What is the role of disturbance in catalyzing spatial shifts in forest composition and tree species biomass under climate change?

Mounting evidence suggests that climate change will cause shifts of tree species range and abundance (biomass). Abundance changes under climate change are likely to occur prior to a detectable range shift. Disturbances are expected to directly affect tree species abundance and composition, and could profoundly influence tree species spatial distribution within a geographical region. However, how multiple disturbance regimes will interact with changing climate to alter the spatial distribution of species abundance remains unclear. We simulated such forest demographic processes using a forest landscape succession and disturbance model (LANDIS-II) parameterized with forest inventory data in the northeastern United States. Our study incorporated climate change under a high-emission future and disturbance regimes varying with gradients of intensities and spatial extents. The results suggest that disturbances catalyze changes in tree species abundance and composition under a changing climate, but the effects of disturbances differ by intensity and extent. Moderate disturbances and large extent disturbances have limited effects, while high-intensity disturbances accelerate changes by removing cohorts of mid- and late-successional species, creating opportunities for early-successional species. High-intensity disturbances result in the northern movement of early-successional species and the southern movement of late-successional species abundances. Our study is among the first to systematically investigate how disturbance extent and intensity interact to determine the spatial distribution of changes in species abundance and forest composition.

Forecasting effects of tree species reintroduction strategies on carbon stocks in a future without historical analog.

American chestnut (Castanea dentata) was once an important component forests in the central Appalachians (USA), but it was functionally extirpated nearly a century ago. Attempts are underway to reintroduce blight-resistant chestnut to its former range, but it is uncertain how current forest composition, climate, and atmospheric changes and disturbance regimes will interact to determine future forest dynamics and ecosystem services. The combination of novel environmental conditions (e.g. climate change), a reintroduced tree species and new disturbance regimes (e.g. exotic insect pests, fire suppression) have no analog in the past that can be used to parameterize phenomenological models. We therefore used a mechanistic approach within the LANDIS-II forest landscape model that relies on physiological first principles to project forest dynamics as the outcome of competition of tree cohorts for light and water as a function of temperature, precipitation, CO2 concentration, and life history traits. We conducted a factorial landscape simulation experiment to evaluate specific hypotheses about future forest dynamics in two study sites in the center of the former range of chestnut. Our results supported the hypotheses that climate change would favor chestnut because of its optimal temperature range and relative drought resistance, and that chestnut would be less competitive in the more mesic Appalachian Plateau province because competitors will be less stressed. The hypothesis that chestnut will increase carbon stocks was supported, although the increase was modest. Our results confirm that aggressive restoration is needed regardless of climate and soils, and that increased aggressiveness of chestnut restoration increased biomass accumulation. The hypothesis that chestnut restoration will increase both compositional and structural richness was not supported because chestnut displaced some species and age cohorts. Although chestnut restoration did not markedly enhance carbon stocks, our findings provide hope that this formerly important species can be successfully reintroduced and associated ecosystem services recovered.

How disturbance, competition, and dispersal interact to prevent tree range boundaries from keeping pace with climate change.

Climate change is expected to cause geographic shifts in tree species' ranges, but such shifts may not keep pace with climate changes because seed dispersal distances are often limited and competition-induced changes in community composition can be relatively slow. Disturbances may speed changes in community composition, but the interactions among climate change, disturbance and competitive interactions to produce range shifts are poorly understood. We used a physiologically based mechanistic landscape model to study these interactions in the northeastern United States. We designed a series of disturbance scenarios to represent varied disturbance regimes in terms of both disturbance extent and intensity. We simulated forest succession by incorporating climate change under a high-emissions future, disturbances, seed dispersal, and competition using the landscape model parameterized with forest inventory data. Tree species range boundary shifts in the next century were quantified as the change in the location of the 5th (the trailing edge) and 95th (the leading edge) percentiles of the spatial distribution of simulated species. Simulated tree species range boundary shifts in New England over the next century were far below (usually <20 km) that required to track the velocity of temperature change (usually more than 110 km over 100 years) under a high-emissions scenario. Simulated species` ranges shifted northward at both the leading edge (northern boundary) and trailing edge (southern boundary). Disturbances may expedite species' recruitment into new sites, but they had little effect on the velocity of simulated range boundary shifts. Range shifts at the trailing edge tended to be associated with photosynthetic capacity, competitive ability for light and seed dispersal ability, whereas shifts at the leading edge were associated only with photosynthetic capacity and competition for light. This study underscores the importance of understanding the role of interspecific competition and disturbance when studying tree range shifts.

Integrating ecophysiology and forest landscape models to improve projections of drought effects under climate change.

Fundamental drivers of ecosystem processes such as temperature and precipitation are rapidly changing and creating novel environmental conditions. Forest landscape models (FLM) are used by managers and policy-makers to make projections of future ecosystem dynamics under alternative management or policy options, but the links between the fundamental drivers and projected responses are weak and indirect, limiting their reliability for projecting the impacts of climate change. We developed and tested a relatively mechanistic method to simulate the effects of changing precipitation on species competition within the LANDIS-II FLM. Using data from a field precipitation manipulation experiment in a piñon pine (Pinus edulis) and juniper (Juniperus monosperma) ecosystem in New Mexico (USA), we calibrated our model to measurements from ambient control plots and tested predictions under the drought and irrigation treatments against empirical measurements. The model successfully predicted behavior of physiological variables under the treatments. Discrepancies between model output and empirical data occurred when the monthly time step of the model failed to capture the short-term dynamics of the ecosystem as recorded by instantaneous field measurements. We applied the model to heuristically assess the effect of alternative climate scenarios on the piñon-juniper ecosystem and found that warmer and drier climate reduced productivity and increased the risk of drought-induced mortality, especially for piñon. We concluded that the direct links between fundamental drivers and growth rates in our model hold great promise to improve our understanding of ecosystem processes under climate change and improve management decisions because of its greater reliance on first principles.

SEARCH: Spatially Explicit Animal Response to Composition of Habitat.

Complex decisions dramatically affect animal dispersal and space use. Dispersing individuals respond to a combination of fine-scale environmental stimuli and internal attributes. Individual-based modeling offers a valuable approach for the investigation of such interactions because it combines the heterogeneity of animal behaviors with spatial detail. Most individual-based models (IBMs), however, vastly oversimplify animal behavior and such behavioral minimalism diminishes the value of these models. We present program SEARCH (Spatially Explicit Animal Response to Composition of Habitat), a spatially explicit, individual-based, population model of animal dispersal through realistic landscapes. SEARCH uses values in Geographic Information System (GIS) maps to apply rules that animals follow during dispersal, thus allowing virtual animals to respond to fine-scale features of the landscape and maintain a detailed memory of areas sensed during movement. SEARCH also incorporates temporally dynamic landscapes so that the environment to which virtual animals respond can change during the course of a simulation. Animals in SEARCH are behaviorally dynamic and able to respond to stimuli based upon their individual experiences. Therefore, SEARCH is able to model behavioral traits of dispersing animals at fine scales and with many dynamic aspects. Such added complexity allows investigation of unique ecological questions. To illustrate SEARCH's capabilities, we simulated case studies using three mammals. We examined the impact of seasonally variable food resources on the weight distribution of dispersing raccoons (Procyon lotor), the effect of temporally dynamic mortality pressure in combination with various levels of behavioral responsiveness in eastern chipmunks (Tamias striatus), and the impact of behavioral plasticity and home range selection on disperser mortality and weight change in virtual American martens (Martes americana). These simulations highlight the relevance of SEARCH for a variety of applications and illustrate benefits it can provide for conservation planning.

Comparing modern and presettlement forest dynamics of a subboreal wilderness: does spruce budworm enhance fire risk?

Insect disturbance is often thought to increase fire risk through enhanced fuel loadings, particularly in coniferous forest ecosystems. Yet insect disturbances also affect successional pathways and landscape structure that interact with fire disturbances (and vice-versa) over longer time scales. We applied a landscape succession and disturbance model (LANDIS-II) to evaluate the relative strength of interactions between spruce budworm (Choristoneura fumiferana) outbreaks and fire disturbances in the Boundary Waters Canoe Area (BWCA) in northern Minnesota (USA). Disturbance interactions were evaluated for two different scenarios: presettlement forests and fire regimes vs. contemporary forests and fire regimes. Forest composition under the contemporary scenario trended toward mixtures of deciduous species (primarily Betula papyrifera and Populus spp.) and shade-tolerant conifers (Picea mariana, Abies balsamea, Thuja occidentalis), with disturbances dominated by a combination of budworm defoliation and high-severity fires. The presettlement scenario retained comparatively more "big pines" (i.e., Pinus strobus, P. resinosa) and tamarack (L. laricina), and experienced less budworm disturbance and a comparatively less-severe fire regime. Spruce budworm disturbance decreased area burned and fire severity under both scenarios when averaged across the entire 300-year simulations. Contrary to past research, area burned and fire severity during outbreak decades were each similar to that observed in non-outbreak decades. Our analyses suggest budworm disturbances within forests of the BWCA have a comparatively weak effect on long-term forest composition due to a combination of characteristics. These include strict host specificity, fine-scaled patchiness created by defoliation damage, and advance regeneration of its primary host, balsam fir (A. balsamea) that allows its host to persist despite repeated disturbances. Understanding the nature of the three-way interaction between budworm, fire, and composition has important ramifications for both fire mitigation strategies and ecosystem restoration initiatives. We conclude that budworm disturbance can partially mitigate long-term future fire risk by periodically reducing live ladder fuel within the mixed forest types of the BWCA but will do little to reverse the compositional trends caused in part by reduced fire rotations.

Predicting global change effects on forest biomass and composition in south-central Siberia.

Multiple global changes such as timber harvesting in areas not previously disturbed by cutting and climate change will undoubtedly affect the composition and spatial distribution of boreal forests, which will, in turn, affect the ability of these forests to retain carbon and maintain biodiversity. To predict future states of the boreal forest reliably, it is necessary to understand the complex interactions among forest regenerative processes (succession), natural disturbances (e.g., fire, wind, and insects), and anthropogenic disturbances (e.g., timber harvest). We used a landscape succession and disturbance model (LANDIS-II) to study the relative effects of climate change, timber harvesting, and insect outbreaks on forest composition, biomass (carbon), and landscape pattern in south-central Siberia. We found that most response variables were more strongly influenced by timber harvest and insect outbreaks than by the direct effects of climate change. Direct climate effects generally increased tree productivity and modified probability of establishment, but indirect effects on the fire regime generally counteracted the direct effects of climate on forest composition. Harvest and insects significantly changed forest composition, reduced living aboveground biomass, and increased forest fragmentation. We concluded that: (1) Global change is likely to significantly change forest composition of south-central Siberian landscapes, with some changes taking ecosystems outside the historic range of variability. (2) The direct effects of climate change in the study area are not as significant as the exploitation of virgin forest by timber harvest and the potential increased outbreaks of the Siberian silk moth. (3) Novel disturbance by timber harvest and insect outbreaks may greatly reduce the aboveground living biomass of Siberian forests and may significantly alter ecosystem dynamics and wildlife populations by increasing forest fragmentation.

Linking linear programming and spatial simulation models to predict landscape effects of forest management alternatives.

Forest management planners require analytical tools to assess the effects of alternative strategies on the sometimes disparate benefits from forests such as timber production and wildlife habitat. We assessed the spatial patterns of alternative management strategies by linking two models that were developed for different purposes. We used a linear programming model (Spectrum) to optimize timber harvest schedules, then a simulation model (HARVEST) to project those schedules in a spatially explicit way and produce maps from which the spatial pattern of habitat could be calculated. We demonstrated the power of this approach by evaluating alternative plans developed for a national forest plan revision in Wisconsin, USA. The amount of forest interior habitat was inversely related to the amount of timber cut, and increased under the alternatives compared to the current plan. The amount of edge habitat was positively related to the amount of timber cut, and increased under all alternatives. The amount of mature northern hardwood interior and edge habitat increased for all alternatives, but mature pine habitat area varied. Mature age classes of all forest types increased, and young classes decreased under all alternatives. The average size of patches (defined by age class) generally decreased. These results are consistent with the design goals of each of the alternatives, but reveal that the spatial differences among the alternatives are modest. These complementary models are valuable for quantifying and comparing the spatial effects of alternative management strategies.

Modeling the influence of dynamic zoning of forest harvesting on ecological succession in a northern hardwoods landscape.

Dynamic zoning (systematic alteration in the spatial and temporal allocation of even-aged forest management practices) has been proposed as a means to change the spatial pattern of timber harvest across a landscape to maximize forest interior habitat while holding timber harvest levels constant. Simulation studies have established that dynamic zoning strategies produce larger tracts of interior, closed canopy forest, thus increasing the value of these landscapes for interior-dependent wildlife. We used the simulation model LANDIS to examine how the implementation of a dynamic zoning strategy would change trajectories of ecological succession in the Great Divide Ranger District of the Chequamegon-Nicolet National Forest in northern Wisconsin over 500 years. The components of dynamic zoning strategies (number of zones in a scenario and the length of the hiatus between successive entries into zones) and their interaction had highly significant impacts on patterns of forest succession. Dynamic zoning scenarios with more zones and shorter hiatus lengths increased the average amount of the forest dominated by early successional aspen (Populus sp.). Dynamic zoning scenarios with two zones produced more late successional mature northern hardwoods than scenarios with four zones. Dynamic zoning scenarios with very short (30 years) or very long (120 years) hiatus lengths resulted in more late successional mature northern hardwoods than scenarios with intermediate hiatus lengths (60 and 90 years). However, none of the dynamic scenarios produced as much late successional mature northern hardwoods as the static alternative. Furthermore, the amounts of all habitat types in all dynamic zoning scenarios fluctuated greatly in time and space relative to static alternatives, which could negatively impact wildlife species that require a stable amount of habitat above some minimum critical threshold. Indeed, implementing dynamic zoning scenarios of different designs would have both positive and negative effects on wildlife species and for other objectives of forest management.