When relationships estimated in the past cannot be used to predict the future: using mechanistic models to predict landscape ecological dynamics in a changing world

Researchers and natural resource managers need predictions of how multiple global changes (e.g., climate change, rising levels of air pollutants, exotic invasions) will affect landscape composition and ecosystem function. Ecological predictive models used for this purpose are constructed using either a mechanistic (process-based) or a phenomenological (empirical) approach, or combination. Given the accelerating pace of global changes, it is becoming increasingly difficult to trust future projections made by phenomenological models estimated under past conditions. Using forest landscape models as an example, I review current modeling approaches and propose principles for developing the next generation of landscape models. First, modelers should increase the use of mechanistic components based on appropriately scaled “first principles” even though such an approach is not without cost and limitations. Second, the interaction of processes within a model should be designed to minimize a priori constraints on process interactions and mimic how interactions play out in real life. Third, when a model is expected to make accurate projections of future system states it must include all of the major ecological processes that structure the system. A completely mechanistic approach to the molecular level is not tractable or desirable at landscape scales. I submit that the best solution is to blend mechanistic and phenomenological approaches in a way that maximizes the use of mechanisms where novel driver conditions are expected while keeping the model tractable. There may be other ways. I challenge landscape ecosystem modelers to seek new ways to make their models more robust to the multiple global changes occurring today.     

The formulations of site-scale processes affect landscape-scale forest change predictions: a comparison between LANDIS PRO and LANDIS-II forest landscape models

Context

Forest landscape models (FLMs) are important tools for simulating forest changes over broad spatial and temporal scales. The ability of FLMs to accurately predict forest changes may be significantly influenced by the formulations of site-scale processes including seedling establishment, tree growth, competition, and mortality.

Objective

The objectives of this study were to investigate the effects of site-scale processes and interaction effects of site-scale processes and harvest on landscape-scale forest change predictions.

Methods

We compared the differences in species’ distribution (quantified by species’ percent area), total aboveground biomass, and species’ biomass derived from two FLMs: (1) a model that explicitly incorporates stand density and size for each species age cohort (LANDIS PRO), and (2) a model that explicitly tracks biomass for each species age cohort (LANDIS-II with biomass succession extension), which are variants from the LANDIS FLM family with different formulations of site-scale processes.

Results

For early successional species, the differences in simulated distribution and biomass were small (mostly less than 5 %). For mid- to late-successional species, the differences in simulated distribution and biomass were relatively large (10–30 %). The differences in species’ biomass predictions were generally larger than those for species’ distribution predictions. Harvest mediated the differences on landscape-scale predictions.

Conclusions

The effects of site-scale processes on landscape-scale forest change predictions are dependent on species’ ecological traits such as shade tolerance, seed dispersal, and growth rates.

     

Coupling ecosystem and landscape models to study the effects of plot number and location on prediction of forest landscape change

A fundamental but unsolved dilemma is that observation and prediction scales are often mismatched. Reconciling this mismatch largely depends on how to design samples on a heterogeneous landscape. In this study, we used a coupled modeling approach to investigate the effects of plot number and location on predicting tree species distribution at the landscape scale. We used an ecosystem process model (LINKAGES) to generate tree species response to the environment (a land type) at the plot scale. To explore realistic parameterization scenarios we used results from LINKAGES simulations on species establishment probabilities under the current and warming climate. This allowed us to design a series of plot number and location scenarios at the landscape scale. Species establishment probabilities for different land types were then used as input for the forest landscape model (LANDIS) that simulated tree species distribution at the landscape scale. To investigate the effects of plot number and location on forest landscape predictions, LANDIS considered effects of climate warming only for the land types in which experimental plots were placed; otherwise inputs for the current climate were used. We then statistically examined the relationships of response variables (species percent area) among these scenarios and the reference scenario in which plots were placed on all land types of the study area. Our results showed that for species highly or moderately sensitive to environmental heterogeneity, increasing plot numbers to cover as many land types as possible is the strategy to accurately predict species distribution at the landscape scale. In contrast, for species insensitive to environmental heterogeneity, plot location was more important than plot number. In this case, placing plots in land types with large area of species distribution is warranted. For some moderately sensitive species that experienced intense disturbance, results were different in different simulation periods. Results from this study may provide insights into sample design for forest landscape predictions.     

Old-growth forest landscape transitions from pre-European settlement to present

We conducted a multi-temporal spatial analysis of forest cover for a 9600 ha landscape in northern Wisconsin, U.S.A., using data from pre-European settlement (1860s), post-settlement (1931), and current (1989) periods. Using GIS we have shown forest landscape changes and trajectories that have been generally described in aggregate for the norther Great Lake States region. We created the pre-European settlement map from the witness tree data of the original federal General Land Office survey notes. The 1931 cover was produced from the Wisconsin Land Economic Inventory, and the 1989 cover map was based on color infrared photography. We used GIS to analyze 1) land area occupied by different forest types at different dates, 2) temporal transitions between dates and their driving proceses, and 3) successional trajectories with landforms and spatial associations of forest types. Over the 120 year period, forest cover has changed from a landscape dominated by old-growth hemlock (Tsuga canadensis) and hardwood forests (Acer saccharum, Betula alleghaniensis) to largely second-growth hardwoods and conifers. The former dominant hemlock is largely eliminated from the landscape. From 1860 to 1931, large-scale disturbances associated with logging were the dominant processes on the landscape. Early successional forest types covered much of the landscape by the 1930s. From 1931 to 1989, succession was the dominant process driving forest transitions as forest types succeeded to a diverse group of upland hardwood and conifer forest types. If successional trajectories continue, a more homogeneous landscape may develop comprised of both a northern hardwood type dominated by sugar maple, and a boreal conifer/hardwood forest.     

Spatial dynamics of coastal forest bird assemblages: the influence of landscape context,forest type,and structural connectivity

Context

Complex structural connectivity patterns can influence the distribution of animals in coastal landscapes, particularly those with relatively large home ranges, such as birds. To understand the nuanced nature of coastal forest avifauna, where there may be considerable overlap in assemblages of adjacent forest types, the concerted influence of regional landscape context and vegetative structural connectivity at multiple spatial scales warrants investigation.

Objectives

This study determined whether species compositions of coastal forest bird assemblages differ with regional landscape context or with forest type, and if this is influenced by structural connectivity patterns measured at multiple spatial scales.

Methods

Three replicate bird surveys were conducted in four coastal forest types at ten survey locations across two regional landscape contexts in northeast Australia. Structural connectivity patterns of 11 vegetation types were quantified at 3, 6, and 12 km spatial scales surrounding each survey location, and differences in bird species composition were evaluated using multivariate ordination analysis.

Results

Bird assemblages differed between regional landscape contexts and most coastal forest types, although Melaleuca woodland bird assemblages were similar to those of eucalypt woodlands and rainforests. Structural connectivity was primarily correlated with differences in bird species composition between regional landscape contexts, and correlation depended on vegetation type and spatial scale.

Conclusions

Spatial scale, landscape context, and structural connectivity have a combined influence on bird species composition. This suggests that effective management of coastal landscapes requires a holistic strategy that considers the size, shape, and configuration of all vegetative components at multiple spatial scales.

     

Elasticity and loop analyses: tools for understanding forest landscape response to climatic change in spatial dynamic models

Spatially explicit dynamic forest landscape models have been important tools to study large-scale forest landscape response under global climatic change. However, the quantification of relative importance of different transition pathways among different forest types to forest landscape dynamics stands as a significant challenge. In this study, we propose a novel approach of elasticity and loop analyses to identify important transition pathways contributing to forest landscape dynamics. The elasticity analysis calculates the elasticity to measure the importance of one-directional transitions (transition from one forest type directly to another forest type); while the loop analysis is employed to measure the importance of different circular transition pathways (transition from one forest type through other forest types back to itself). We apply the proposed approach to a spatially explicit dynamic model, LANDIS-II, in a study of forest landscape response to climatic change in the Boundary Waters Canoe Area (BWCA) incorporating the uncertainties in climatic change predictions. Our results not only corroborate the findings of the previous studies on the most likely future forest compositions under simulated climatic variability, but also, through the novel application of the elasticity and loop analyses concepts, provide a quantitative assessment of the specific mechanisms leading to particular forest compositions, some of which might remain undetected with conventional model evaluation methods. By quantifying the importance of specific processes (transitions among forest types) to forest composition dynamics, the proposed approach can be a valuable tool for a more quantitative understanding of the relationship between processes and landscape composition/patterns.     

Soil nitrogen and carbon dynamics in a fragmented landscape experiencing forest succession

Forest fragmentation is an increasingly common feature across the globe, but few studies examine its influence on biogeochemical fluxes. We assessed the influence of differences in successional trajectory and stem density with forest patch size on biomass quantity and quality and N transformations in the soil at an experimentally fragmented landscape in Kansas, USA. We measured N-related fluxes in the laboratory, not the field, to separate effects of microclimate and fragment edges from the effects of inherent biomass differences with patch size. We measured net N mineralization and N₂O fluxes in soil incubations, gross rates of ammonification and nitrification, and microbial biomass in soils. We also measured root and litterfall biomass, C:N ratios, and δ¹³C and δ¹⁵N signatures; litterfall [cellulose] and [lignin]; and [C], [N], and δ¹³C and δ¹⁵N of soil organic matter. Rates of net N mineralization and N₂O fluxes were greater (by 113% and 156%, respectively) in small patches than in large, as were gross rates of nitrification. These differences were associated with greater quantities of root biomass in small patch soil profiles (664.2 ± 233.3 vs 192.4 ± 66.2 g m⁻² for the top 15 cm). These roots had greater N concentration than in large patches, likely generating greater root derived organic N pools in small patches. These data suggest greater rates of N cycling in small forested patches compared to large patches, and that gaseous N loss from the ecosystem may be related to forest patch size. The study indicates that the differences in successional trajectory with forest patch size can impart significant influence on soil N transformations in fragmented, aggrading woodlands.     

Statistical models of landscape pattern metrics,with applications to regional scale dynamic forest simulations

Forest managers in Canada need to model landscape pattern or spatial configurationoverlarge (100,000 km²) regions. This presents a scalingproblem, as landscape configuration is measured at a high spatial resolution,but a low spatial resolution is indicated for regional simulation. We present astatistical solution to this scaling problem by showing how a wide range oflandscape pattern metrics can be modelled from low resolution data. Our studyarea comprises about 75,000 km² of boreal mixedwoodforest in northeast Alberta, Canada. Within this area we gridded a sample of 84digital forest cover maps, each about 9500 ha in size, to aresolution of 1 ha and used FRAGSTATS to compute a suite oflandscape pattern metrics for each map. We then used multivariate dimensionreduction techniques and canonical correlation analysis to model therelationship between landscape pattern metrics and simpler stand table metricsthat are easily obtained from non-spatial forest inventories. These analyseswere performed on four habitat types common in boreal mixedwood forests: youngdeciduous, old deciduous, white spruce, and mixedwood types. Using only threelandscape variables obtained directly from stand attribute tables (totalhabitatarea, and the mean and standard deviation of habitat patch size), ourstatistical models explained more than 73% of the joint variation in fivelandscape pattern metrics (representing patch shape, forest interior habitat,and patch isolation). By PCA, these five indices captured much of the totalvariability in the rich set of landscape pattern metrics that FRAGSTATS cangenerate. The predictor variables and strengths of association were highlyconsistent across habitat classes. We illustrate the potential use of suchstatistical relationships by simulating the regional, cumulative effects ofwildfire and forest management on the spatial arrangement of forest patches,using non-spatial stand attribute tables.This revised version was published online in May 2005 with corrections to the Cover Date.     

Interactions of landscape disturbances and climate change dictate ecological pattern and process: spatial modeling of wildfire,insect, and disease dynamics under future climates

Context

Interactions among disturbances, climate, and vegetation influence landscape patterns and ecosystem processes. Climate changes, exotic invasions, beetle outbreaks, altered fire regimes, and human activities may interact to produce landscapes that appear and function beyond historical analogs.

Objectives

We used the mechanistic ecosystem-fire process model FireBGCv2 to model interactions of wildland fire, mountain pine beetle (Dendroctonus ponderosae), and white pine blister rust (Cronartium ribicola) under current and future climates, across three diverse study areas.

Methods

We assessed changes in tree basal area as a measure of landscape response over a 300-year simulation period for the Crown of the Continent in north-central Montana, East Fork of the Bitterroot River in western Montana, and Yellowstone Central Plateau in western Wyoming, USA.

Results

Interacting disturbances reduced overall basal area via increased tree mortality of host species. Wildfire decreased basal area more than beetles or rust, and disturbance interactions modeled under future climate significantly altered landscape basal area as compared with no-disturbance and current climate scenarios. Responses varied among landscapes depending on species composition, sensitivity to fire, and pathogen and beetle suitability and susceptibility.

Conclusions

Understanding disturbance interactions is critical for managing landscapes because forest responses to wildfires, pathogens, and beetle attacks may offset or exacerbate climate influences, with consequences for wildlife, carbon, and biodiversity.

     

Validating landtype associations using forest inventory and analysis data and neutral landscape models

One of the principal uses of land type associations (LTAs) is to provide information on ecological patterns and potentials useful for identifying alternatives and setting vegetation management objectives at landscape and watershed scales. Since LTA identification and delineation is an iterative process involving subjective decisions, it is imperative that LTAs accurately capture measurable distinctions in vegetation. In this study, we provided a framework for validating LTAs in Missouri’s. We chose a suite of variables from the Forest Inventory and Analysis database of the US Forest Service, an independent source, to capture a wide range of characteristics of the forest ecosystem. These variables included forest type, species composition, species diversity, species richness, site index, density of all trees 1 inch dbh and greater, and density of all trees 5 inch dbh and greater. First, the appropriateness of grouping LTAs into broader LTA types (identifications) was examined, and results suggest that species composition is more similar within LTA Type than among LTA Types. Second, a neutral model approach was used to evaluate the variables used in the study, and it was shown that forest type and the two density measures validated LTA adjacencies no more often than randomly delineated LTAs. Therefore, these three variables were removed from the analysis. Finally, comparisons of adjacent LTAs using statistical analysis of remaining variables resulted in the validation of 480 of the 623 compared adjacencies, showing a significant difference in at least one variable. Results of this study provide a quantitative measure of Missouri LTAs and reveal LTAs that have less distinction and need further refinements.     

Integrating models across temporal and spatial scales to simulate landscape patterns and dynamics in mountain pasture-woodlands

Context

Pasture-woodlands are semi-natural landscapes that result from the combined influences of climate, management, and intrinsic vegetation dynamics. These landscapes are sensitive to future changes in land use and climate, but our ability to predict the impact on ecosystem service provisioning is limited due to the disparate scales in time and space that govern their dynamics.

Objectives

To develop a process-based model to simulate pasture-woodland landscapes and the provisioning of ecosystem services (i.e., livestock forage, woody biomass and landscape heterogeneity).

Methods

We modified a dynamic forest landscape model to simulate pasture-woodland landscapes in Switzerland. This involved including an annual herbaceous layer, selective grazing from cattle, and interactions between grazing and tree recruitment. Results were evaluated within a particular pasture, and then the model was used to simulate regional vegetation patterns and livestock suitability for a ~198,000 ha landscape in the Jura Vaudois region.

Results

The proportion of vegetation cover types at the pasture level (i.e., open, semi-open and closed forests) was well represented, but the spatial distribution of trees was only broadly similar. The entire Jura Vaudois region was simulated to be highly suitable for livestock, with only a small proportion being unsuitable due to steep slopes and high tree cover. High and low elevation pastures were equally suitable for livestock, as lower forage production at higher elevations was compensated by reduced tree cover.

Conclusions

The modified model is valuable for assessing landscape to regional patterns in vegetation and livestock, and offers a platform to evaluate how climate and management impact ecosystem services.

     

An ecological classification of forest landscape simulation models: tools and strategies for understanding broad-scale forested ecosystems

Computer models are increasingly being used by forest ecologists and managers to simulate long-term forest landscape change. We review models of forest landscape change from an ecological rather than methodological perspective. We developed a classification based on the representation of three ecological criteria: spatial interactions, tree species community dynamics, and ecosystem processes. Spatial interactions are processes that spread across a landscape and depend upon spatial context and landscape configuration. Communities of tree species may change over time or can be defined a priori. Ecosystem process representation may range from no representation to a highly mechanistic, detailed representation. Our classification highlights the implicit assumptions of each model group and helps define the problem set for which each model group is most appropriate. We also provide a brief history of forest landscape simulation models, summarize the current trends in methods, and consider how forest landscape models may evolve and continue to contribute to forest ecology and management. Our classification and review can provide novice modelers with the ecological context for understanding or choosing an appropriate model for their specific hypotheses. In addition, our review clarifies the challenges and opportunities that confront practicing model users and model developers.     

Collaborative scenario modeling reveals potential advantages of blending strategies to achieve conservation goals in a working forest landscape

Context

Broad-scale land conservation and management often involve applying multiple strategies in a single landscape. However, the potential outcomes of such arrangements remain difficult to evaluate given the interactions of ecosystem dynamics, resource extraction, and natural disturbances. The costs and potential risks of implementing these strategies make robust evaluation critical.

Objectives

We used collaborative scenario modeling to compare the potential outcomes of alternative management strategies in the Two Hearted River watershed in Michigan’s Upper Peninsula to answer key questions: Which management strategies best achieve conservation goals of maintaining landscape spatial heterogeneity and conserving mature forests and wetlands? And how does an increase in wildfire and windthrow disturbances influence these outcomes?

Methods

Scenarios were modeled using the VDDT/TELSA state-and-transition modeling suite, and resulting land cover maps were analyzed using ArcGIS, FRAGSTATS, and R statistical software.

Results

Results indicate that blending conservation strategies, such as single-ownership forest reserves and working forest conservation easements in targeted areas of the landscape, may better achieve these goals than applying a single strategy across the same area. However, strategies that best achieve these conservation goals may increase the sensitivity of the landscape to changes in wildfire and windthrow disturbance regimes.

Conclusions

These results inform decision-making about which conservation strategy or combination of strategies to apply in specific locations on the landscape to achieve optimum conservation outcomes, how to best utilize scarce financial resources, and how to reduce the financial and ecological risks associated with the application of innovative strategies in an uncertain future.

     

Soil saturation effects on forest dynamics: scaling across a southern boreal/northern hardwood landscape

Patch modeling can be used to scale-up processes to portray landscape-level dynamics. Via direct extrapolation, a heterogeneous landscape is divided into its constituent patches; dynamics are simulated on each representative patch and are weighted and aggregated to formulate the higher level response. Further extrapolation may be attained by coarsening the resolution of or lumping environmental data (e.g., climatic, edaphic, hydrologic, topographic) used to delimit a patch.Forest patterns at the southern boreal/northern hardwood transition zone are often defined by soil heterogeneity, determined primarily by the extent and duration of soil saturation. To determine how landscape-level dynamics predicted from direct extrapolation compare when coarsening soil parameters, we simulated forest dynamics for soil series representing a range of drainage classes from east- central Maine. Responses were aggregated according to the distribution of soil associations comprising a 600 ha area based on local- (1:12,000), county- (1:120,000) and state- (1:250,000) scale soil maps. At the patch level, simulated aboveground biomass accumulated more slowly in poorer draining soils. Different soil series yielded different communities comprised of species with various tolerances for soil saturation. When aggregated, removal of waterlogging caused a 20–60% increase in biomass accumulation during the first 50 years of simulation. However, this early successional increase and the maximum level of biomass accumulation over a 200 year period varied by as much as 40% depending on the geospatial data. This marked discrepancy suggests caution when extrapolating with forest patch models by coarsening parameters and demonstrates how rules used to rescale environmental data need to be evaluated for consistency.     

Correction to: Coupling landscape graph modeling and biological data: a review

Landscape Ecology - In the original publication of the article, the sixth author name has been misspelt. The correct name is given in this Correction. The original article has been corrected.     

Long-term,landscape patterns of past fire events in a montane ponderosa pine forest of central Colorado

Parameters of fire regimes, including fire frequency, spatial extent of burned areas, fire severity, and season of fire occurrence, influence vegetation patterns over multiple scales. In this study, centuries-long patterns of fire events in a montane ponderosa pine – Douglas-fir forest landscape surrounding Cheesman Lake in central Colorado were reconstructed from fire-scarred trees and inferences from forest stand ages. We crossdated 153 fire-scarred trees from an approximately 4000 ha study area that recorded 77 total fire years from 1197 to the present. Spatial extent of burned areas during fire years varied from the scale of single trees or small clusters of trees to fires that burned across the entire landscape. Intervals between fire years varied from 1 to 29 years across the entire landscape to 3 to 58 years in one stand, to over 100 years in other stands. Large portions of the landscape did not record any fire for a 128 year-long period from 1723 to 1851. Fire severity varied from low-intensity surface fires to large-scale, stand-destroying fires, especially during the 1851 fire year but also possibly during other years. Fires occurred throughout tree growing seasons and both before and after growing seasons. These results suggest that the fire regime has varied considerably across the study area during the past several centuries. Since fires influence plant establishment and mortality on the landscape, these results further suggest that vegetation patterns changed at multiple scales during this period. The fire history from Cheesman Lake documents a greater range in fire behavior in ponderosa pine forests than generally has been found in previous studies.     

Cultural landscapes of the future: using agent-based modeling to discuss and develop the use and management of the cultural landscape of South West Devon

Context

Hedgerows are typical landscape features of high environmental and cultural value that often have been sacrificed for agricultural intensification and scale enlargement.

Objectives

We studied the dynamics of hedgerow quality over time in a case study area renowned for its hedgerow landscapes: South West Devon (UK) answering the following research questions: (1) how does the imperative of scale enlargement affect hedgerow quality? and (2) to what extent can cultural landscape degradation be countered by targeted policies?

Methods

We applied an agent-based modeling approach, parameterized with a site specific survey, to explore and discuss outcomes of future landscape change with stakeholders and co-designed preferred scenarios of landscape change during a workshop.

Results

Outcomes suggested that in the case-study area, scale enlargement has a negative effect on hedgerow quality when agri-environment scheme subsidies (AES) are low. In contrast, if the level of AES enrollment is high, scale enlargement can have a positive effect on hedgerow quality, as large holders are more likely to enroll for AES. Stakeholders acknowledged the need for agricultural intensification, but at the same time valued biodiversity and environmental value of the landscape in South West Devon.

Conclusion

Current AES are able to retain a decent hedgerow quality. With lower AES, scale enlargement can have an invigorative effect on hedgerow quality as land managers of larger farms will be less likely to join AES As an addition to AES, harvesting wood fuel from coppiced hedgerows appears a promising way to incentivize rejuvenating hedgerow management without governmental subsidies.

     

Contrasting land use legacy effects on forest landscape dynamics in the Italian Alps and the Apennines

Landscape Ecology - Land use legacies of human activities and recent post-abandonment forest expansion have extensively modified numerous forest landscapes throughout the European mountain ranges....