Links between plant species’ spatial and temporal responses to a warming climate

To generate realistic projections of species’ responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species’s climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.     

Range and niche shifts in response to past climate change in the desert horned lizard Phrynosoma platyrhinos

During climate change, species are often assumed to shift their geographic distributions (geographic ranges) in order to track environmental conditions – niches – to which they are adapted. Recent work, however, suggests that the niches do not always remain conserved during climate change but shift instead, allowing populations to persist in place or expand into new areas. We assessed the extent of range and niche shifts in response to the warming climate after the Last Glacial Maximum (LGM) in the desert horned lizard Phrynosoma platyrhinos, a species occupying the western deserts of North America. We used a phylogeographic approach with mitochondrial DNA sequences to approximate the species range during the LGM by identifying populations that exhibit a genetic signal of population stability versus those that exhibit a signal of a recent (likely post‐LGM) geographic expansion. We then compared the climatic niche that the species occupies today with the niche it occupied during the LGM using two models of simulated LGM climate. The genetic analyses indicated that P. platyrhinos persisted within the southern Mojave and Sonoran deserts throughout the latest glacial period and expanded from these deserts northwards, into the western and eastern Great Basin, after the LGM. The climatic niche comparisons revealed that P. platyrhinos expanded its climatic niche after the LGM towards novel, warmer and drier climates that allowed it to persist within the southern deserts. Simultaneously, the species shifted its climatic niche towards greater temperature and precipitation fluctuations after the LGM. We concluded that climatic changes at the end of the LGM promoted both range and niche shifts in this lizard. The mechanism that allowed the species to shift its niche remains unknown, but phenotypic plasticity likely contributes to the species ability to adjust to climate change.     

Temperature dependent sex determination and climate change

One possible response of species to climate change is shifting their geographical range so as to track their climatic niche. Many concerns have been raised about the species ability to disperse effectively. I argue that species may have mechanisms, like temperature-dependent sex determination (TSD), that are responsive to climate change and may facilitate an appropriate shift in their geographical range. More specifically, I hypothesize that, under stable climatic conditions, populations of some TSD species at the edge of their range are regulated by reduced growth rate (due to skewed sex ratios or due to limited availability of suitable nesting sites). Under climate change, these populations face new climatic conditions that trigger fast population growth (e.g. by more balanced sex ratio, or greater availability of nesting sites). Increased population size may lead to increased dispersal, and thus efficient colonization of the newly created habitat patches. So, the species rapidly tracks the geographical position of its climatic niche. This conceptual model is speculative but it leads to specific hypotheses, and opens up new research questions about the existence of prior adaptations that will enable the appropriate response to climate change.     

Intra‐specific variability and plasticity influence potential tree species distributions under climate change

Aim To assess the effect of local adaptation and phenotypic plasticity on the potential distribution of species under future climate changes. Trees may be adapted to specific climatic conditions; however, species range predictions have classically been assessed by species distribution models (SDMs) that do not account for intra‐specific genetic variability and phenotypic plasticity, because SDMs rely on the assumption that species respond homogeneously to climate change across their range, i.e. a species is equally adapted throughout its range, and all species are equally plastic. These assumptions could cause SDMs to exaggerate or underestimate species at risk under future climate change. Location The Iberian Peninsula. Methods Species distributions are predicted by integrating experimental data and modelling techniques. We incorporate plasticity and local adaptation into a SDM by calibrating models of tree survivorship with adaptive traits in provenance trials. Phenotypic plasticity was incorporated by calibrating our model with a climatic index that provides a measure of the differences between sites and provenances. Results We present a new modelling approach that is easy to implement and makes use of existing tree provenance trials to predict species distribution models under global warming. Our results indicate that the incorporation of intra‐population genetic diversity and phenotypic plasticity in SDMs significantly altered their outcome. In comparing species range predictions, the decrease in area occupancy under global warming conditions is smaller when considering our survival–adaptation model than that predicted by a ‘classical SDM’ calibrated with presence–absence data. These differences in survivorship are due to both local adaptation and plasticity. Differences due to the use of experimental data in the model calibration are also expressed in our results: we incorporate a null model that uses survival data from all provenances together. This model always predicts less reduction in area occupancy for both species than the SDM calibrated with presence–absence. Main conclusions We reaffirm the importance of considering adaptive traits when predicting species distributions and avoiding the use of occurrence data as a predictive variable. In light of these recommendations, we advise that existing predictions of future species distributions and their component populations must be reconsidered.     

The effects of phenotypic plasticity and local adaptation on forecasts of species range shifts under climate change

Species are the unit of analysis in many global change and conservation biology studies; however, species are not uniform entities but are composed of different, sometimes locally adapted, populations differing in plasticity. We examined how intraspecific variation in thermal niches and phenotypic plasticity will affect species distributions in a warming climate. We first developed a conceptual model linking plasticity and niche breadth, providing five alternative intraspecific scenarios that are consistent with existing literature. Secondly, we used ecological niche‐modeling techniques to quantify the impact of each intraspecific scenario on the distribution of a virtual species across a geographically realistic setting. Finally, we performed an analogous modeling exercise using real data on the climatic niches of different tree provenances. We show that when population differentiation is accounted for and dispersal is restricted, forecasts of species range shifts under climate change are even more pessimistic than those using the conventional assumption of homogeneously high plasticity across a species' range. Suitable population‐level data are not available for most species so identifying general patterns of population differentiation could fill this gap. However, the literature review revealed contrasting patterns among species, urging greater levels of integration among empirical, modeling and theoretical research on intraspecific phenotypic variation.     

Plasticity in Dendroclimatic Response across the Distribution Range of Aleppo Pine (Pinus halepensis)

We investigated the variability of the climate-growth relationship of Aleppo pine across its distribution range in the Mediterranean Basin. We constructed a network of tree-ring index chronologies from 63 sites across the region. Correlation function analysis identified the relationships of tree-ring index to climate factors for each site. We also estimated the dominant climatic gradients of the region using principal component analysis of monthly, seasonal, and annual mean temperature and total precipitation from 1,068 climatic gridpoints. Variation in ring width index was primarily related to precipitation and secondarily to temperature. However, we found that the dendroclimatic relationship depended on the position of the site along the climatic gradient. In the southern part of the distribution range, where temperature was generally higher and precipitation lower than the regional average, reduced growth was also associated with warm and dry conditions. In the northern part, where the average temperature was lower and the precipitation more abundant than the regional average, reduced growth was associated with cool conditions. Thus, our study highlights the substantial plasticity of Aleppo pine in response to different climatic conditions. These results do not resolve the source of response variability as being due to either genetic variation in provenance, to phenotypic plasticity, or a combination of factors. However, as current growth responses to inter-annual climate variability vary spatially across existing climate gradients, future climate-growth relationships will also likely be determined by differential adaptation and/or acclimation responses to spatial climatic variation. The contribution of local adaptation and/or phenotypic plasticity across populations to the persistence of species under global warming could be decisive for prediction of climate change impacts across populations. In this sense, a more complex forest dynamics modeling approach that includes the contribution of genetic variation and phenotypic plasticity can improve the reliability of the ecological inferences derived from the climate-growth relationships.     

Incorporating evolutionary adaptation in species distribution modelling reduces projected vulnerability to climate change

Based on the sensitivity of species to ongoing climate change, and numerous challenges they face tracking suitable conditions, there is growing interest in species' capacity to adapt to climatic stress. Here, we develop and apply a new generic modelling approach (AdaptR) that incorporates adaptive capacity through physiological limits, phenotypic plasticity, evolutionary adaptation and dispersal into a species distribution modelling framework. Using AdaptR to predict change in the distribution of 17 species of Australian fruit flies (Drosophilidae), we show that accounting for adaptive capacity reduces projected range losses by up to 33% by 2105. We identify where local adaptation is likely to occur and apply sensitivity analyses to identify the critical factors of interest when parameters are uncertain. Our study suggests some species could be less vulnerable than previously thought, and indicates that spatiotemporal adaptive models could help improve management interventions that support increased species' resilience to climate change.     

Climatic and historical factors controlling horizontal and vertical distribution patterns of two sympatric beech species, Fagus crenata Blume and Fagus japonica Maxim., in eastern Japan

The Japanese Archipelago is unique when viewed in terms of beech flora, since two native species, Fagus crenata Blume and Fagus japonica Maxim., occur sympatrically there. In order to examine the most important environmental or historical factors restricting the geographical ranges of these beech species in eastern Japan (34–44°N and 137–143°E), horizontal and vertical distributions were comparatively examined in detail. The study used two kinds of data sets: (1) DS1, constructed by assembling data from literature, herbarium specimens, etc., and (2) DS2, based on a mesh vegetation database. The upper range limit was expected to be in equilibrium with the current climatic conditions for both species (correlated with temperature factors for F. crenata and with snow depth for F. japonica). The lower range limit was also expected to be in equilibrium with the present climatic condition for both species, but the importance of competition with evergreen trees was also suggested. Dispersal limitation and a topo-geological barrier (for F. japonica) were expected to strongly restrict the northern range limit. Under contrasting climatic conditions in winter (between the Pacific and the Sea of Japan sides in eastern Japan) the geographic ranges of the two beech species are differentiated because of a difference in tolerance to heavy snowfall. A range shift model that assumes a migration lag along a horizontal direction because of dispersal limitation can explain the observed distribution patterns of these beech species in relation to climatic change in the Quaternary.     

Intraspecific trait variation across elevation predicts a widespread tree species' climate niche and range limits

Global change is widely altering environmental conditions which makes accurately predicting species range limits across natural landscapes critical for conservation and management decisions. If climate pressures along elevation gradients influence the distribution of phenotypic and genetic variation of plant functional traits, then such trait variation may be informative of the selective mechanisms and adaptations that help define climatic niche limits. Using extensive field surveys along 16 elevation transects and a large common garden experiment, we tested whether functional trait variation could predict the climatic niche of a widespread tree species (Populus angustifolia) with a double quantile regression approach. We show that intraspecific variation in plant size, growth, and leaf morphology corresponds with the species' total climate range and certain climatic limits related to temperature and moisture extremes. Moreover, we find evidence of genetic clines and phenotypic plasticity at environmental boundaries, which we use to create geographic predictions of trait variation and maximum values due to climatic constraints across the western US. Overall, our findings show the utility of double quantile regressions for connecting species distributions and climate gradients through trait‐based mechanisms. We highlight how new approaches like ours that incorporate genetic variation in functional traits and their response to climate gradients will lead to a better understanding of plant distributions as well as identifying populations anticipated to be maladapted to future environments.     

Non-equilibrium succession dynamics indicate continued northern migration of lodgepole pine

Because species affect ecosystem functioning, understanding migration processes is a key component of predicting future ecosystem responses to climate change. This study provides evidence of range expansion under current climatic conditions of an indigenous species with strong ecosystem effects. Surveys of stands along the northern distribution limit of lodgepole pine (Pinus contorta var. latifolia) in central Yukon Territory, Canada showed consistent increases in pine dominance following fire. These patterns differed strongly from those observed at sites where pine has been present for several thousand years. Differences in species thinning rates are unlikely to account for the observed increases in pine dominance. Rates of pine regeneration at its range limits were equivalent to those of spruce, indicating a capacity for rapid local population expansion. The study also found no evidence of strong climatic limitation of pine population growth at the northern distribution limit. We interpret these data as evidence of current pine expansion at its range limits and conclude that the northern distribution of lodgepole pine is not in equilibrium with current climate. This study has implications for our ability to predict vegetation response to climate change when populations may lag in their response to climate.     

Higher susceptibility of beech to drought in comparison to oak

The expected increase in drought severity and frequency as a result of anthropogenic climate change leads to concerns about the ability of native tree species to cope with these changes. To determine the susceptibility of Fagus sylvatica (European beech) and Quercus robur (pedunculate oak) – the two dominant deciduous tree species in Central Europe – to drought, we quantified the climate sensitivity and drought-response of radial growth for both species using an array of dendroecological methods. Tree-ring data were collected from a site east of Coburg, Bavaria which had shown pronounced stress-symptoms (early leaf coloration) during the record drought of 2018. Climate-growth relationships were used to establish the sensitivity of radial growth to multiple climatic variables. The impact of specific drought events on tree growth was quantified using tolerance indices. In addition, we employed a Principal Component Gradient Analysis (PCGA) and remote sensing data (MODIS Normalized Difference Vegetation Index (NDVI)) to delineate the species specific drought responses. Using these methods we were able to show a clear difference in drought susceptibility between beech and oak. Beech displayed a higher sensitivity to temperature and the standardized precipitation evapotranspiration index (SPEI) and showed lower resistance and resilience to drought events than oak. In particular, beech was unable to fully recover from the 2003 drought, after which it expressed a stark growth decline, i.e. drought legacies, which was not observed for oak. The PCGA revealed a clear differentiation in the grouping of drought responses between beech and oak, supporting the findings of the climate-growth analysis and the tolerance indices. Correlations of NDVI and ring-width indices (RWI) indicated that under normal climatic conditions NDVI variability is linked to the start of the growing season. This is in contrast to drought years, such as 2003, where summer NDVI mirrored the drought response of beech and oak. These results reveal beech to have both a higher sensitivity to summer temperature and SPEI and a higher susceptibility to drought events. Although, in the past high plasticity and adaptability to drought have been attributed to both beech and oak, our study assigns beech a higher risk than oak to suffer from anticipated increases in drought frequency and intensity as a consequence of climate change.     

Challenges for growth of beech and co-occurring conifers in a changing climate context

Given recent climatic trends, it is crucial to understand the plasticity and adaptiveness of forest trees in order to evaluate their current and future responses to changing climatic conditions. We investigated inter- and intra-annual xylem growth and its relation to weather factors in European beech (Fagus sylvatica) and a co-occurring conifer, either Norway spruce (Picea abies) or Scots pine (Pinus sylvestris), at five sites with different elevations and climatic conditions, in Spain, Slovenia and the Czech Republic. The selected sites were located in central and marginal parts across the distribution range of the investigated species. The results showed that climate-growth relationships vary between different species at the same site and within the same species at different sites, indicating diverse survival strategies among the observed species but also high plastic capacity within the same species. Moreover, xylogenesis analysis revealed the capacity of trees to adapt the beginning and cessation of wood formation depending on conditions. In case of beech, shortening growth duration in dryer and warmer environments and the opposite in the case of conifers, with which the growing period was extended in such conditions. Consequently, the plasticity capacity of beech was limited due to short growth duration, while conifers (especially pine) proved to be able to compensate climatic constrictions by growing over a longer period and becoming more adapted to survival in drought-prone environments.     

Climate-driven range dynamics of the freshwater limpet, Ancylus fluviatilis (Pulmonata, Basommatophora)

Aim Our aim was to understand the processes that have shaped the present‐day distribution of the freshwater limpet Ancylus fluviatilis sensu stricto in order to predict the consequences of global climate change for the geographical range of this species. Location North‐western Europe. Methods We sampled populations of A. fluviatilis sensu stricto over the entire range of the species (north‐western Europe) and sequenced 16S ribosomal RNA (16S) and cytochrome oxidase subunit I (COI) mitochondrial fragments to perform phylogenetic and phylogeographical analyses. Climatic niche modelling allowed us to infer the climatic preferences of the species. A principal components analysis identified the most important climatic factors explaining the actual range of A. fluviatilis. We also identified which climatic factor was the most limiting at range margins, and predicted the species’ geographical range under a climate change scenario [Community Climate Model 3 (CCM3)]. Results By means of the phylogeographical analysis, we infer that A. fluviatilis sensu stricto occupied northern refuges during the Last Glacial Maximum. We show that the climatic preferences of Baltic populations are significantly different from those of Central European populations. The projection of the occupied area under the CCM3 climate model predicts a moderate poleward shift of the northern range limits, but a dramatic loss of areas currently occupied, for instance in northern Germany and in southern Great Britain. Main conclusions The post‐glacial range dynamics of A. fluviatilis are not governed by niche conservatism. Therefore, we must be cautious about bioclimatic model predictions: the expected impact of climate change could be tempered by the adaptive potential this species has already shown in its evolutionary history. Thus, modelling approaches should rather be seen as conservative forecasts of altered species ranges as long as the adaptive potential of the organisms in question cannot be predicted.     

Climatic niche breadth determines the response of bumblebees (<Emphasis Type="Italic">Bombus</Emphasis> spp.) to climate warming in mountain areas of the Northern Iberian Peninsula

Studies examining species range shifts in the face of climate change have consistently found that response patterns are complex and varied, suggesting that ecological traits might be affecting species response. However, knowledge of how the traits of a species determine its response to climate change is still poorly understood. Here we investigate the role of species-specific climate niche breadth in forecasting bumblebee (Bombus spp.) responses to regional climate warming in the Cantabrian Range (north-western Iberian Peninsula). Climate niche breadth was defined using known data for occurrences of specific species at their continental (i.e., European) scale of distribution. For each bumblebee species, climate niche breadth was found to be related to (1) the elevational range shifts of species between their historical (1988–1989) and recent (2007–2009) distribution and (2) the variation in the climatic conditions of the localities they inhabited (i.e., the local climate space) between both study periods. Our results show a strong relationship between climate niche breadth, particularly thermal niche breadth, and the response of bumblebee species to climate warming, but only when this response was determined as variations in local climate space. The main conclusions of our work are thus twofold. First, variations in the climatic conditions underlying range shifts are useful in making accurate assessments of the impact of climate change on species distributions. Second, climate niche breadth is a particularly informative ecological trait for forecasting variations in species responses to climate change.     

Increased Phenotypic Plasticity to Climate May Have Boosted the Invasion Success of Polyploid Centaurea stoebe

Phenotypic plasticity may allow organisms to cope with altered environmental conditions as e.g. after the introduction into a new range. In particular polyploid organisms, containing more than two sets of chromosomes, may show high levels of plasticity, which could in turn increase their environmental tolerance and invasiveness. Here, we studied the role of phenotypic plasticity in the invasion of Centaurea stoebe (Asteraceae), which in the native range in Europe occurs as diploids and tetraploids, whereas in the introduced range in North America so far only tetraploids have been found. In a common garden experiment at two sites in the native range, we grew half-sibs of the three geo-cytotypes (native European diploids, European tetraploids and invasive North American tetraploids) from a representative sample of 27 populations. We measured the level and the adaptive significance of phenotypic plasticity in eco-physiological and life-history traits in response to the contrasting climatic conditions at the two study sites as well as three different soil conditions in pots, simulating the most crucial abiotic differences between the native and introduced range. European tetraploids showed increased levels of phenotypic plasticity as compared to diploids in response to the different climatic conditions in traits associated with rapid growth and fast phenological development. Moreover, we found evidence for adaptive plasticity in these traits, which suggests that increased plasticity may have contributed to the invasion success of tetraploid C. stoebe by providing an advantage under the novel climatic conditions. However, in invasive tetraploids phenotypic plasticity was similar to that of native tetraploids, indicating no evolution of increased plasticity during invasions. Our findings provide the first empirical support for increased phenotypic plasticity associated with polyploids, which may contribute to their success as invasive species in novel environments.     

Climatic limits for the present distribution of beech (Fagus L.) species in the world

Aim Beech (Fagus L., Fagaceae) species are representative trees of temperate deciduous broadleaf forests in the Northern Hemisphere. We focus on the distributional limits of beech species, in particular on identifying climatic factors associated with their present range limits. Location Beech species occur in East Asia, Europe and West Asia, and North America. We collated information on both the southern and northern range limits and the lower and upper elevational limits for beech species in each region. Methods In total, 292 lower/southern limit and 310 upper/northern limit sites with available climatic data for all 11 extant beech species were collected by reviewing the literature, and 13 climatic variables were estimated for each site from climate normals at nearby stations. We used principal components analysis (PCA) to detect climatic variables most strongly associated with the distribution of beech species and to compare the climatic spaces for the different beech species. Results Statistics for thermal and moisture climatic conditions at the lower/southern and upper/northern limits of all world beech species are presented. The first two PCA components accounted for 70% and 68% of the overall variance in lower/southern and upper/northern range limits, respectively. The first PCA axis represented a thermal gradient, and the second a moisture gradient associated with the world‐wide distribution pattern of beech species. Among thermal variables, growing season warmth was most important for beech distribution, but winter low temperature (coldness and mean temperature for the coldest month) and climatic continentality were also coupled with beech occurrence. The moisture gradient, indicated by precipitation and moisture indices, showed regional differences. American beech had the widest thermal range, Japanese beeches the most narrow; European beeches occurred in the driest climate, Japanese beeches the most humid. Climatic spaces for Chinese beech species were between those of American and European species. Main conclusions The distributional limits of beech species were primarily associated with thermal factors, but moisture regime also played a role. There were some regional differences in the climatic correlates of distribution. The growing season temperature regime was most important in explaining distribution of Chinese beeches, whilst their northward distribution was mainly limited by shortage of precipitation. In Japan, distribution limits of beech species were correlated with summer temperature, but the local dominance of beech was likely to be dependent on snowfall and winter low temperature. High summer temperature was probably a limiting factor for southward extension of American beech, while growing season warmth seemed critical for its northward distribution. Although the present distribution of beech species corresponded well to the contemporary climate in most areas, climatic factors could not account for some distributions, e. g., that of F. mexicana compared to its close relative F. grandifolia. It is likely that historical factors play a secondary role in determining the present distribution of beech species. The lack of F. grandifolia on the island of Newfoundland, Canada, may be due to inadequate growing season warmth. Similarly, the northerly distribution of beech in Britain has not reached its potential limit, perhaps due to insufficient time since deglaciation to expand its range.     

Back from a predicted climatic extinction of an island endemic: a future for the Corsican Nuthatch

The Corsican Nuthatch (Sitta whiteheadi) is red-listed as vulnerable to extinction by the IUCN because of its endemism, reduced population size, and recent decline. A further cause is the fragmentation and loss of its spatially-restricted favourite habitat, the Corsican pine (Pinus nigra laricio) forest. In this study, we aimed at estimating the potential impact of climate change on the distribution of the Corsican Nuthatch using species distribution models. Because this species has a strong trophic association with the Corsican and Maritime pines (P. nigra laricio and P. pinaster), we first modelled the current and future potential distribution of both pine species in order to use them as habitat variables when modelling the nuthatch distribution. However, the Corsican pine has suffered large distribution losses in the past centuries due to the development of anthropogenic activities, and is now restricted to mountainous woodland. As a consequence, its realized niche is likely significantly smaller than its fundamental niche, so that a projection of the current distribution under future climatic conditions would produce misleading results. To obtain a predicted pine distribution at closest to the geographic projection of the fundamental niche, we used available information on the current pine distribution associated to information on the persistence of isolated natural pine coppices. While common thresholds (maximizing the sum of sensitivity and specificity) predicted a potential large loss of the Corsican Nuthatch distribution by 2100, the use of more appropriate thresholds aiming at getting closer to the fundamental distribution of the Corsican pine predicted that 98% of the current presence points should remain potentially suitable for the nuthatch and its range could be 10% larger in the future. The habitat of the endemic Corsican Nuthatch is therefore more likely threatened by an increasing frequency and intensity of wildfires or anthropogenic activities than by climate change.     

Forecasting Distributional Responses of Limber Pine to Climate Change at Management-Relevant Scales in Rocky Mountain National Park

Resource managers at parks and other protected areas are increasingly expected to factor climate change explicitly into their decision making frameworks. However, most protected areas are small relative to the geographic ranges of species being managed, so forecasts need to consider local adaptation and community dynamics that are correlated with climate and affect distributions inside protected area boundaries. Additionally, niche theory suggests that species'' physiological capacities to respond to climate change may be underestimated when forecasts fail to consider the full breadth of climates occupied by the species rangewide. Here, using correlative species distribution models that contrast estimates of climatic sensitivity inferred from the two spatial extents, we quantify the response of limber pine (Pinus flexilis) to climate change in Rocky Mountain National Park (Colorado, USA). Models are trained locally within the park where limber pine is the community dominant tree species, a distinct structural-compositional vegetation class of interest to managers, and also rangewide, as suggested by niche theory. Model forecasts through 2100 under two representative concentration pathways (RCP 4.5 and 8.5 W/m²) show that the distribution of limber pine in the park is expected to move upslope in elevation, but changes in total and core patch area remain highly uncertain. Most of this uncertainty is biological, as magnitudes of projected change are considerably more variable between the two spatial extents used in model training than they are between RCPs, and novel future climates only affect local model predictions associated with RCP 8.5 after 2091. Combined, these results illustrate the importance of accounting for unknowns in species'' climatic sensitivities when forecasting distributional scenarios that are used to inform management decisions. We discuss how our results for limber pine may be interpreted in the context of climate change vulnerability and used to help guide adaptive management.     

Patterns of Cross-Continental Variation in Tree Seed Mass in the Canadian Boreal Forest

Seed mass is an adaptive trait affecting species distribution, population dynamics and community structure. In widely distributed species, variation in seed mass may reflect both genetic adaptation to local environments and adaptive phenotypic plasticity. Acknowledging the difficulty in separating these two aspects, we examined the causal relationships determining seed mass variation to better understand adaptability and/or plasticity of selected tree species to spatial/climatic variation. A total of 504, 481 and 454 seed collections of black spruce (Picea mariana (Mill.) B.S.P.), white spruce (Picea glauca (Moench) Voss) and jack pine (Pinus banksiana Lamb) across the Canadian Boreal Forest, respectively, were selected. Correlation analyses were used to determine how seed mass vary with latitude, longitude, and altitude. Structural Equation Modeling was used to examine how geographic and climatic variables influence seed mass. Climatic factors explained a large portion of the variation in seed mass (34, 14 and 29%, for black spruce, white spruce and jack pine, respectively), indicating species-specific adaptation to long term climate conditions. Higher annual mean temperature and winter precipitation caused greater seed mass in black spruce, but annual precipitation was the controlling factor for white spruce. The combination of factors such as growing season temperature and evapotranspiration, temperature seasonality and annual precipitation together determined seed mass of jack pine. Overall, sites with higher winter temperatures were correlated with larger seeds. Thus, long-term climatic conditions, at least in part, determined spatial variation in seed mass. Black spruce and Jack pine, species with relatively more specific habitat requirements and less plasticity, had more variation in seed mass explained by climate than did the more plastic species white spruce. As traits such as seed mass are related to seedling growth and survival, they potentially influence forest species composition in a changing climate and should be included in future modeling of vegetation shifts.     

Contrasting growth forecasts across the geographical range of Scots pine due to altitudinal and latitudinal differences in climatic sensitivity

Ongoing changes in global climate are altering ecological conditions for many species. The consequences of such changes are typically most evident at the edge of a species’ geographical distribution, where differences in growth or population dynamics may result in range expansions or contractions. Understanding population responses to different climatic drivers along wide latitudinal and altitudinal gradients is necessary in order to gain a better understanding of plant responses to ongoing increases in global temperature and drought severity. We selected Scots pine (Pinus sylvestris L.) as a model species to explore growth responses to climatic variability (seasonal temperature and precipitation) over the last century through dendrochronological methods. We developed linear models based on age, climate and previous growth to forecast growth trends up to year 2100 using climatic predictions. Populations were located at the treeline across a latitudinal gradient covering the northern, central and southernmost populations and across an altitudinal gradient at the southern edge of the distribution (treeline, medium and lower elevations). Radial growth was maximal at medium altitude and treeline of the southernmost populations. Temperature was the main factor controlling growth variability along the gradients, although the timing and strength of climatic variables affecting growth shifted with latitude and altitude. Predictive models forecast a general increase in Scots pine growth at treeline across the latitudinal distribution, with southern populations increasing growth up to year 2050, when it stabilizes. The highest responsiveness appeared at central latitude, and moderate growth increase is projected at the northern limit. Contrastingly, the model forecasted growth declines at lowland‐southern populations, suggesting an upslope range displacement over the coming decades. Our results give insight into the geographical responses of tree species to climate change and demonstrate the importance of incorporating biogeographical variability into predictive models for an accurate prediction of species dynamics as climate changes.