Moderate disturbances accelerate forest transition dynamics under climate change in the temperate–boreal ecotone of eastern North America

Several temperate tree species are expected to migrate northward and colonize boreal forests in response to climate change. Tree migrations could lead to transitions in forest types, but these could be influenced by several non‐climatic factors, such as disturbances and soil conditions. We analysed over 10,000 forest inventory plots, sampled from 1970 to 2018 in meridional Québec, Canada, to identify what environmental conditions promote or prevent regional‐scale forest transitions. We used a continuous‐time multi‐state Markov model to quantify the probabilities of transitions between forest states (temperate, boreal, mixed, pioneer) as a function of climate (mean temperature and climate moisture index during the growing season), soil conditions (pH and drainage) and disturbances (severity levels of natural disturbances and logging). We further investigate how different disturbance types and severities impact forests' short‐term transient dynamics and long‐term equilibrium using properties of Markov transition matrices. The most common transitions observed during the study period were from mixed to temperate states, as well as from pioneer to boreal forests. In our study, transitions were mainly driven by natural and anthropogenic disturbances and secondarily by climate, whereas soil characteristics exerted relatively minor constraints. While major disturbances only promoted transitions to the pioneer state, moderate disturbances increased the probability of transition from mixed to temperate states. Long‐term projections of our model under the current environmental conditions indicate that moderate disturbances would promote a northward shift of the temperate forest. Moreover, disturbances reduced turnover and convergence time for all transitions, thereby accelerating forest dynamics. Contrary to our expectation, mixed to temperate transitions were not driven by temperate tree recruitment but by mortality and growth. Overall, our results suggest that moderate disturbances could catalyse rapid forest transitions and accelerate broad‐scale biome shifts.     

Disturbance maintains alternative biome states

Understanding the mechanisms controlling the distribution of biomes remains a challenge. Although tropical biome distribution has traditionally been explained by climate and soil, contrasting vegetation types often occur as mosaics with sharp boundaries under very similar environmental conditions. While evidence suggests that these biomes are alternative states, empirical broad‐scale support to this hypothesis is still lacking. Using community‐level field data and a novel resource‐niche overlap approach, we show that, for a wide range of environmental conditions, fire feedbacks maintain savannas and forests as alternative biome states in both the Neotropics and the Afrotropics. In addition, wooded grasslands and savannas occurred as alternative grassy states in the Afrotropics, depending on the relative importance of fire and herbivory feedbacks. These results are consistent with landscape scale evidence and suggest that disturbance is a general factor driving and maintaining alternative biome states and vegetation mosaics in the tropics.     

Sixteen hundred years of increasing tree cover prior to modern deforestation in Southern Amazon and Central Brazilian savannas

Tropical ecosystems are under increasing pressure from land‐use change and deforestation. Changes in tropical forest cover are expected to affect carbon and water cycling with important implications for climatic stability at global scales. A major roadblock for predicting how tropical deforestation affects climate is the lack of baseline conditions (i.e., prior to human disturbance) of forest–savanna dynamics. To address this limitation, we developed a long‐term analysis of forest and savanna distribution across the Amazon–Cerrado transition of central Brazil. We used soil organic carbon isotope ratios as a proxy for changes in woody vegetation cover over time in response to fluctuations in precipitation inferred from speleothem oxygen and strontium stable isotope records. Based on stable isotope signatures and radiocarbon activity of organic matter in soil profiles, we quantified the magnitude and direction of changes in forest and savanna ecosystem cover. Using changes in tree cover measured in 83 different locations for forests and savannas, we developed interpolation maps to assess the coherence of regional changes in vegetation. Our analysis reveals a broad pattern of woody vegetation expansion into savannas and densification within forests and savannas for at least the past ~1,600 years. The rates of vegetation change varied significantly among sampling locations possibly due to variation in local environmental factors that constrain primary productivity. The few instances in which tree cover declined (7.7% of all sampled profiles) were associated with savannas under dry conditions. Our results suggest a regional increase in moisture and expansion of woody vegetation prior to modern deforestation, which could help inform conservation and management efforts for climate change mitigation. We discuss the possible mechanisms driving forest expansion and densification of savannas directly (i.e., increasing precipitation) and indirectly (e.g., decreasing disturbance) and suggest future research directions that have the potential to improve climate and ecosystem models.     

Synergistic effects of drought and deforestation on the resilience of the south-eastern Amazon rainforest

The south-eastern Amazon rainforest is subject to ongoing deforestation and is expected to become drier due to climate change. Recent analyses of the distribution of tree cover in the tropics show three modes that have been interpreted as representing alternative stable states: forest, savanna and treeless states. This situation implies that a change in environmental conditions, such as in the climate, could cause critical transitions from a forest towards a savanna ecosystem. Shifts to savanna might also occur if perturbations such as deforestation exceed a critical threshold. Recovering the forest would be difficult as the savanna will be stabilized by a feedback between tree cover and fire. Here we explore how environmental changes and perturbations affect the forest by using a simple model with alternative tree-cover states. We focus on the synergistic effects of precipitation reduction and deforestation on the probability of regime shifts in the south-eastern Amazon rainforest. The analysis indicated that in a large part of the south-eastern Amazon basin rainforest and savanna could be two alternative states, although massive forest dieback caused by mean-precipitation reduction alone is unlikely. However, combinations of deforestation and climate change triggered up to 6.6 times as many local regime shifts than the two did separately, causing large permanent forest losses in the studied region. The results emphasize the importance of reducing deforestation rates in order to prevent a climate-induced dieback of the south-eastern Amazon rainforest.     

Land‐use change outweighs projected effects of changing rainfall on tree cover in sub‐Saharan Africa

Global change will likely affect savanna and forest structure and distributions, with implications for diversity within both biomes. Few studies have examined the impacts of both expected precipitation and land use changes on vegetation structure in the future, despite their likely severity. Here, we modeled tree cover in sub‐Saharan Africa, as a proxy for vegetation structure and land cover change, using climatic, edaphic, and anthropic data (R² = 0.97). Projected tree cover for the year 2070, simulated using scenarios that include climate and land use projections, generally decreased, both in forest and savanna, although the directionality of changes varied locally. The main driver of tree cover changes was land use change; the effects of precipitation change were minor by comparison. Interestingly, carbon emissions mitigation via increasing biofuels production resulted in decreases in tree cover, more severe than scenarios with more intense precipitation change, especially within savannas. Evaluation of tree cover change against protected area extent at the WWF Ecoregion scale suggested areas of high biodiversity and ecosystem services concern. Those forests most vulnerable to large decreases in tree cover were also highly protected, potentially buffering the effects of global change. Meanwhile, savannas, especially where they immediately bordered forests (e.g. West and Central Africa), were characterized by a dearth of protected areas, making them highly vulnerable. Savanna must become an explicit policy priority in the face of climate and land use change if conservation and livelihoods are to remain viable into the next century.     

Tipping points in tropical tree cover: linking theory to data

It has recently been found that the frequency distribution of remotely sensed tree cover in the tropics has three distinct modes, which seem to correspond to forest, savanna, and treeless states. This pattern has been suggested to imply that these states represent alternative attractors, and that the response of these systems to climate change would be characterized by critical transitions and hysteresis. Here, we show how this inference is contingent upon mechanisms at play. We present a simple dynamical model that can generate three alternative tree cover states (forest, savanna, and a treeless state), based on known mechanisms, and use this model to simulate patterns of tree cover under different scenarios. We use these synthetic data to show that the hysteresis inferred from remotely sensed tree cover patterns will be inflated by spatial heterogeneity of environmental conditions. On the other hand, we show that the hysteresis inferred from satellite data may actually underestimate real hysteresis in response to climate change if there exists a positive feedback between regional tree cover and precipitation. Our results also indicate that such positive feedback between vegetation and climate should cause direct shifts between forest and a treeless state (rather than through an intermediate savanna state) to become more likely. Finally, we show how directionality of historical change in conditions may bias the observed relationship between tree cover and environmental conditions.     

The combined role of dispersal and niche evolution in the diversification of Neotropical lizards

Ecological requirements and environmental conditions can influence diversification across temporal and spatial scales. Understanding the role of ecological niche evolution under phylogenetic contexts provides insights on speciation mechanisms and possible responses to future climatic change. Large‐scale phyloclimatic studies on the megadiverse Neotropics, where biomes with contrasting vegetation types occur in narrow contact, are rare. We integrate ecological and biogeographic data with phylogenetic comparative methods, to investigate the relative roles of biogeographic events and niche divergence and conservatism on the diversification of the lizard genus Kentropyx Spix, 1825 (Squamata: Teiidae), distributed in South American rainforests and savannas. Using five molecular markers, we estimated a dated species tree, which recovered three clades coincident with previously proposed species groups diverging during the mid‐Miocene. Biogeography reconstruction indicates a role of successive dispersal events from an ancestral range in the Brazilian Shield and western Amazonia. Ancestral reconstruction of climatic tolerances and niche overlap metrics indicates a trend of conservatism during the diversification of groups from the Amazon Basin and Guiana Shield, and a strong signal of niche divergence in the Brazilian Shield savannas. Our results suggest that climatic‐driven divergence at dynamic forest‐savanna borders might have resulted in adaptation to new environmental niches, promoting habitat shifts and shaping speciation patterns of Neotropical lizards. Dispersal and ecological divergence could have a more important role in Neotropical diversification than previously thought.     

Decadal dynamics of tree cover in an Australian tropical savanna

Spatio‐temporal variation in tropical savanna tree cover remains poorly understood. We aimed to quantify the drivers of tree cover in tropical mesic savannas in Kakadu National Park by relating changes in tree cover over 40 years to: mean annual rainfall, fire activity, initial tree cover and prior changes in tree cover. Aerial photography, acquired in 1964, 1984 and 2004, was obtained for fifty sites in Kakadu that spanned a rainfall gradient from approximately 1200 to 1600 mm. The remotely sensed estimates of tree cover were validated via field survey. Linear mixed effects modelling and multi‐model inference were used to assess the strength and form of the relationships between tree cover and predictor variables. Over the 40 years, tree cover across these savannas increased on average by 4.94 ± 0.88%, but was spatio‐temporally variable. Tree cover showed a positive albeit weak trend across the rainfall gradient. The strength of this positive relationship varied over the three measurement times, and this suggests that other factors are important in controlling tree cover. Tree cover was positively related to prior tree cover, and negatively correlated with fire activity. Over 20 years tree cover was more likely to increase if (i) tree cover was initially low or (ii) had decreased in the previous 20‐year interval or (iii) there had been fewer fires. Across the examined rainfall gradient, the greater variability in fire activity and inherently higher average tree cover at the wetter latitudes resulted in greater dynamism of tree cover compared with the drier latitudes. This is consistent with savanna tree cover being determined by interactions between mean annual rainfall, tree competition and frequent fire in these tropical mesic savannas.     

Drought-induced tree death in savanna

Increasing densities of woody plants in savannas has been attributed to both elevated atmospheric CO₂ and reduced burning with grazing management, such that the biome could represent a substantial carbon sink. However, we show that extreme droughts (less than two-thirds expected rainfall over 3 years) occur in the drier half of the savanna biome and can cause substantial tree death. An Australian case study reveals that a net increase in tree cover over five decades of above-average rainfall was offset by sudden tree death during drought. The relationship between woody cover change and rainfall is moderated by competition with growth being facilitated by low woody cover and drought-induced death more likely as the woody component of savanna increases. The results are not supportive of a sustained increase in the woody component of xeric savannas resulting from CO₂ fertilization or land management. Extensive tree death in savanna regions will become a stark consequence of climate change if predictions of increasing severity and frequency of drought are realized.     

Analysis of stable states in global savannas: is the CART pulling the horse?

Multiple stable states, bifurcations and thresholds are fashionable concepts in the ecological literature, a recognition that complex ecosystems may at times exhibit the interesting dynamic behaviours predicted by relatively simple biomathematical models. Recently, several papers in Global Ecology and Biogeography, Proceedings of the National Academy of Sciences USA, Science and elsewhere have attempted to quantify the prevalence of alternate stable states in the savannas of Africa, Australia and South America, and the tundra–taiga–grassland transitions of the circum‐boreal region using satellite‐derived woody canopy cover. While we agree with the logic that basins of attraction can be inferred from the relative frequencies of ecosystem states observed in space and time, we caution that the statistical methodologies underlying the satellite product used in these studies may confound our ability to infer the presence of multiple stable states. We demonstrate this point using a uniformly distributed ‘pseudo‐tree cover’ database for Africa that we use to retrace the steps involved in creation of the satellite tree‐cover product and subsequent analysis. We show how classification and regression tree (CART)‐based products may impose discontinuities in satellite tree‐cover estimates even when such discontinuities are not present in reality. As regional and global remote sensing and geospatial data become more easily accessible for ecological studies, we recommend careful consideration of how error distributions in remote sensing products may interact with the data needs and theoretical expectations of the ecological process under study.     

How do disturbances and environmental heterogeneity affect the pace of forest distribution shifts under climate change?

Although it is widely predicted that the geographic distributions of tree species and forest types will undergo substantial shifts in future, modelling approaches used to date are largely unable to project the pace at which forest distributions will respond to environmental change. The expansion and contraction of forest distributions act against considerable demographic inertia in the present composition and size‐structure of forest stands as climate‐induced changes in growth, mortality, and recruitment alter population dynamics through time. We aimed to better understand how shifts in forest distributions reflect long‐term changes in tree demographic rates and population dynamics, and how such shifts are influenced by 1) disturbance from forest harvesting and 2) local environmental heterogeneity. Using a simple, data‐constrained gap model, we simulated regional forest dynamics in the eastern United States over the next 500 yr. We then compared the geographic distributions of five different forest types through time under present and altered climatic conditions, in scenarios that variously included and excluded forest harvesting and environmental heterogeneity. Although we held climate fixed after 100 yr, it took another 160 yr after this for these forest types to collectively experience 90% of their eventual climate‐related distribution gains and losses. Competition strongly affected the nature of responses to climate change. Harvesting accelerated and amplified gains by an early‐successional forest type at the expense of a late‐successional one, but these gains did not occur faster than those for other forest types. Environmental heterogeneity had little effect on distribution gains or losses through time. These findings indicate that forest distributions should respond quite slowly to climate change, with the leading and trailing edges of different forest types shifting over a span of centuries. Disturbances can expedite some transitions, but are unlikely to lead to wholesale changes in forest types in the coming decades.     

Woody cover in African savannas: the role of resources, fire and herbivory

Aim To determine the functional relationships between, and the relative importance of, different driver variables (mean annual precipitation, soil properties, fire and herbivory) in regulating woody plant cover across broad environmental gradients in African savannas. Location Savanna grasslands of East, West and Southern Africa. Methods The dependence of woody cover on mean annual precipitation (MAP), soil properties (texture, nitrogen mineralization potential and total phosphorus), fire regimes, and herbivory (grazer, browser + mixed feeder, and elephant biomass) was determined for 161 savanna sites across Africa using stochastic gradient boosting, a refinement of the regression tree analysis technique. Results All variables were significant predictors of woody cover, collectively explaining 71% of the variance in our data set. However, their relative importance as regulators of woody cover varied. MAP was the most important predictor, followed by fire return periods, soil characteristics and herbivory regimes. Woody cover showed a strong positive dependence on MAP between 200 and 700 mm, but no dependence on MAP above this threshold when the effects of other predictors were accounted for. Fires served to reduce woody cover below rainfall‐determined levels. Woody cover showed a complex, non‐linear relationship with total soil phosphorus, and was negatively correlated with clay content. There was a strong negative dependence of woody cover on soil nitrogen (N) availability, suggesting that increased N‐deposition may cause shifts in savannas towards more grassy states. Elephants, mixed feeders and browsers had negative effects on woody cover. Grazers, on the other hand, depressed woody cover at low biomass, but favoured woody vegetation when their biomass exceeded a certain threshold. Main conclusions Our results indicate complex and contrasting relationships between woody cover, rainfall, soil properties and disturbance regimes in savannas, and suggest that future environmental changes such as altered precipitation regimes, N‐enrichment and elevated levels of CO₂ are likely to have opposing, and potentially interacting, influences on the tree–grass balance in savannas.     

A continental-scale analysis of tree cover in African savannas

Aim We present a continental‐scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. Location African savannas. Methods Tree cover is represented as a resource‐driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi‐empirical modelling and information theory to identify the best models. Results Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. Main conclusions The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These ‘perturbation’ effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi‐arid and mesic savannas (400–1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600–2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24–34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.     

Can savannas become forests? A coupled analysis of nutrient stocks and fire thresholds in central Brazil

Aims

The effects of fire ensure that large areas of the seasonal tropics are maintained as savannas. The advance of forests into these areas depends on shifts in species composition and the presence of sufficient nutrients. Predicting such transitions, however, is difficult due to a poor understanding of the nutrient stocks required for different combinations of species to resist and suppress fires.

Methods

We compare the amounts of nutrients required by congeneric savanna and forest trees to reach two thresholds of establishment and maintenance: that of fire resistance, after which individual trees are large enough to survive fires, and that of fire suppression, after which the collective tree canopy is dense enough to minimize understory growth, thereby arresting the spread of fire. We further calculate the arboreal and soil nutrient stocks of savannas, to determine if these are sufficient to support the expansion of forests following initial establishment.

Results

Forest species require a larger nutrient supply to resist fires than savanna species, which are better able to reach a fire-resistant size under nutrient limitation. However, forest species require a lower nutrient supply to attain closed canopies and suppress fires; therefore, the ingression of forest trees into savannas facilitates the transition to forest. Savannas have sufficient N, K, and Mg, but require additional P and Ca to build high-biomass forests and allow full forest expansion following establishment.

Conclusions

Tradeoffs between nutrient requirements and adaptations to fire reinforce savanna and forest as alternate stable states, explaining the long-term persistence of vegetation mosaics in the seasonal tropics. Low-fertility limits the advance of forests into savannas, but the ingression of forest species favors the formation of non-flammable states, increasing fertility and promoting forest expansion.     

Scale matters: fire–vegetation feedbacks are needed to explain tropical tree cover at the local scale

At a broad (regional to global) spatial scale, tropical vegetation is controlled by climate; at the local scale, it is believed to be determined by interactions between disturbance, vegetation and local conditions (soil and topography) through feedback processes. It has recently been suggested that strong fire–vegetation feedback processes may not be needed to explain tree‐cover patterns in tropical ecosystems and that climate–fire determinism is an alternative possibility. This conclusion was based on the fact that it is possible to reproduce observed patterns in tropical regions (e.g. a trimodal frequency distribution of tree cover) using a simple model that does not explicitly incorporate fire–vegetation feedback processes. We argue that these two mechanisms (feedbacks versus fire–climate control) operate at different spatial and temporal scales; it is not possible to evaluate the role of a process acting at fine scales (e.g. fire–vegetation feedbacks) using a model designed to reproduce regional‐scale pattern (scale mismatch). While the distributions of forest and savannas are partially determined by climate, many studies are providing evidence that the most parsimonious explanation for their environmental overlaps is the existence of feedback processes. Climate is unlikely to be an alternative to feedback processes; rather, climate and fire–vegetation feedbacks are complementary processes at different spatial and temporal scales.     

Phylogeography and paleodistribution models of a widespread birch (Betula platyphylla Suk.) across East Asia: Multiple refugia,multidirectional expansion,and heterogeneous genetic pattern

Widespread tree species cover large geographical areas and play important roles in various vegetation types. Understanding how these species responded to historical climatic changes is important for understanding community assembly mechanisms with evolutionary and conservation implications. However, the location of refugial areas and postglacial history of widespread trees in East Asia remain poorly known. We combined microsatellite data (63 populations, 1756 individuals) and ecological niche modeling to examine the range‐wide population diversity, genetic structure, and historical demography of a pioneer tree species, Asian white birch (Betula platyphylla Suk.) across East Asia. We found a north‐to‐south trend of declining genetic diversity and five clusters, corresponding to geographical regions. Different clusters were inferred to have diverged through Pleistocene climatic oscillations and have different expansion routes, leading to genetic admixture in some populations. Ecological niche models indicated that the distribution of B. platyphylla during the last glacial maximum still had a large latitude span with slight shifts toward southeast, and northern populations had more variable distribution ranges than those in the south during later climatic oscillations. Our results reflect the relatively stable distribution through the last glacial–interglacial cycles and recent multidirectional expansion of B. platyphylla, providing new hypotheses for the response pattern of widespread tree species to climate change. The gradual genetic pattern from northeast to southwest and alternative distribution dynamics possibly resulted from environmental differences caused by latitude and topographic heterogeneity.     

Modelling biome shifts and tree cover change for 2050 in West Africa

Aim Africa is expected to face severe changes in climatic conditions. Our objectives are: (1) to model trends and the extent of future biome shifts that may occur by 2050, (2) to model a trend in tree cover change, while accounting for human impact, and (3) to evaluate uncertainty in future climate projections. Location West Africa. Methods We modelled the potential future spatial distribution of desert, grassland, savanna, deciduous and evergreen forest in West Africa using six bioclimatic models. Future tree cover change was analysed with generalized additive models (GAMs). We used climate data from 17 general circulation models (GCMs) and included human population density and fire intensity to model tree cover. Consensus projections were derived via weighted averages to: (1) reduce inter‐model variability, and (2) describe trends extracted from different GCM projections. Results The strongest predicted effect of climate change was on desert and grasslands, where the bioclimatic envelope of grassland is projected to expand into the desert by an area of 2 million km². While savannas are predicted to contract in the south (by 54 ± 22 × 10⁴ km²), deciduous and evergreen forest biomes are expected to expand (64 ± 13 × 10⁴ km² and 77 ± 26 × 10⁴ km²). However, uncertainty due to different GCMs was particularly high for the grassland and the evergreen biome shift. Increasing tree cover (1–10%) was projected for large parts of Benin, Burkina Faso, Côte d’Ivoire, Ghana and Togo, but a decrease was projected for coastal areas (1–20%). Furthermore, human impact negatively affected tree cover and partly changed the direction of the projected change from increase to decrease. Main conclusions Considering climate change alone, the model results of potential vegetation (biomes) show a ‘greening’ trend by 2050. However, the modelled effects of human impact suggest future forest degradation. Thus, it is essential to consider both climate change and human impact in order to generate realistic future tree cover projections.     

The relative roles of environment and history as controls of tree species composition and richness in Europe

Aim This study investigates the determinants of European‐scale patterns in tree species composition and richness, addressing the following questions: (1) What is the relative importance of environment and history? History refers to lasting effects of past large‐scale events and time‐dependent cumulative effects of ongoing processes, notably dispersal limited range dynamics. (2) Among the environmental determinants, what is the relative importance of climate, soils, and forest cover? (3) Do the answers to questions 1 and 2 differ between conifers and Fagales, the two major monophyletic groups of European trees? Location The study area comprises most of Europe (34° N–72° N and 11° W–32° E). Methods Atlas data on native distributions of 54 large tree species at 50 × 50 km resolution were linked with climatic, edaphic, and forest cover maps in a geographical information system. Unconstrained (principal components analysis using Hellinger distance transformation and detrended correspondence analysis) and constrained ordinations (redundancy analysis using Hellinger distance transformation and canonical correspondence analysis) and multiple linear regressions were used to investigate the determinants of species composition and species richness, respectively. History is expected to leave its mark as broad spatial patterns and was represented by the nine spatial terms of a cubic trend surface polynomial. Results The main floristic pattern identified by all ordinations was a latitude‐temperature gradient, while the lower axes corresponded mostly to spatial variables. Partitioning the floristic variation using constrained ordinations showed the mixed spatial‐environmental and pure spatial fractions to be much greater than the pure environmental fraction. Biplots, forward variable selection, and partial analyses all suggested climatic variables as more important floristic determinants than forest cover or soil variables. Tree species richness peaked in the mountainous regions of East‐Central and Southern Europe, except the Far West. Variation partitioning of species richness found the mixed spatial‐environmental and pure spatial fractions to be much greater than the pure environmental fraction for all species combined and Fagales, but not for conifers. The scaled regression coefficients indicated climate as a stronger determinant of richness than soils or forest cover. While the dominant patterns were similar for conifers and Fagales, conifers exhibited less predictable patterns overall, a smaller pure spatial variation fraction relative to pure environmental fraction, and a greater relative importance of climate; all differences being more pronounced for species richness than for species composition. Main conclusions The analyses suggest that history is at least as important as current environment in controlling species composition and richness of European trees, with the exception of conifer species richness. Strong support for interpreting the spatial patterns as outcomes of historical processes, notably dispersal limitation, came from the observation that many European tree species naturalize extensively outside their native ranges. Furthermore, it was confirmed that climate predominates among environmental determinants of distribution and diversity patterns at large spatial scales. Finally, the particular patterns exhibited by conifers probably reflect greater environmental specialization and greater human impact. These findings warn against expecting the European tree flora to be able track fast future climate changes on its own.     

Scale dependency of metapopulation models used to predict climate change impacts on small mammals

To investigate potential range shifts in a changing climate it is becoming increasingly common to develop models that account for demographic processes. Metapopulation models incorporate the spatial configuration of occupied habitat (i.e. arrangement, size and quality), population demographics, and inter‐patch dispersal making them suitable for investigating potential threats to small mammal range and abundance. However, the spatial scale (resolution) used to represent species–environment dynamics may affect estimates of range shift and population resilience by failing to realistically represent the spatial configuration of suitable habitat, including stepping stones and refugia. We aimed to determine whether relatively fine‐scale environmental information influenced predictions of metapopulation persistence and range shift. Species distribution models were constructed for four small terrestrial mammals from southern Australia using environmental predictors measured at 0.1 × 0.1 km (0.01 km²) or 1.0 × 1.0 km (1 km²) resolution, and combined with demographic information to parameterise coupled niche‐population models. These models were used to simulate population dynamics projected over 40‐yr under a stable and changing climate. Initial estimates of the area of available habitat were similar at both spatial scales. However, at the fine‐scale, habitat configuration comprised a greater number of patches (ca 12 times), that were more irregular in shape (ca 8 times the perimeter:area), and separated by a tenth of the distance than at the coarse‐scale. While small patches were not more prone to extinction, populations generally declined at a higher rate and were associated with a lower expected minimum abundance. Despite increased species vulnerability at the fine‐scale, greater range shifts were measured at the coarse‐scale (for species illustrating a shift at both scales). These results highlight the potential for range shifts and species vulnerability information to be misrepresented if advanced modelling techniques incorporating species demographics and dispersal inadequately represent the scale at which these processes occur.     

An ecoregion assessment of projected tree species vulnerabilities in western North America through the 21st century

Forest ecosystems across western North America will likely see shifts in both tree species dominance and composition over the rest of this century in response to climate change. Our objective in this study was to identify which ecological regions might expect the greatest changes to occur. We used the process‐based growth model 3‐PG, to provide estimates of tree species responses to changes in environmental conditions and to evaluate the extent that species are resilient to shifts in climate over the rest of this century. We assessed the vulnerability of 20 tree species in western North America using the Canadian global circulation model under three different emission scenarios. We provided detailed projections of species shifts by including soil maps that account for the spatial variation in soil water availability and soil fertility as well as by utilizing annual climate projections of monthly changes in air temperature, precipitation, solar radiation, vapor pressure deficit and frost at a spatial resolution of one km. Projected suitable areas for tree species were compared to their current ranges based on observations at >40 000 field survey plots. Tree species were classified as vulnerable if environmental conditions projected in the future appear outside that of their current distribution ≥70% of the time. We added a migration constraint that limits species dispersal to <200 m yr^?1 to provide more realistic projections on species distributions. Based on these combinations of constraints, we predicted the greatest changes in the distribution of dominant tree species to occur within the Northwest Forested Mountains and the highest number of tree species stressed will likely be in the North American Deserts. Projected climatic changes appear especially unfavorable for species in the subalpine zone, where major shifts in composition may lead to the emergence of new forest types.