold or vice versa, indicating time‐lags in abundance patterns of the species. Conclusions Our results show a potential importance of past and present microclimate heterogeneity for the co‐existence of species with different temperature preferences in the same landscape and highlight the possibility to manage microclimates to mitigate climate change impacts on forest biodiversity.
... An outstanding question in studying the responses of forest reproduction to climate change is whether there are individual traits that can explain the variation in masting breakdown. Unlike our past studies , we here test how changes in masting patterns are modulated by individual tree size, which is a key life-history trait related to plant growth, survival, and reproduction (Bennett et al., 2015;Qiu et al., 2021;Stovall et al., 2019). ...
... Seed supply can be optimized by stand structure management, but such management is unlikely to fully compensate for the fecundity decline we observed. Depending on the climate change effects on later life stages, local extinctions may follow (Hylander & Ehrlén, 2013;Talluto et al., 2017;Tilman et al., 1994). These results set priorities for further research and management. ...
... Seed supply can be optimized by stand structure management, but such management is unlikely to fully compensate for the fecundity decline we observed. Depending on the climate change effects on later life stages, local extinctions may follow (Hylander & Ehrlén, 2013;Talluto et al., 2017;Tilman et al., 1994). These results set priorities for further research and management. ...
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