Potential effects of climate change on ecosystem and tree species distribution in British Columbia

A new ecosystem-based climate envelope modeling approach was applied to assess potential climate change impacts on forest communities and tree species. Four orthogonal canonical discriminant functions were used to describe the realized climate space for British Columbia's ecosystems and to model portions of the realized niche space for tree species under current and predicted future climates. This conceptually simple model is capable of predicting species ranges at high spatial resolutions far beyond the study area, including outlying populations and southern range limits for many species. We analyzed how the realized climate space of current ecosystems changes in extent, elevation, and spatial distribution under climate change scenarios and evaluated the implications for potential tree species habitat. Tree species with their northern range limit in British Columbia gain potential habitat at a pace of at least 100 km per decade, common hardwoods appear to be generally unaffected by climate change, and some of the most important conifer species in British Columbia are expected to lose a large portion of their suitable habitat. The extent of spatial redistribution of realized climate space for ecosystems is considerable, with currently important sub-boreal and montane climate regions rapidly disappearing. Local predictions of changes to tree species frequencies were generated as a basis for systematic surveys of biological response to climate change.     

Modelling phytoclimatic versatility as a large scale indicator of adaptive capacity to climate change in forest ecosystems

CLIMPAIR is a new phytoclimatic model, correlative and niche-based, which simultaneously assesses non-linear, non-statistical and dual measurements of proximity/potentiality of a site with respect to a number of climatic ranges of species, defined by convex hulls, within a suitability space. This set of phytoclimatic distances makes it possible to evaluate the degree to which each species is suitable for that site. Considering not only the number of species compatible (expected species richness), but also all those compatible covers presenting a high level of suitability evenness and finally applying an indicator derived from Shannon's classic entropy index to the set of standardized phytoclimatic coordinates in the suitability hyperspace, we can evaluate the phytoclimatic entropy which may be considered as a means of estimating the phytoclimatic versatility of the site. A site with high phytoclimatic entropy would promise versatile future behaviour, characterized by a wide range of possibilities of adaptation to climate change, and hence versatility can be used as an index of resilience and ability of a forest ecosystem to adapt to climate change. The model has been applied to peninsular Spain for 18 forest tree species and 12 climatic variables between the current mean climate (period 1951-1999) and a future climatic scenario (period 2040-2069). The results generally point to a significant decrease in the versatility of forest tree formations in the area studied, which is not homogeneous owing to a dual altitudinal/latitudinal decoupling. The decrease in versatility is greater in Mediterranean biogeographical areas than in Euro-Siberian ones, where in some cases it actually increases. In altitudinal terms, areas at elevations of less than 1500 m tend to become less versatile than areas situated at higher elevations, where versatility increases partly as a result of enrichment of alpine conifer forests with broadleaf species.     

The influence of land use and climate change on forest biomass and composition in Massachusetts, USA

Land use and climate change have complex and interacting effects on naturally dynamic forest landscapes. To anticipate and adapt to these changes, it is necessary to understand their individual and aggregate impacts on forest growth and composition. We conducted a simulation experiment to evaluate regional forest change in Massachusetts, USA over the next 50 years (2010-2060). Our objective was to estimate, assuming a linear continuation of recent trends, the relative and interactive influence of continued growth and succession, climate change, forest conversion to developed uses, and timber harvest on live aboveground biomass (AGB) and tree species composition. We examined 20 years of land use records in relation to social and biophysical explanatory variables and used regression trees to create "probability-of-conversion" and "probability-of-harvest" zones. We incorporated this information into a spatially interactive forest landscape simulator to examine forest dynamics as they were affected by land use and climate change. We conducted simulations in a full-factorial design and found that continued forest growth and succession had the largest effect on AGB, increasing stores from 181.83 Tg to 309.56 Tg over 50 years. The increase varied from 49% to 112% depending on the ecoregion within the state. Compared to simulations with no climate or land use, forest conversion reduced gains in AGB by 23.18 Tg (or 18%) over 50 years. Timber harvests reduced gains in AGB by 5.23 Tg (4%). Climate change (temperature and precipitation) increased gains in AGB by 17.3 Tg (13.5%). Pinus strobus and Acer rubrum were ranked first and second, respectively, in terms of total AGB throughout all simulations. Climate change reinforced the dominance of those two species. Timber harvest reduced Quercus rubra from 10.8% to 9.4% of total AGB, but otherwise had little effect on composition. Forest conversion was generally indiscriminate in terms of species removal. Under the naive assumption that future land use patterns will resemble the recent past, we conclude that continued forest growth and recovery will be the dominant mechanism driving forest dynamics over the next 50 years, and that while climate change may enhance growth rates, this will be more than offset by land use, primarily forest conversion to developed uses.     

Site-specific and integrated adaptation to climate change in the coastal mangrove zone of Soc Trang Province, Viet Nam

The dynamic coastline of Soc Trang Province in the Mekong Delta of Viet Nam is in most parts protected from erosion, storms and flooding by a narrow belt of mangroves. However, the unsustainable use of natural resources and development in the coastal zone is threatening the protection function of this forest belt. This situation is exacerbated by the impacts of climate change, particularly by the increased intensity and frequency of storms, floods and by rising sea levels. Based on analysis of past experience of mangrove planting and historical changes in mangrove cover, an integrated and site-specific approach to adaptation to climate change has been put in place, which comprises mangrove planting and rehabilitation with emphasis on resilience to climate change, and participatory involvement of local communities in effective mangrove management and protection through co-management. To address uncertainties associated with the impacts of climate change, testing of new mangrove planting techniques has started. This includes mimicking successful natural regeneration for small-scale planting in sites with high wave energy and transformation of existing even-aged plantations into more diverse forests—both in terms of structure and species composition. The pre-requisite for mangrove rehabilitation in erosion sites has successfully been put in place: breakwaters made from bamboo have reduced erosion and stimulated sedimentation. The design and construction of the wave-breaking structures, which was based on a numerical model which simulates hydrodynamics and shoreline development, ensures that downdrift erosion can be avoided as far as possible. A comprehensive monitoring program has been established and initial results provide evidence for the effectiveness of the bamboo breakwaters. Early experience shows that co-management is an effective way of maintaining and enhancing the protection function of the mangrove forest belt and at the same time providing livelihood for local communities. Payment for ecosystem services contributes to sustainability of co-management as well as livelihood improvement.     

Frequency, not relative abundance, of temperate tree species varies along climate gradients in eastern North America

There have been many attempts to model the impacts of climate change on the distributions of temperate tree species, but empirical analyses of the effects of climate on the distribution and abundance of tree species have lagged far behind the models. Here, we used forest inventory data to characterize variation in adult tree abundance along climate gradients for the 24 most common tree species in the northeastern United States. The two components of our measure of species abundance--local frequency vs. relative abundance--showed dramatically different patterns of variation along gradients of mean annual temperature and precipitation. Local frequency (i.e., the percentage of plots in a given climate in which a species occurred) varied strongly for all 24 species, particularly as a function of temperature. Relative abundance when present in a plot, on the other hand, was effectively constant for most species right up to their estimated climatic range limits. Although the range limits for both temperature and precipitation were quite broad for all of the species, the range of climates within which a species was common (i.e., high frequency) was much narrower. Because frequency in sites within a given climate shows a strong sensitivity to temperature, at least, this suggests that the processes determining canopy tree recruitment on new sites also vary strongly with climate.     

Incorporating geodiversity into conservation decisions

In a rapidly changing climate, conservation practitioners could better use geodiversity in a broad range of conservation decisions. We explored selected avenues through which this integration might improve decision making and organized them within the adaptive management cycle of assessment, planning, implementation, and monitoring. Geodiversity is seldom referenced in predominant environmental law and policy. With most natural resource agencies mandated to conserve certain categories of species, agency personnel are challenged to find ways to practically implement new directives aimed at coping with climate change while retaining their species‐centered mandate. Ecoregions and ecological classifications provide clear mechanisms to consider geodiversity in plans or decisions, the inclusion of which will help foster the resilience of conservation to climate change. Methods for biodiversity assessment, such as gap analysis, climate change vulnerability analysis, and ecological process modeling, can readily accommodate inclusion of a geophysical component. We adapted others’ approaches for characterizing landscapes along a continuum of climate change vulnerability for the biota they support from resistant, to resilient, to susceptible, and to sensitive and then summarized options for integrating geodiversity into planning in each landscape type. In landscapes that are relatively resistant to climate change, options exist to fully represent geodiversity while ensuring that dynamic ecological processes can change over time. In more susceptible landscapes, strategies aiming to maintain or restore ecosystem resilience and connectivity are paramount. Implementing actions on the ground requires understanding of geophysical constraints on species and an increasingly nimble approach to establishing management and restoration goals. Because decisions that are implemented today will be revisited and amended into the future, increasingly sophisticated forms of monitoring and adaptation will be required to ensure that conservation efforts fully consider the value of geodiversity for supporting biodiversity in the face of a changing climate.     

Linking species- and ecosystem-level impacts of climate change in lakes with a complex and a minimal model

To study the interaction between species- and ecosystem-level impacts of climate change, we focus on the question of how climate-induced shifts in key species affect the positive feedback loops that lock shallow lakes either in a transparent, macrophyte-dominated state or, alternatively, in a turbid, phytoplankton-dominated state. We hypothesize that climate warming will weaken the resilience of the macrophyte-dominated clear state. For the turbid state, we hypothesize that climate warming and climate-induced eutrophication will increase the dominance of cyanobacteria. Climate change will also affect shallow lakes through a changing hydrology and through climate change-induced eutrophication. We study these phenomena using two models, the full ecosystem model PCLake and a minimal dynamic model of lake phosphorus dynamics. Quantitative predictions with the complex model show that changes in nutrient loading, hydraulic loading and climate warming can all lead to shifts in ecosystem state. The minimal model helped in interpreting the non-linear behaviour of the complex model. The main output parameters of interest for water quality managers are the critical nutrient loading at which the system will switch from clear to turbid and the much lower critical nutrient loading – due to hysteresis – at which the system switches back. Another important output parameter is the chlorophyll-a level in the turbid state. For each of these three output parameters we performed a sensitivity analysis to further understand the dynamics of the complex model PCLake. This analysis showed that our model results are most sensitive to changes in temperature-dependence of cyanobacteria, planktivorous fish and zooplankton. We argue that by combining models at various levels of complexity and looking at multiple aspects of climate changes simultaneously we can develop an integrated view of the potential impact of climate change on freshwater ecosystems.     

Tropical amphibians in shifting thermal landscapes under land‐use and climate change

Land‐cover and climate change are both expected to alter species distributions and contribute to future biodiversity loss. However, the combined effects of land‐cover and climate change on assemblages, especially at the landscape scale, remain understudied. Lowland tropical amphibians may be particularly susceptible to changes in land cover and climate warming because many species have narrow thermal safety margins resulting from air and body temperatures that are close to their critical thermal maxima (CT_max). We examined how changing thermal landscapes may alter the area of thermally suitable habitat (TSH) for tropical amphibians. We measured microclimates in 6 land‐cover types and CT_max of 16 frog species in lowland northeastern Costa Rica. We used a biophysical model to estimate core body temperatures of frogs exposed to habitat‐specific microclimates while accounting for evaporative cooling and behavior. Thermally suitable habitat area was estimated as the portion of the landscape where species CT_max exceeded their habitat‐specific maximum body temperatures. We projected changes in TSH area 80 years into the future as a function of land‐cover change only, climate change only, and combinations of land‐cover and climate‐change scenarios representing low and moderate rates of change. Projected decreases in TSH area ranged from 16% under low emissions and reduced forest loss to 30% under moderate emissions and business‐as‐usual land‐cover change. Under a moderate emissions scenario (A1B), climate change alone contributed to 1.7‐ to 4.5‐fold greater losses in TSH area than land‐cover change only, suggesting that future decreases in TSH from climate change may outpace structural habitat loss. Forest‐restricted species had lower mean CT_max than species that occurred in altered habitats, indicating that thermal tolerances will likely shape assemblages in changing thermal landscapes. In the face of ongoing land‐cover and climate change, it will be critical to consider changing thermal landscapes in strategies to conserve ectotherm species.     

温带落叶林的植物物候特征及其对气候变化的响应

植被物候是气候变化对生物圈产生长期或短期影响的重要指示因子。气候变化已经明显改变了许多物种的营养生长和繁殖物候,尤其是在温带地区。研究温带森林物候变化及其对全球变暖的响应,对认识森林物种共存,协同进化以及森林保护和经营等有重要意义。通过概述温带森林下物候研究的进展发现,光照和积温是影响木本植物展叶及繁殖物候的关键因素,林下层树木通过更早展叶,以尽量减少生长季林冠层遮阴对下层树木生长的影响,更早时期开花的树木具有从顶部向四周次第开花的时空格局,林冠层树种开花具有较好的同步性。而草本植物的物候通常受融雪时间和冠层动态的影响更大,并且,温带森林下不同生活史对策的草本植物的物候特征对气候变化的响应也不尽相同,存在明显的季节动态。繁殖物候、光照的季节变化、光合特征、授粉成功之间的联系决定了林下不同繁殖特性的草本植物的繁殖成功率。量化的、多指标、多对象的定位监测是精准物候研究的基础,物候变化的机理和建立可预测的物候模型将是未来研究的重点。     

Phenological tracking enables positive species responses to climate change

Earlier spring phenology observed in many plant species in recent decades provides compelling evidence that species are already responding to the rising global temperatures associated with anthropogenic climate change. There is great variability among species, however, in their phenological sensitivity to temperature. Species that do not phenologically "track" climate change may be at a disadvantage if their growth becomes limited by missed interactions with mutualists, or a shorter growing season relative to earlier-active competitors. Here, we set out to test the hypothesis that phenological sensitivity could be used to predict species performance in a warming climate, by synthesizing results across terrestrial warming experiments. We assembled data for 57 species across 24 studies where flowering or vegetative phenology was matched with a measure of species performance. Performance metrics included biomass, percent cover, number of flowers, or individual growth. We found that species that advanced their phenology with warming also increased their performance, whereas those that did not advance tended to decline in performance with warming. This indicates that species that cannot phenologically "track" climate may be at increased risk with future climate change, and it suggests that phenological monitoring may provide an important tool for setting future conservation priorities.     

Antagonistic effect of natural habitat conversion on community adjustment to climate warming in nonbreeding waterbirds

Although the impacts of climate and land-use changes on biodiversity have been widely documented, their joint effects remain poorly understood. We evaluated how nonbreeding waterbird communities adjust to climate warming along a gradient of land-use change. Using midwinter waterbird counts (132 species) at 164 major nonbreeding sites in 22 Mediterranean countries, we assessed the changes in species composition from 1991 to 2010, relative to thermal niche position and breadth, in response to regional and local winter temperature anomalies and conversion of natural habitats. We observed a low-level, nonsignificant community adjustment to the temperature increase where natural habitat conversion occurred. At the sites affected by natural habitat conversion, the relative increase of warm-dwelling species in response to climate warming was 6 times lower and the relative species decline was 3 times higher than in the sites without natural habitat conversion. We found no evidence of community adjustment to climate warming when natural habitat conversion was >5% over 15 years. This strong negative effect suggests an antagonistic interaction between climate warming and habitat change. These results underline the importance of habitat conservation in community adjustment to climate warming.     

Sensitivity of plant-pollinator-herbivore communities to changes in phenology

For many species in seasonal environments, warmer springs associated with anthropogenic climate change are causing phenological changes. Within ecological communities, the timing of interactions among species may be altered if the species experience asymmetrical phenological shifts. We present a model that examines the consequences of changes in the relative timing of herbivory and pollination in a community of herbivores and pollinators that share a host plant. Our model suggests that phenological shifts can alter the abundances of these species and, in some cases, their population dynamics. If historical patterns of interactions in a community change and herbivores become active before pollinators, the community could see a reduction in pollinators and an increase in herbivores, while if pollinators become active before herbivores, there could be a loss of stable coexistence. Previous studies have warned of the potential for climate change to cause large phenological mismatches whereby species that depend on one another become so separated in time that they can no longer interact. Our results suggest that climate change-induced phenological shifts can have a major impact on communities even in cases where complete phenological mismatches do not occur.     

Trophic Flexibility and the Persistence of Understory Birds in Intensively Logged Rainforest

Effects of logging on species composition in tropical rainforests are well known but may fail to reveal key changes in species interactions. We used nitrogen stable‐isotope analysis of 73 species of understory birds to quantify trophic responses to repeated intensive logging of rainforest in northern Borneo and to test 4 hypotheses: logging has significant effects on trophic positions and trophic‐niche widths of species, and the persistence of species in degraded forest is related to their trophic positions and trophic‐niche widths in primary forest. Species fed from higher up the food chain and had narrower trophic‐niche widths in degraded forest. Species with narrow trophic‐niche widths in primary forest were less likely to persist after logging, a result that indicates a higher vulnerability of dietary specialists to local extinction following habitat disturbance. Persistence of species in degraded forest was not related to a species’ trophic position. These results indicate changes in trophic organization that were not apparent from changes in species composition and highlight the importance of focusing on trophic flexibility over the prevailing emphasis on membership of static feeding guilds. Our results thus support the notion that alterations to trophic organization and interactions within tropical forests may be a pervasive and functionally important hidden effect of forest degradation. Flexibilidad Trófica y la Persistencia de Aves de Sotobosque en un Bosque Lluvioso Talado Intensivamente     

Applying assessments of adaptive capacity to inform natural-resource management in a changing climate

Adaptive capacity (AC)—the ability of a species to cope with or accommodate climate change—is a critical determinant of species vulnerability. Using information on species’ AC in conservation planning is key to ensuring successful outcomes. We identified connections between a list of species’ attributes (e.g., traits, population metrics, and behaviors) that were recently proposed for assessing species’ AC and management actions that may enhance AC for species at risk of extinction. Management actions were identified based on evidence from the literature, a review of actions used in other climate adaptation guidance, and our collective experience in diverse fields of global-change ecology and climate adaptation. Selected management actions support the general AC pathways of persist in place or shift in space, in response to contemporary climate change. Some actions, such as genetic manipulations, can be used to directly alter the ability of species to cope with climate change, whereas other actions can indirectly enhance AC by addressing ecological or anthropogenic constraints on the expression of a species’ innate abilities to adapt. Ours is the first synthesis of potential management actions directly linked to AC. Focusing on AC attributes helps improve understanding of how and why aspects of climate are affecting organisms, as well as the mechanisms by which management interventions affect a species’ AC and climate change vulnerability. Adaptive-capacity-informed climate adaptation is needed to build connections among the causes of vulnerability, AC, and proposed management actions that can facilitate AC and reduce vulnerability in support of evolving conservation paradigms.     

Species' traits predict phenological responses to climate change in butterflies

How do species' traits help identify which species will respond most strongly to future climate change? We examine the relationship between species' traits and phenology in a well-established model system for climate change, the U.K. Butterfly Monitoring Scheme (UKBMS). Most resident U.K. butterfly species have significantly advanced their dates of first appearance during the past 30 years. We show that species with narrower larval diet breadth and more advanced overwintering stages have experienced relatively greater advances in their date of first appearance. In addition, species with smaller range sizes have experienced greater phenological advancement. Our results demonstrate that species' traits can be important predictors of responses to climate change, and they suggest that further investigation of the mechanisms by which these traits influence phenology may aid in understanding species' responses to current and future climate change.     

Predicting extinctions as a result of climate change

Widespread extinction is a predicted ecological consequence of global warming. Extinction risk under climate change scenarios is a function of distribution breadth. Focusing on trees and birds of the eastern United States, we used joint climate and environment models to examine fit and climate change vulnerability as a function of distribution breadth. We found that extinction vulnerability increases with decreasing distribution size. We also found that model fit decreases with decreasing distribution size, resulting in high prediction uncertainty among narrowly distributed species. High prediction uncertainty creates a conservation dilemma in that excluding these species under-predicts extinction risk and favors mistaken inaction on global warming. By contrast, including narrow endemics results in over-predicting extinction risk and promotes mistaken inaction on behalf of individual species prematurely considered doomed to extinction.     

A model of surface fire,climate and forest pattern in the Sierra Nevada,California

A spatially explicit forest gap model was developed for the Sierra Nevada, California, and is the first of its kind because it integrates climate, fire and forest pattern. The model simulates a forest stand as a grid of 15×15 m forest plots and simulates the growth of individual trees within each plot. Fuel inputs are generated from each individual tree according to tree size and species. Fuel moisture varies both temporally and spatially with the local site water balance and forest condition, thus linking climate with the fire regime. Fires occur as a function of the simulated fuel loads and fuel moisture, and the burnable area is simulated as a result of the spatially heterogeneous fuel bed conditions. We demonstrate the model’s ability to couple the fire regime to both climate and forest pattern. In addition, we use the model to investigate the importance of climate and forest pattern as controls on the fire regime. Comparison of model results with independent data indicate that the model performs well in several areas. Patterns of fuel accumulation, climatic control of fire frequency and the influence of fuel loads on the spatial extent of fires in the model are particularly well-supported by data. This model can be used to examine the complex interactions among climate, fire and forest pattern across a wide range of environmental conditions and vegetation types. Our results suggest that, in the Sierra Nevada, fuel moisture can exert an important control on fire frequency and this control is especially pronounced at sites where most of the annual precipitation is in the form of snow. Fuel loads, on the other hand, may limit the spatial extent of fire, especially at elevations below 1500 m. Above this elevation, fuel moisture may play an increasingly important role in limiting the area burned.     

Bird extirpations and community dynamics in an Andean cloud forest over 100 years of land-use change

Long-term studies to understand biodiversity changes remain scarce—especially so for tropical mountains. We examined changes from 1911 to 2016 in the bird community of the cloud forest of San Antonio, a mountain ridge in the Colombian Andes. We evaluated the effects of past land-use change and assessed species vulnerability to climate disruption. Forest cover decreased from 95% to 50% by 1959, and 33 forest species were extirpated. From 1959 to 1990, forest cover remained stable, and an additional 15 species were lost—a total of 29% of the forest bird community. Thereafter, forest cover increased by 26% and 17 species recolonized the area. The main cause of extirpations was the loss of connections to adjacent forests. Of the 31 (19%) extirpated birds, 25 have ranges peripheral to San Antonio, mostly in the lowlands. Most still occurred regionally, but broken forest connections limited their recolonization. Other causes of extirpation were hunting, wildlife trade, and water diversion. Bird community changes included a shift from predominantly common species to rare species; forest generalists replaced forest specialists that require old growth, and functional groups, such as large-body frugivores and nectarivores, declined disproportionally. All water-dependent birds were extirpated. Of the remaining 122 forest species, 19 are vulnerable to climate disruption, 10 have declined in abundance, and 4 are threatened. Our results show unequivocal species losses and changes in community structure and abundance at the local scale. We found species were extirpated after habitat loss and fragmentation, but forest recovery stopped extirpations and helped species repopulate. Land-use changes increased species vulnerability to climate change, and we suggest reversing landscape transformation may restore biodiversity and improve resistance to future threats.     

Response of tree growth to a changing climate in boreal central Canada: A comparison of empirical,process-based,and hybrid modelling approaches

The impact of 2 × CO₂ driven climate change on radial growth of boreal tree species Pinus banksiana Lamb., Populus tremuloides Michx. and Picea mariana (Mill.) BSP growing in the Duck Mountain Provincial Forest of Manitoba (DMPF), Canada, is simulated using empirical and process-based model approaches. First, empirical relationships between growth and climate are developed. Stepwise multiple-regression models are conducted between tree-ring growth increments (TRGI) and monthly drought, precipitation and temperature series. Predictive skills are tested using a calibration–verification scheme. The established relationships are then transferred to climates driven by 1× and 2 × CO₂ scenarios using outputs from the Canadian second-generation coupled global climate model. Second, empirical results are contrasted with process-based projections of net primary productivity allocated to stem development (NPP_s). At the finest scale, a leaf-level model of photosynthesis is used to simulate canopy properties per species and their interaction with the variability in radiation, temperature and vapour pressure deficit. Then, a top-down plot-level model of forest productivity is used to simulate landscape-level productivity by capturing the between-stand variability in forest cover. Results show that the predicted TRGI from the empirical models account for up to 56.3% of the variance in the observed TRGI over the period 1912–1999. Under a 2 × CO₂ scenario, the predicted impact of climate change is a radial growth decline for all three species under study. However, projections obtained from the process-based model suggest that an increasing growing season length in a changing climate could counteract and potentially overwhelm the negative influence of increased drought stress. The divergence between TRGI and NPP_s simulations likely resulted, among others, from assumptions about soil water holding capacity and from calibration of variables affecting gross primary productivity. An attempt was therefore made to bridge the gap between the two modelling approaches by using physiological variables as TRGI predictors. Results obtained in this manner are similar to those obtained using climate variables, and suggest that the positive effect of increasing growing season length would be counteracted by increasing summer temperatures. Notwithstanding uncertainties in these simulations (CO₂ fertilization effect, feedback from disturbance regimes, phenology of species, and uncertainties in future CO₂ emissions), a decrease in forest productivity with climate change should be considered as a plausible scenario in sustainable forest management planning of the DMPF.