Functional heterogencity in resources within landscapes and herbivore population dynamics

Large mammalian herbivores are notorious for their propensity towards population irruptions and crashes, yet many herbivore populations remain relatively stable. I explore how resource heterogeneity within landscapes dampens population instability, using a metaphysiological modelling approach condidering patch state distributions. Resource heterogeneity is functionally stabilizing through spreading consumption away from preferred resources before these critically depleted. Lower-quality resources act as a buffer against starvation during critical critical periods of the seasonal cycle. Enriching resource quality is destabilizing, even if patch diversity is maintained, because food quantity then becomes the limitation. The potential consequences of landscape fragmentation are explored using the Serengeti ecosystem, characterised by broadscale resource gradients, as a hypothetical example. Further insights provided by the model are illustrated with specific examples concerning the effects of patch scales and waterpoint distribution. A metaphysiological modelling approach enables the basic consequences of landscape heterogeneity to be distinguished from further effects that may arise from specific patch scales and configurations, without the distracting detail of spatially explicit models.     

Effects of increasing landscape heterogeneity on local plant species richness: how much is enough?

Contemporary landscape ecology continues to explore the causes and consequences of landscape heterogeneity across a range of scales, and demands for the scientific underpinnings of landscape planning and management still remains high. The spatial distribution of resources can be a key element in determining habitat quality, and that in turn is directly related to the level of heterogeneity in the system. In this sense, forest habitat mosaics may be more affected by lack of heterogeneity than by structural fragmentation. Nonetheless, increasing spatial heterogeneity at a given spatial scale can also decrease habitat patch size, with potential negative consequences for specialist species. Such dual effect may lead to hump-backed shape relationships between species diversity and heterogeneity, leading to three related assumptions: (i) at low levels of heterogeneity, an increase in heterogeneity favours local and regional species richness, (ii) there is an optimum heterogeneity level at which a maximum number of species is reached, (iii) further increase in spatial heterogeneity has a negative effect on local and regional species richness, due to increasing adverse effects of habitat fragmentation. In this study, we investigated the existence of a hump-shaped relationship between local plant species richness and increasing forest landscape heterogeneity on a complex mosaic in the French Alps. Forest landscape heterogeneity was quantified with five independent criteria. We found significant quadratic relationships between local forest species richness and two heterogeneity criteria indicators, showing a slight decrease of forest species richness at very high heterogeneity levels. Species richness–landscape heterogeneity relationships varied according to the heterogeneity metrics involved and the type of species richness considered. Our results support the assumption that intermediate levels of heterogeneity may support more species than very high levels of heterogeneity, although we were not able to conclude for a systematic negative effect of very high levels of heterogeneity on local plant species richness.     

Interaction of food resources and landscape structure in determining the probability of patch use by carnivores in fragmented landscapes

Studies on the distribution of mammalian carnivores in fragmented landscapes have focused mainly on structural aspects such as patch and landscape features; similarly, habitat connectivity is usually associated with landscape structure. The influence of food resources on carnivore patch use and the important effect on habitat connectivity have been overlooked. The aim of this study is to evaluate the relative importance of food resources on patch use patterns and to test if food availability can overcome structural constraints on patch use. We carried out a patch-use survey of two carnivores: the beech marten (Martes foina) and the badger (Meles meles) in a sample of 39 woodland patches in a fragmented landscape in central Italy. We used the logistic model to investigate the relative effects on carnivore distribution of patch, patch neighbourhood and landscape scale variables as well as the relative abundance of food resources. Our results show how carnivore movements in fragmented landscapes are determined not only by patch/landscape structure but also by the relative abundance of food resources. The important take-home message of our research is that, within certain structural limits (e.g. within certain limits of patch isolation), by modifying the relative amount of resources and their distribution, it is possible to increase suitability in smaller/relatively isolated patches. Conversely, however, there are certain thresholds above which an increase in resources will not achieve high probability of presence. Our findings have important and generalizable consequences for highly fragmented landscapes in areas where it may not be possible to increase patch sizes and/or reduce isolation so, for instance, forest regimes that will increase resource availability could be implemented. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Animal movements and population dynamics in heterogeneous landscapes

Organisms respond to environmental heterogeneity at different scales and in different ways. These differences are consequences of how the movement characteristics of animals—their movement rates, directionality, turning frequencies, and turning angles—interact with patch and boundary features in landscape mosaics. The interactions of movement patterns with landscape features in turn produce spatial patterns in individual space-use, population dynamics and dispersion, gene flow, and the redistribution of nutrients and other materials. We describe several theoretical approaches for modeling the diffusion, foraging behavior, and population dynamics of animals in heterogeneous landscapes, including: (1) scaling relationships derived from percolation theory and fractal geometry, (2) extensions of traditional patch-based metapopulation models, and (3) individual-based, spatially explicit models governed by local rules. We conclude by emphasizing the need to couple theoretical models with empirical studies and the usefulness of ‘microlandscape’ investigations.     

Scaling properties in landscape patterns: New Zealand experience

In this paper we present a case study of spatial structure in landscape patterns for the North and South Islands of New Zealand. The aim was to characterise quantitatively landscape heterogeneity and investigate its possible scaling properties. The study examines spatial heterogeneity, in particular patchiness, at a range of spatial scales, to help build understanding on the effects of landscape heterogeneity on water movement in particular, and landscape ecology in general.We used spatial information on various landscape properties (soils, hydrogeology, vegetation, topography) generated from the New Zealand Land Resource Inventory. To analyse this data set we applied various methods of fractal analyses following the hypothesis that patchiness in selected landscape properties demonstrates fractal scaling behaviour at two structural levels: (1) individual patches; and (2) mosaics (sets) of patches.Individual patches revealed scaling behaviour for both patch shape and boundary. We found self-affinity in patch shape with Hurst exponent H from 0.75 to 0.95. We also showed that patch boundaries in most cases were self-similar and in a few cases of large patches were self-affine. The degree of self-affinity was lower for finer patches. Similarly, when patch scale decreases the orientation of patches tends to be uniformly distributed, though patch orientation on average is clearly correlated with broad scale geological structures. These results reflect a tendency to isotropic behaviour of individual patches from broad to finer scales. Mosaics of patches also revealed fractal scaling in the total patch boundaries, patch centers of mass, and in patch area distribution. All these reflect a special organisation in patchiness represented in fractal patch clustering. General relationships which interconnect fractal scaling exponents were derived and tested. These relationships show how scaling properties of individual patches affect those for mosaics of patches and vice-versa. To explain similarity in scaling behaviour in patchiness of different types we suggest that the Self-Organised Criticality concept should be used. Also, potential applications of our results in landscape ecology are discussed, especially in relation to improved neutral landscape models.     

A comparison of quantitative methods for examining landscape pattern and scale

Ecologists have long recognized the importance of spatial and temporal patterns that characterize heterogeneity in landscapes. However, despite the realization that inferences about ecological phenomena are scale dependent, little attention has been paid to determining appropriate scales of measurement (e.g., plot or grain size) in studies of landscape dynamics or ecosystem change. This paper compares the results from three data sets using several quantitative methods available for characterizing landscape heterogeneity and/or for determining scale of measurement. Methods evaluated include tests of non-randomness, estimation of patch size, spectral analysis, fractals, variance ratio analysis, and correlation analysis. The results showed that no one method provides consistently good estimates of scale. Thus, sampling strategies for landscape studies should be derived from estimates of patch size and/or scale of pattern obtained from more than one of these methods.     

Landscape dynamics in crown fire ecosystems

Crown fires create broad-scale patterns in vegetation by producing a patch mosaic of stand age classes, but the spread and behavior of crown fires also may be constrained by spatial patterns in terrain and fuels across the landscape. In this review, we address the implications of landscape heterogeneity for crown fire behavior and the ecological effects of crown fires over large areas. We suggest that fine-scale mechanisms of fire spread can be extrapolated to make broad-scale predictions of landscape pattern by coupling the knowledge obtained from mechanistic and empirical fire behavior models with spatially-explicit probabilistic models of fire spread. Climatic conditions exert a dominant control over crown fire behavior and spread, but topographic and physiographic features in the landscape and the spatial arrangement and types of fuels have a strong influence on fire spread, especially when burning conditions (e.g., fuel moisture and wind) are not extreme. General trends in crown fire regimes and stand age class distributions can be observed across continental, latitudinal, and elevational gradients. Crown fires are more frequent in regions having more frequent and/or severe droughts, and younger stands tend to dominate these landscapes. Landscapes dominated by crown fires appear to be nonequilibrium systems. This nonequilibrium condition presents a significant challenge to land managers, particularly when the implications of potential changes in the global climate are considered. Potential changes in the global climate may alter not only the frequency of crown fires but also their severity. Crown fires rarely consume the entire forest, and the spatial heterogeneity of burn severity patterns creates a wide range of local effects and is likely to influence plant reestablishment as well as many other ecological processes. Increased knowledge of ecological processes at regional scales and the effects of landscape pattern on fire dynamics should provide insight into our understanding of the behavior and consequences of crown fires.     

Environmental factors at different spatial scales governing soil fauna community patterns in fragmented forests

Spatial and temporal changes in community structure of soil organisms may result from a myriad of processes operating at a hierarchy of spatial scales, from small-scale habitat conditions to species movements among patches and large-sale landscape features. To disentangle the relative importance of spatial and environmental factors at different scales (plot, patch and landscape), we analyzed changes in Collembola community structure along a gradient of forest fragmentation, testing predictions of the Hierarchical Patch Dynamics Paradigm (HPDP) in different European biogeographic regions (Boreal, Continental, Atlantic, Mediterranean, Alpine). Using variance partitioning methods, based on partial CCAs, we observed that the independent effect of environmental processes was significantly explaining Collembola community variance in all regions, while the relative effect of spatial variables was not significant, due to the observed high levels of landscape heterogeneity along the gradient. Environmental factors at the patch and plot scales were generally significant and explained the larger part of community changes. Landscape variables were not significant across all study sites. Yet, at the landscape level, an increase in forest habitat and proximity of forest patches were showed to have an indirect influence on local community changes, by influencing microhabitat heterogeneity at lower spatial scales in all studied regions. In line with HPDP, large-scale landscape features influenced spatio-temporal changes in soil fauna communities by constraining small-scale environmental processes. In turn, these provided mechanistic understanding for diversity patterns operating at the patch scale, via shifts in community weighted mean of Collembola life-forms occurring in local communities along the fragmentation gradient.     

Resource utilization scales and landscape pattern

The spatial patterning of resources constrains the movement of consumers on the landscape. Percolation theory predicts that an organism can move freely if its critical resource or habitat occupies 59.28% of the landscape. Sparse resources require an organism to operate on larger resource utilization scales. Multiple critical resources necessitate larger scales, while substitutable resources ease the scale requirements. Contagious spatial patterns require larger scales to permit movement between resource clusters. The study indicates a strong link between spatial pattern and ecological processes on a landscape.     

Multi-scaled habitat considerations for conserving urban biodiversity: native reptiles and small mammals in Brisbane,Australia

The rapid expansion of the world’s urban population is a major driver of contemporary landscape change and ecosystem modification. Urbanisation destroys, degrades and fragments native ecosystems, replacing them with a heterogeneous matrix of urban development, parks, roads, and isolated remnant fragments of varying size and quality. This presents a major challenge for biodiversity conservation within urban areas. To make spatially explicit decisions about urban biodiversity conservation actions, urban planners and managers need to be able to separate the relative influence of landscape composition and configuration from patch and local (site)-scale variables for a range of fauna species. We address this problem using a hierarchical landscape approach for native, terrestrial reptiles and small mammals living in a fragmented semi-urban landscape of Brisbane, Australia. Generalised linear modelling and hierarchical partitioning analysis were applied to quantify the relative influence of landscape composition and configuration, patch size and shape, and local habitat composition and structure on the species’ richness of mammal and reptile assemblages. Landscape structure (composition and configuration) and local-scale habitat structure variables were found to be most important for influencing reptile and mammal assemblages, although the relative importance of specific variables differed between reptile and mammal assemblages. These findings highlight the importance of considering landscape composition and configuration in addition to local habitat elements when planning and/or managing for the conservation of native, terrestrial fauna diversity in urban landscapes.     

Assessing the influence of resource covariates at multiple spatial scales: an application to forest-dwelling caribou faced with intensive human activity

Efforts in isolating the relative effects of resources and disturbances on animal-distribution patterns remain hindered by the difficulty of accounting for multiple scales of resource selection by animals with seasonally dynamic drivers. We developed multi-scale, seasonal models to explore how local resource selection by the threatened forest-dwelling woodland caribou (Rangifer tarandus caribou) was influenced by both broad-scale landscape context and local resource heterogeneity in the intensively managed region of Charlevoix, Québec, Canada, located on the southern border of the North American caribou range. We estimated resource selection functions using 23 GPS-collared caribou monitored from 2004 to 2006 and landscape data on vegetation classes, terrain conditions, and roads. We found evidence of thresholds in road “proximity” effects (up to 1.25 km), which underscores the importance of including landscape context variables in addition to locally measured variables, and of fitting seasonal-specific models given temporal variation in the magnitude of selection and optimal scale of measurement. Open lichen woodlands were an important cover type for caribou during winter and spring, whereas deciduous forests, wetlands, and even young disturbed stands became important during calving and summer. Caribou consistently avoided roads and rugged terrain conditions at both local and landscape levels. Landscape context fundamentally constrains the choices available to animals, and we showed that failing to consider landscape context, or arbitrarily choosing an inappropriate scale for measuring covariates, may provide biased inferences with respect to habitat selection patterns. Effective habitat management for rare or declining species should carefully consider the hierarchical nature of habitat selection.     

Multi-scale predictive habitat suitability modeling based on hierarchically delineated patches: an example for yellow-billed cuckoos nesting in riparian forests,California, USA

The discipline of landscape ecology recognizes the importance of measuring habitat suitability variables at spatial scales relevant to specific organisms. This paper uses a novel multi-scale hierarchical patch delineation method, PatchMorph, to measure landscape patch characteristics at two distinct spatial scales and statistically relate them to the presence of state-listed endangered yellow-billed cuckoos (Coccyzus americanus occidentalis) nesting in forest patches along the Sacramento River, California, USA. The landscape patch characteristics calculated were: patch thickness, area of cottonwood forest, area of riparian scrub, area of other mixed riparian forest, and total patch area. A third, regional spatial variable, delineating the north and south portions of study area was also analyzed for the effect of regional processes. Using field surveys, the landscape characteristics were related to patch occupancy by yellow-billed cuckoos. The area of cottonwood forest measured at the finest spatial scale of patches was found to be the most important factor determining yellow-billed cuckoo presence in the forest patches, while no patch characteristics at the larger scale of habitat patches were important. The regional spatial variable was important in two of the three analysis techniques. Model validation using an independent data set of surveys (conducted 1987–1990) found 76–82% model accuracy for all the statistical techniques used. Our results show that the spatial scale at which habitat characteristics are measured influences the suitability of forest patches. This multi-scale patch and model selection approach to habitat suitability analysis can readily be generalized for use with other organisms and systems.     

A leakiness index for assessing landscape function using remote sensing

The cover, number, size, shape, spatial arrangement and orientation of vegetation patches are attributes that have been used to indicate how well landscapes function to retain, not ‘leak’, vital system resources such as rainwater and soil. We derived and tested a directional leakiness index (DLI) for this resource retention function. We used simulated landscape maps where resource flows over map surfaces were directional and where landscape patch attributes were known. Although DLI was most strongly related to patch cover, it also logically related to patch number, size, shape, arrangement and orientation. If the direction of resource flow is multi-directional, a variant of DLI, the multi-directional leakiness index (MDLI) can be used. The utility of DLI and MDLI was demonstrated by applying these indices to three Australian savanna landscapes differing in their remotely sensed vegetation patch attributes. These leakiness indices clearly positioned these three landscapes along a function-dysfunction continuum, where dysfunctional landscapes are leaky (poorly retain resources). This revised version was published online in July 2006 with corrections to the Cover Date.     

Creating landscape patterns by forest cutting: Ecological consequences and principles

Landscape structural characteristics, such as patch size, edge length, and configuration, are altered markedly when management regimes are imposed on primeval landscapes. The ecological consequences of clearcutting patterns were explored by using a model of the dispersed patch or checkerboard system currently practiced on federal forest lands in the western United States. Thresholds in landscape structure were observed on a gradient of percentages of landscape cutover. Probability of disturbance,e.g., wildfire and windthrow, and biotic components,e.g., species diversity and game populations, are highly sensitive to these structural changes. Altering the spatial configuration and size of clearcuts provides an opportunity to create alternative landscapes that differ significantly in their ecological characteristics. Both ecosystem and heterogeneous landscape perspectives are critical in resource management.     

Quantifying patch distribution at multiple spatial scales: Applications to wildlife-habitat models

Multiscale analyses are widely employed for wildlife-habitat studies. In most cases, however, each scale is considered discrete and little emphasis is placed on incorporating or measuring the responses of wildlife to resources across multiple scales. We modeled the responses of three Arctic wildlife species to vegetative resources distributed at two spatial scales: patches and collections of patches aggregated across a regional area. We defined a patch as a single or homogeneous collection of pixels representing 1 of 10 unique vegetation types. We employed a spatial pattern technique, three-term local quadrat variance, to quantify the distribution of patches at a larger regional scale. We used the distance at which the variance for each of 10 vegetation types peaked to define a moving window for calculating the density of patches. When measures of vegetation patch and density were applied to resource selection functions, the most parsimonious models for wolves and grizzly bears included covariates recorded at both scales. Seasonal resource selection by caribou was best described using a model consisting of only regional scale covariates. Our results suggest that for some species and environments simple patch-scale models may not capture the full range of spatial variation in resources to which wildlife may respond. For mobile animals that range across heterogeneous areas we recommend selection models that integrate resources occurring at a number of spatial scales. Patch density is a simple technique for representing such higher-order spatial patterns.     

Quantifying patch distribution at multiple spatial scales: Applications to wildlife-habitat models

Multiscale analyses are widely employed for wildlife-habitat studies. In most cases, however, each scale is considered discrete and little emphasis is placed on incorporating or measuring the responses of wildlife to resources across multiple scales. We modeled the responses of three Arctic wildlife species to vegetative resources distributed at two spatial scales: patches and collections of patches aggregated across a regional area. We defined a patch as a single or homogeneous collection of pixels representing 1 of 10 unique vegetation types. We employed a spatial pattern technique, three-term local quadrat variance, to quantify the distribution of patches at a larger regional scale. We used the distance at which the variance for each of 10 vegetation types peaked to define a moving window for calculating the density of patches. When measures of vegetation patch and density were applied to resource selection functions, the most parsimonious models for wolves and grizzly bears included covariates recorded at both scales. Seasonal resource selection by caribou was best described using a model consisting of only regional scale covariates. Our results suggest that for some species and environments simple patch-scale models may not capture the full range of spatial variation in resources to which wildlife may respond. For mobile animals that range across heterogeneous areas we recommend selection models that integrate resources occurring at a number of spatial scales. Patch density is a simple technique for representing such higher-order spatial patterns.     

Spatial Patterns in European Rabbit Abundance After a Population Collapse

Assessing the associations between spatial patterns in population abundance and environmental heterogeneity is critical for understanding various population processes and for managing species and communities. This study evaluates responses in the abundance of the European rabbit (Oryctolagus cuniculus), an important prey for predators of conservation concern in Mediterranean ecosystems, to environmental heterogeneity at different spatial scales. Multi-scale habitat models of rabbit abundance in three areas of Doñana, south-western Spain, were developed using a spatially extensive dataset of faecal pellet counts as an abundance index. The best models included habitat variables at the three spatial scales examined: distance from lagoons (broad scale), mean landscape shrub coverage and interspersion of pastures (home-range scale), and shrub and pasture cover (microhabitat scale). These variables may well have been related to the availability of food and refuge for the species at the different scales. However, the models’ fit to data and their predictive accuracy for an independent sample varied among the study regions. Accurate predictions in some areas showed that the combination of variables at various spatial scales can provide a reliable method for assessing the abundance of ecologically complex species such as the European rabbit over large areas. On the other hand, the models failed to identify abundance patterns in a population that suffered the strongest demographic collapse after viral epidemics, underlining the difficulty of generalizing this approach. In the latter case, factors difficult to implement in static models such as disease history and prevalence, predator regulation and others may underlie the lack of association. Habitat models can provide useful guidelines for the management of landscape attributes relevant to rabbits and help improve the conservation of Mediterranean communities. However, other influential factors not obviously related to environmental heterogeneity should also be analyzed in more detail.     

Dispersal behaviour in fragmented landscapes: Deriving a practical formula for patch accessibility

Dispersal has been shown to be a key determinant of spatially structured populations. One crucial aspect is predicting patch accessibility: the probability r_ij of a certain patch j being reached by individuals starting at another patch i. Patch accessibility r_ij depends on both the landscape structure and the individuals’ dispersal behaviour. To investigate the effects of these factors on r_ij, we developed a simulation model focusing on animal dispersal. Our model analyses show that there is an important intrinsic effect of the interplay between landscape structure and dispersal behaviour on patch accessibility: the competition between patches for migrants. We derive a formula for patch accessibility. This formula is very simple because it just takes distances into account: not only the distance between start patch and target patch, but also between the start patch and all the other patches in the landscape. Despite its simplicity, the formula is able to cover effects such as the competition for migrants. The formula was found to have high predictive power for a variety of movement behaviours (random walk with various degrees of correlation, Archimedean spirals and loops) in any given landscape. The formula can be interpreted as a generic function for patch accessibility for further population dynamics analyses. It also delivers insights into the consequences of dispersal in fragmented landscapes.