Should heterogeneity be the basis for conservation? Grassland bird response to fire and grazing.

In tallgrass prairie, disturbances such as grazing and fire can generate patchiness across the landscape, contributing to a shifting mosaic that presumably enhances biodiversity. Grassland birds evolved within the context of this shifting mosaic, with some species restricted to one or two patch types created under spatially and temporally distinct disturbance regimes. Thus, management-driven reductions in heterogeneity may be partly responsible for declines in numbers of grassland birds. We experimentally altered spatial heterogeneity of vegetation structure within a tallgrass prairie by varying the spatial and temporal extent of fire and by allowing grazing animals to move freely among burned and unburned patches (patch treatment). We contrasted this disturbance regime with traditional agricultural management of the region that promotes homogeneity (traditional treatment). We monitored grassland bird abundance during the breeding seasons of 2001-2003 to determine the influence of altered spatial heterogeneity on the grassland bird community. Focal disturbances of patch burning and grazing that shifted through the landscape over several years resulted in a more heterogeneous pattern of vegetation than uniform application of fire and grazing. Greater spatial heterogeneity in vegetation provided greater variability in the grassland bird community. Some bird species occurred in greatest abundance within focally disturbed patches, while others occurred in relatively undisturbed patches in our patch treatment. Henslow's Sparrow, a declining species, occurred only within the patch treatment. Upland Sandpiper and some other species were more abundant on recently disturbed patches within the same treatment. The patch burn treatment created the entire gradient of vegetation structure required to maintain a suite of grassland bird species that differ in habitat preferences. Our study demonstrated that increasing spatial and temporal heterogeneity of disturbance in grasslands increases variability in vegetation structure that results in greater variability at higher trophic levels. Thus, management that creates a shifting mosaic using spatially and temporally discrete disturbances in grasslands can be a useful tool in conservation. In the case of North American tallgrass prairie, discrete fires that capitalize on preferential grazing behavior of large ungulates promote a shifting mosaic of habitat types that maintain biodiversity and agricultural productivity.     

A spatially explicit model to predict future landscape composition of aspen woodlands under various management scenarios

Quaking aspen (Populus tremuloides) is declining across the western United States. Aspen habitats are among the most diverse plant communities in this region and loss of these habitats can result in shifts in biodiversity, productivity, and hydrology across a range of spatial scales. Western aspen occurs on the majority of sites seral to conifer species, and long-term maintenance of these aspen woodlands requires periodic fire. Over the past century, fire intervals, extents, and intensities have been insufficient to regenerate aspen stands at historic rates; however the effects of various fire regimes and management scenarios on aspen vegetation dynamics at broad spatial and temporal scales are unexplored. Here we use field data, remotely sensed data, and fire atlas information to develop a spatially explicit landscape simulation model to assess the effects of current and historic wildfire regimes and prescribed burning programs on landscape vegetation composition across two mountain ranges in the Owyhee Plateau, Idaho. Model outputs depict the future structural makeup and species composition of the landscape at selected time steps under simulated management scenarios. We found that under current fire regimes and in the absence of management activities, loss of seral aspen stands will continue to occur over the next two centuries. However, a return to historic fire regimes (burning 12–14% of the modeled landscape per decade) would maintain the majority of aspen stands in early and mid seral woodland stages and minimizes the loss of aspen. A fire rotation of 70–80 years was estimated for the historic fire regime while the current fire regime resulted in a fire rotation of 340–450 years, underscoring the fact that fire is currently lacking in the system. Implementation of prescribed burning programs, treating aspen and young conifer woodlands according to historic fire occurrence probabilities, are predicted to prevent conifer dominance and loss of aspen stands.     

A hierarchical,multi-scale,management-responsive model of Mediterranean vegetation dynamics

Ecosystems of the Mediterranean basin are characterized by a heterogeneous and dynamic landscape mosaic of vegetation formations. This landscape has been shaped over millenia by disturbances associated with agropastoral land use: clearing, grazing and burning, and by regeneration processes of the natural woody vegetation. The ability to predict the effects of management decisions on the structure and composition of the vegetation is essential for present and future land management. To improve this ability we developed a hierarchical multi-scale, management-responsive model of vegetation dynamics.     

The effects of shifting vegetation mosaics on habitat suitability for coastal dune fauna—a case study on sand lizards (Lacerta agilis)

Encroachment of tall grasses and shrubs in coastal dunes has resulted in loss of vegetation heterogeneity. This is expected to have negative effects on animal diversity. To counteract encroachment and develop structural heterogeneity grazing is a widely used management practice. Here, we aim to functionally interpret changes in vegetation composition and configuration following grazing management on habitat suitability for sand lizards. Aerial photographs taken over a period of 16 years were used to quantify changes in vegetation composition. A GIS-based method was developed to calculate habitat suitability for sand lizards in a spatially explicit manner, encompassing differences in vegetation structure and patch size. From 1987 to 2003 dune vegetation shifted from small patches of moss and sand to larger patches covered by shrubs and grasses. Grazing management did not have any significant effect on the overall level of heterogeneity, measured as habitat suitability for sand lizards. However, on a more local scale highly suitable patches in 1987 were deteriorating whereas unsuitable patches became more suitable in 2003. This inversion results from a broad shift with shrubs being a limiting habitat element in 1987 to sandy patches being the limiting element in 2003. Future changes are believed to negatively impact sand lizards. The habitat suitability model has proven to be a useful tool to functionally interpret changes in coastal dune vegetation heterogeneity from an animal’s perspective. Further research should aim to include multiple species operating on different scale levels to fully capture the natural landscape dynamics.     

Land‐use history as a guide for forest conservation and management

Conservation efforts to protect forested landscapes are challenged by climate projections that suggest substantial restructuring of vegetation and disturbance regimes in the future. In this regard, paleoecological records that describe ecosystem responses to past variations in climate, fire, and human activity offer critical information for assessing present landscape conditions and future landscape vulnerability. We illustrate this point drawing on 8 sites in the northwestern United States, New Zealand, Patagonia, and central and southern Europe that have undergone different levels of climate and land‐use change. These sites fall along a gradient of landscape conditions that range from nearly pristine (i.e., vegetation and disturbance shaped primarily by past climate and biophysical constraints) to highly altered (i.e., landscapes that have been intensely modified by past human activity). Position on this gradient has implications for understanding the role of natural and anthropogenic disturbance in shaping ecosystem dynamics and assessments of present biodiversity, including recognizing missing or overrepresented species. Dramatic vegetation reorganization occurred at all study sites as a result of postglacial climate variations. In nearly pristine landscapes, such as those in Yellowstone National Park, climate has remained the primary driver of ecosystem change up to the present day. In Europe, natural vegetation–climate–fire linkages were broken 6000–8000 years ago with the onset of Neolithic farming, and in New Zealand, natural linkages were first lost about 700 years ago with arrival of the Maori people. In the U.S. Northwest and Patagonia, the greatest landscape alteration occurred in the last 150 years with Euro‐American settlement. Paleoecology is sometimes the best and only tool for evaluating the degree of landscape alteration and the extent to which landscapes retain natural components. Information on landscape‐level history thus helps assess current ecological change, clarify management objectives, and define conservation strategies that seek to protect both natural and cultural elements.     

Influence of environment, disturbance, and ownership on forest vegetation of coastal Oregon.

Information about how vegetation composition and structure vary quantitatively and spatially with physical environment, disturbance history, and land ownership is fundamental to regional conservation planning. However, current knowledge about patterns of vegetation variability across large regions that is spatially explicit (i.e., mapped) tends to be general and qualitative. We used spatial predictions from gradient models to examine the influence of environment, disturbance, and ownership on patterns of forest vegetation biodiversity across a large forested region, the 3-million-ha Oregon Coast Range (USA). Gradients in tree species composition were strongly associated with environment, especially climate, and insensitive to disturbance, probably because many dominant tree species are long-lived and persist throughout forest succession. In contrast, forest structure was strongly correlated with disturbance and only weakly with environmental gradients. Although forest structure differed among ownerships, differences were blurred by the presence of legacy trees that originated prior to current forest management regimes. Our multi-ownership perspective revealed biodiversity concerns and benefits not readily visible in single-ownership analyses, and all ownerships contributed to regional biodiversity values. Federal lands provided most of the late-successional and old-growth forest. State lands contained a range of forest ages and structures, including diverse young forest, abundant legacy dead wood, and much of the high-elevation true fir forest. Nonindustrial private lands provided diverse young forest and the greatest abundance of hardwood trees, including almost all of the foothill oak woodlands. Forest industry lands encompassed much early-successional forest, most of the mixed hardwood-conifer forest, and large amounts of legacy down wood. The detailed tree- and species-level data in the maps revealed regional trends that would be masked in traditional coarse-filter assessment. Although abundant, most early-successional forests originated after timber harvest and lacked legacy live and dead trees important as habitat and for other ecological functions. Many large-conifer forests that might be classified as old growth using a generalized forest cover map lacked structural features of old growth such as multilayered canopies or dead wood. Our findings suggest that regional conservation planning include all ownerships and land allocations, as well as fine-scale elements of vegetation composition and structure.     

Distinguishing stressors acting on land bird communities in an urbanizing environment

Urbanization has profound influences on ecological communities, but our understanding of causal mechanisms is limited by a lack of attention to its component stressors. Published research suggests that at landscape scales, habitat loss and fragmentation are the major drivers of community change, whereas at local scales, human activity and vegetation management are the primary stressors. Little research has focused on whether urbanization stressors may supplant natural factors as dominant forces structuring communities. We used model selection to determine the relative importance of urban development, human activity, local and landscape vegetation, topography, and geographical location in explaining land bird species richness, abundance, and dominance. We analyzed the entire community and groups of species based on ecological characteristics, using data collected in remnant forests along a gradient of urban development in the Lake Tahoe basin, California and Nevada, USA. Urbanization stressors were consistently among the principal forces structuring the land bird community. Strikingly, disturbance from human activity was the most important factor for richness in many cases, surpassing even habitat loss from development. Landscape-scale factors were consistently more important than local-scale factors for abundance. In demonstrating considerable changes in land bird community structure, our results suggest that ecosystem function in urban areas may be severely compromised. Such changes compel local- and landscape-scale management, focused research, and long-term monitoring to retain biodiversity in urban areas to the extent possible.     

Accounting for system dynamics in reserve design.

Systematic conservation plans have only recently considered the dynamic nature of ecosystems. Methods have been developed to incorporate climate change, population dynamics, and uncertainty in reserve design, but few studies have examined how to account for natural disturbance. Considering natural disturbance in reserve design may be especially important for the world's remaining intact areas, which still experience active natural disturbance regimes. We developed a spatially explicit, dynamic simulation model, CONSERV, which simulates patch dynamics and fire, and used it to evaluate the efficacy of hypothetical reserve networks in northern Canada. We designed six networks based on conventional reserve design methods, with different conservation targets for woodland caribou habitat, high-quality wetlands, vegetation, water bodies, and relative connectedness. We input the six reserve networks into CONSERV and tracked the ability of each to maintain initial conservation targets through time under an active natural disturbance regime. None of the reserve networks maintained all initial targets, and some over-represented certain features, suggesting that both effectiveness and efficiency of reserve design could be improved through use of spatially explicit dynamic simulation during the planning process. Spatial simulation models of landscape dynamics are commonly used in natural resource management, but we provide the first illustration of their potential use for reserve design. Spatial simulation models could be used iteratively to evaluate competing reserve designs and select targets that have a higher likelihood of being maintained through time. Such models could be combined with dynamic planning techniques to develop a general theory for reserve design in an uncertain world.     

Disturbance and landscape dynamics in a changing world

Disturbance regimes are changing rapidly, and the consequences of such changes for ecosystems and linked social-ecological systems will be profound. This paper synthesizes current understanding of disturbance with an emphasis on fundamental contributions to contemporary landscape and ecosystem ecology, then identifies future research priorities. Studies of disturbance led to insights about heterogeneity, scale, and thresholds in space and time and catalyzed new paradigms in ecology. Because they create vegetation patterns, disturbances also establish spatial patterns of many ecosystem processes on the landscape. Drivers of global change will produce new spatial patterns, altered disturbance regimes, novel trajectories of change, and surprises. Future disturbances will continue to provide valuable opportunities for studying pattern-process interactions. Changing disturbance regimes will produce acute changes in ecosystems and ecosystem services over the short (years to decades) and long-term (centuries and beyond). Future research should address questions related to (1) disturbances as catalysts of rapid ecological change, (2) interactions among disturbances, (3) relationships between disturbance and society, especially the intersection of land use and disturbance, and (4) feedbacks from disturbance to other global drivers. Ecologists should make a renewed and concerted effort to understand and anticipate the causes and consequences of changing disturbance regimes.     

Conservation planning for successional landscapes

The systematic conservation planning literature invariably assumes that the biodiversity features being preserved in sites do not change through time. We develop a conservation planning framework for ecosystems where disturbance events and succession drive vegetation dynamics. The framework incorporates three key attributes of disturbance theory: heterogeneity in disturbance rates, spatial correlation between disturbance events and different impacts of disturbance. In our conservation problem we wish to maximise the chance that we represent a certain number of successional types given a cap on the number of sites we can conserve. Correlation between disturbance events dramatically complicates the problem of choosing the optimal suite of sites. However, in our problem we discover that spatial correlation in disturbances affects the optimal reserve network very little. The reason is twofold: (i) through our probabilistic framework we focus on the long-term effectiveness of reserve networks and (ii) in the dynamics considered in our model the state of a site is not only affected by the most recent (correlated) disturbance event but also by the site's long-term stochastic history which blurs the impact of spatial correlation. If successional states are the conservation target rather than individual species then, conserving a site can only contribute to meeting one target. However, given that correlation of disturbance events may be ignored, we show that if the number of candidate reserves is sufficiently large the statistical dependence of different conservation targets may be ignored, too. We conclude that the computational complexity of reserve selection methods for dynamic ecosystems can be much simpler than they first appear.     

Conserving and managing the subnivium

In regions where snowfall historically has been a defining seasonal characteristic of the landscape, warming winters have reduced the depth, duration, and extent of snowpack. However, most management and conservation has focused on how aboveground wildlife will be affected by altered snow conditions, even though the majority of species that persist through the winter do so under the snowpack in a thermally stable refugium: the subnivium. Shortened winters, forest management practices, and winter recreation can alter subnivium conditions by increasing snow compaction and compromising thermal stability at the soil–snow interface. To help slow the loss of the subnivium in the face of rapidly changing winter conditions, we suggest managers adopt regional conservation plans for identifying threatened snow‐covered environments; measure and predict the effects land cover and habitat management has on local subnivium conditions; and control the timing and distribution of activities that disturb and compact snow cover (e.g., silvicultural practices, snow recreation, and road and trail maintenance). As a case study, we developed a spatially explicit model of subnivium presence in a working landscape of the Chequamegon National Forest, Wisconsin. We identified landscapes where winter recreation and management practices could threaten potentially important areas for subnivium persistence. Similar modeling approaches could inform management decisions related to subnivium conservation. Current climate projections predict that snow seasons will change rapidly in many regions, and as result, we advocate for the immediate recognition, conservation, and management of the subnivium and its dependent species.     

Pyric Herbivory: Rewilding Landscapes through the Recoupling of Fire and Grazing

Abstract:  Our understanding of fire and grazing is largely based on small-scale experimental studies in which treatments are uniformly applied to experimental units that are considered homogenous. Any discussion of an interaction between fire and grazing is usually based on a statistical approach that ignores the spatial and temporal interactions on complex landscapes. We propose a new focus on the ecological interaction of fire and grazing in which each disturbance is spatially and temporally dependent on the other and results in a landscape where disturbance is best described as a shifting mosaic (a landscape with patches that vary with time since disturbance) that is critical to ecological structure and function of many ecosystems. We call this spatiotemporal interaction pyric herbivory (literal interpretation means grazing driven by fire). Pyric herbivory is the spatial and temporal interaction of fire and grazing, where positive and negative feedbacks promote a shifting pattern of disturbance across the landscape. We present data we collected from the Tallgrass Prairie Preserve in the southern Great Plains of North America that demonstrates that the interaction between free-roaming bison ( Bison bison ) and random fires promotes heterogeneity and provides the foundation for biological diversity and ecosystem function of North American and African grasslands. This study is different from other studies of fire and grazing because the fires we examined were random and grazing animals were free to roam and select from burned and unburned patches. For ecosystems across the globe with a long history of fire and grazing, pyric herbivory with any grazing herbivore is likely more effective at restoring evolutionary disturbance patterns than a focus on restoring any large vertebrate while ignoring the interaction with fire and other disturbances .     

Increasing heterogeneity of sensitive assemblages as a consequence of human impact in submarine caves

Submarine caves are considered as a top priority for conservation, but the effects of common pressures are poorly known for these habitats. Here, we examined the effect of recreational human visitation on a selection of submarine caves in a Mediterranean Marine Protected Area (40°35′40″N; 8°11′39″E) where diving activities are regulated. Sampling was conducted in visited and not visited caves to assess whether diving activities have a significant effect on cave habitats, what are the components of biodiversity most affected by this disturbance, and its potential effects on spatial heterogeneity of benthic assemblages. Results clearly showed that human visitation could significantly affect spatial patterns of benthic assemblages. Organisms with erect growth forms were significantly more abundant and homogeneously distributed where diving activities are forbidden. An increase in the small-scale heterogeneity of assemblages and a decrease in their three-dimensional structure could be the ultimate consequences of human visitation. The interaction between specific stressors and the patterns of distribution of species and assemblages can drive their spatial heterogeneity also in unique habitats like marine caves, representing an early warning for the development of appropriate management measures.     

Simulating the effects of frequent fire on southern California coastal shrublands.

Fire disturbance is a primary agent of change in the mediterranean-climate chaparral shrublands of southern California, USA. However, fire frequency has been steadily increasing in coastal regions due to ignitions at the growing wildland-urban interface. Although chaparral is resilient to a range of fire frequencies, successively short intervals between fires can threaten the persistence of some species, and the effects may differ according to plant functional type. California shrublands support high levels of biological diversity, including many endangered and endemic species. Therefore, it is important to understand the long-term effects of altered fire regimes on these communities. A spatially explicit simulation model of landscape disturbance and succession (LANDIS) was used to predict the effects of frequent fire on the distribution of dominant plant functional types in a study area administered by the National Park Service. Shrubs dependent on fire-cued seed germination were most sensitive to frequent fire and lost substantial cover to other functional types, including drought-deciduous subshrubs that typify coastal sage scrub and nonnative annual grasses. Shrubs that resprout were favored by higher fire frequencies and gained in extent under these treatments. Due to this potential for vegetation change, caution is advised against the widespread use of prescribed fire in the region.     

Patch-Occupancy Modeling as a Method for Monitoring Changes in Forest Floristics: a Case Study in Southeastern Australia

Abstract:  The ability to monitor changes in biodiversity is fundamental to demonstrating sustainable management practices of natural resources. Disturbance studies generally focus on responses at the plot scale, whereas landscape-scale responses are directly relevant to the development of sustainable forest management. Modeling changes in occupancy is one way to monitor landscape-scale responses. We used understory vegetation data collected over 16 years from a long-term study site in southeastern Australia. The site was subject to timber harvesting and frequent prescribed burning. We used occupancy models to examine the impacts of these disturbances on the distribution of 50 species of plants during the study. Timber harvesting influenced the distribution of 9 species, but these effects of harvesting were generally lost within 14 years. Repeated prescribed fire affected 22 species, but the heterogeneity of the burns reduced the predicted negative effects. Twenty-two species decreased over time independent of treatment, and only 5 species increased over time. These changes probably represent a natural response to a wildfire that occurred in 1973, 13 years before the study began. Occupancy modeling is a useful and flexible technique for analyzing monitoring data and it may also be suitable for inclusion within an adaptive-management framework for forest management.     

Interactive effects of natural and human disturbances on vegetation dynamics across landscapes.

Accurate measures of human effects on landscape processes require consideration of both the direct impacts from human activities and the indirect consequences of the interactions between humans and the landscape. This is particularly evident in systems experiencing regular natural disturbances such as in the mountainous areas of southwestern China, where the remaining population of giant pandas (Ailuropoda melanoleuca) is supported. Here the spatiotemporal patterns of human impacts, forests, and bamboo episodic die-offs combine to determine the distribution of panda habitat. To study the complex interactions of humans and landscapes, we developed an integrated spatiotemporally explicit model of household activities, natural vegetation dynamics, and their impacts on panda habitat. Using this model we examined the direct consequences of local fuelwood collection and household creation on areas of critical giant panda habitat and the indirect impacts when coupled with vegetation dynamics. Through simulations, we found that over the next 30 years household impacts would result in the loss of up to 30% of the habitat relied on by pandas during past bamboo die-offs. The accumulation and spatial distribution of household impacts would also have a considerable indirect influence on the spatial distribution of understory bamboo. While human impacts influence both bamboo die-off and regeneration, over 19% of pre-existing low-elevation bamboo habitat may be lost following an episodic die-off depending on the severity of the impacts and timing of the die-offs. Our study showed not only the importance of the spatial distribution of direct household impacts on habitat, but also the far-reaching effects of the indirect interactions between humans and the landscapes they are modifying.     

Scaling-up spatially-explicit ecological models using graphics processors

How the properties of ecosystems relate to spatial scale is a prominent topic in current ecosystem research. Despite this, spatially explicit models typically include only a limited range of spatial scales, mostly because of computing limitations. Here, we describe the use of graphics processors to efficiently solve spatially explicit ecological models at large spatial scale using the CUDA language extension. We explain this technique by implementing three classical models of spatial self-organization in ecology: a spiral-wave forming predator-prey model, a model of pattern formation in arid vegetation, and a model of disturbance in mussel beds on rocky shores. Using these models, we show that the solutions of models on large spatial grids can be obtained on graphics processors with up to two orders of magnitude reduction in simulation time relative to normal pc processors. This allows for efficient simulation of very large spatial grids, which is crucial for, for instance, the study of the effect of spatial heterogeneity on the formation of self-organized spatial patterns, thereby facilitating the comparison between theoretical results and empirical data. Finally, we show that large-scale spatial simulations are preferable over repetitions at smaller spatial scales in identifying the presence of scaling relations in spatially self-organized ecosystems. Hence, the study of scaling laws in ecology may benefit significantly from implementation of ecological models on graphics processors.     

Simulating dynamic and mixed-severity fire regimes: A process-based fire extension for LANDIS-II

Fire regimes result from reciprocal interactions between vegetation and fire that may be further affected by other disturbances, including climate, landform, and terrain. In this paper, we describe fire and fuel extensions for the forest landscape simulation model, LANDIS-II, that allow dynamic interactions among fire, vegetation, climate, and landscape structure, and incorporate realistic fire characteristics (shapes, distributions, and effects) that can vary within and between fire events. We demonstrate the capabilities of the new extensions using two case study examples with very different ecosystem characteristics: a boreal forest system from central Labrador, Canada, and a mixed conifer system from the Sierra Nevada Mountains (California, USA). In Labrador, comparison between the more complex dynamic fire extension and a classic fire simulator based on a simple fire size distribution showed little difference in terms of mean fire rotation and potential severity, but cumulative burn patterns created by the dynamic fire extension were more heterogeneous due to feedback between fuel types and fire behavior. Simulations in the Sierra Nevada indicated that burn patterns were responsive to topographic features, fuel types, and an extreme weather scenario, although the magnitude of responses depended on elevation. In both study areas, simulated fire size and resulting fire rotation intervals were moderately sensitive to parameters controlling the curvilinear response between fire spread and weather, as well as to the assumptions underlying the correlation between weather conditions and fire duration. Potential fire severity was more variable within the Sierra Nevada landscape and also was more sensitive to the correlation between weather conditions and fire duration. The fire modeling approach described here should be applicable to questions related to climate change and disturbance interactions, particularly within locations characterized by steep topography, where temporally or spatially dynamic vegetation significantly influences spread rates, where fire severity is variable, and where multiple disturbance types of varying severities are common.     

SEIB–DGVM: A new Dynamic Global Vegetation Model using a spatially explicit individual-based approach

We report the development of a new spatially explicit individual-based Dynamic Global Vegetation Model (SEIB–DGVM), the first DGVM that can simulate the local interactions among individual trees within a spatially explicit virtual forest. In the model, a sample plot is placed at each grid box, and then the growth, competition, and decay of each individual tree within each plot is calculated by considering the environmental conditions for that tree as it relates to the trees that surround it. Based on these parameters only, the model simulated time lags between climate change and vegetation change. This time lags elongated when original biome was forest, because existing trees prevent newly establish trees from receiving enough sunlight and space to quickly replace the original vegetation. This time lags also elongated when horizontal heterogeneity of sunlight distribution was ignored, indicating the potential importance of horizontal heterogeneity for predicting transitional behavior of vegetation under changing climate. On a local scale, the model reproduced climate zone-specific patterns of succession, carbon dynamics, and water flux, although on a global scale, simulations were not always in agreement with observations. Because the SEIB–DGVM was formulated to the scale at which field biologists work, the measurements of relevant parameters and data comparisons are relatively straightforward, and the model should enable more robust modeling of terrestrial ecosystems.