Underestimating the effects of spatial heterogeneity due to individual movement and spatial scale: infectious disease as an example

Many ecological and epidemiological studies occur in systems with mobile individuals and heterogeneous landscapes. Using a simulation model, we show that the accuracy of inferring an underlying biological process from observational data depends on movement and spatial scale of the analysis. As an example, we focused on estimating the relationship between host density and pathogen transmission. Observational data can result in highly biased inference about the underlying process when individuals move among sampling areas. Even without sampling error, the effect of host density on disease transmission is underestimated by approximately 50 % when one in ten hosts move among sampling areas per lifetime. Aggregating data across larger regions causes minimal bias when host movement is low, and results in less biased inference when movement rates are high. However, increasing data aggregation reduces the observed spatial variation, which would lead to the misperception that a spatially targeted control effort may not be very effective. In addition, averaging over the local heterogeneity will result in underestimating the importance of spatial covariates. Minimizing the bias due to movement is not just about choosing the best spatial scale for analysis, but also about reducing the error associated with using the sampling location as a proxy for an individual’s spatial history. This error associated with the exposure covariate can be reduced by choosing sampling regions with less movement, including longitudinal information of individuals’ movements, or reducing the window of exposure by using repeated sampling or younger individuals.

     

Ownership and ecosystem as sources of spatial heterogeneity in a forested landscape,Wisconsin, USA

The interaction between physical environment and land ownership in creating spatial heterogeneity was studied in largely forested landscapes of northern Wisconsin, USA. A stratified random approach was used in which 2500-ha plots representing two ownerships (National Forest and private non-industrial) were located within two regional ecosystems (extremely well-drained outwash sands and moderately well-drained moraines). Sixteen plots were established, four within each combination of ownership and ecosystem, and the land cover on the plots was classified from aerial photographs using a modified form of the Anderson (U.S. Geological Survey) land use and land cover classification system.Upland deciduous forests dominated by northern hardwoods were common on the moraines for both ownerships. On the outwash, the National Forest was dominated by pine plantations, upland deciduous forests, and upland regenerating forests (as defined by <50% canopy coverage). In contrast, a more even distribution among the classes of upland forest existed on private land/outwash. A strong interaction between ecosystem and ownership was evident for most comparisons of landscape structure. On the moraine, the National Forest ownership had a finer grain pattern with more complex patch shapes compared to private land. On the outwash, in contrast, the National Forest had a coarser grain pattern with less complex patch shapes compared to private land. When patch size and shape were compared between ecosystems within an ownership, statistically significant differences in landscape structure existed on public land but not on private land. On public land, different management practices on the moraine and outwash, primarily related to timber harvesting and road building, created very different landscape patterns. Landscape structure on different ecosystems on private land tended to be similar because ownership was fragmented in both ecosystems and because ownership boundaries often corresponded to patch boundaries on private land. A complex relationship exits between ownership, and related differences in land use, and the physical environment that ultimately constrains land use. Studies that do not consider these interactions may misinterpret the importance of either variable in explaining variation in landscape patterns.     

Partitioning spatial,environmental, and community drivers of ecosystem functioning

Landscape Ecology - Community composition, environmental variation, and spatial structuring can influence ecosystem functioning, and ecosystem service delivery. While the role of space in...     

Physical drivers of seagrass spatial configuration: the role of thresholds

Context

Seagrass landscapes vary substantially in extent and pattern, resulting from depth zonation and hydrodynamic stress gradients and may exhibit threshold behavior in response to changes in physical drivers. Seagrass landscapes persist in a delicate balance between processes of disturbance and recovery and therefore may exhibit behavior typical of classic critical systems.

Objectives

Examine how hydrodynamic drivers and physical setting influence seagrass landscape composition and configuration. Determine if seagrass patch size distributions typify patterns observed for critical systems.

Methods

We used landscape metrics to quantify the spatial configuration of seagrass and then modeled the response of these metrics to wave energy, tidal current speed, and water depth at 62 estuarine sites in North Carolina, USA. Seagrass landscapes were representative of cover types observed in the estuary generated by wave energy.

Results

Percent cover, patch size, and number of patches all declined with increasing wave energy. Threshold behavior occurred at wave energy change points between 675–774 J m^?1. Seagrass landscapes differed in spatial configuration and physical setting, above and below change points. There was moderate support for a power law relationship for patch size distribution across a wide range of seagrass landscape cover and wave energy.

Conclusions

With weather extremes on the rise, much of this estuarine seagrass will be exposed to increased wave energy. Where seagrass exists just below the wave energy change points, increases in wave energy could tip those habitats into a new stable state of lower cover resulting in less cover overall in the estuary.

     

Quantifying spatiotemporal drivers of environmental heterogeneity in Kruger National Park,South Africa

Context

Environmental heterogeneity is considered an important mechanism of biodiversity. How environmental heterogeneity is characterised by the compositional, structural and functional variation of biotic and abiotic components is a central research theme in conservation.

Objectives

We explore how environmental heterogeneity relates to the underlying physical landscape template and how that relationship changes over space and time. We examine how, in some areas, environmental heterogeneity may also be driven by dynamic ecological processes, and how this relates to patterns of plant species richness.

Method

We use local geographically weighted regression to spatially partition environmental heterogeneity, measured as Landsat spectral variance, into the portion explained by stable physical landscape properties (R²) and the portion unexplained (1?R²) which we term landscape complexity. We explore how this relationship varies spatially and temporally as a function of dynamic ecological processes such as rainfall and season in Kruger National Park, as well as plant species richness at landscape scales.

Results

The significance and direction of relationships varied over space and time and as a function of rainfall and season. R² values generally decreased in higher rainfall summer months and revealed patterns describing the importance of known stable factors relative to unknown dynamic factors. Landscape complexity (1?R²) explained over 70 % of variation in species richness.

Conclusions

Rainfall and seasonality are important drivers of environmental heterogeneity. The spatial arrangement and magnitude of model agreement helped disentangle the relative influence of the physical landscape template on environmental heterogeneity. Given the high correlation with species richness, landscape complexity provides complementary guidance to biodiversity research and monitoring prioritization.

     

Shifting concepts of urban spatial heterogeneity and their implications for sustainability

Context

Spatial heterogeneity has myriad influences on ecosystem processes, ecosystem services, and thus the sustainability of urban areas. It acts as a medium for urban design, planning, and management to determine how processes affecting sustainability can operate and interact. Therefore, how spatial heterogeneity is conceptualized and measured in cities is crucial for enhancing sustainability.

Objectives

We show that the two most commonly used, but contrasting paradigms of urban ecology, ecology IN versus ecology OF the city, determine how spatial heterogeneity is thought of and used in different ways. We identify the key implications of these theoretical contrasts for the practice and assessment of sustainability in urban areas.

Methods

We review and compare the different ways in which ecology IN versus ecology OF the city affect how to conceptualize, model and map urban spatial heterogeneity. We present a new framework to guide the comparison of spatial heterogeneity under the two paradigms.

Results and conclusion

The integrative nature of this new framework becomes apparent under the ecology OF the city paradigm, because it recognizes the hybrid social and bioecological nature of heterogeneity in urban ecosystems. The hybrid approach to patchiness resonates with the three pillars of sustainability—environment, society, and economy. We exemplify how the more comprehensive and integrated framework of spatial heterogeneity under the ecology OF the city paradigm (1) supports more effective measurement and integration of the three components of sustainability, (2) improves management of heterogeneous urban ecosystems, and (3) satisfies calls for improved ecological tools to support urban ecosystem design.

     

Summer spatial patterning of chukars in relation to free water in western Utah

Free water is considered important to wildlife in arid regions. In the western United States, thousands of water developments have been built to benefit wildlife in arid landscapes. Agencies and researchers have yet to clearly demonstrate their effectiveness. We combined a spatial analysis of summer chukar (Alectoris chukar) covey locations with dietary composition analysis in western Utah. Our specific objectives were to determine if chukars showed a spatial pattern that suggested association with free water in four study areas and to document summer dietary moisture content in relation to average distance from water. The observed data for the Cedar Mountains study area fell within the middle of the random mean distance to water distribution suggesting no association with free water. The observed mean distance to water for the other three areas was much closer than expected compared to a random spatial process, suggesting the importance of free water to these populations. Dietary moisture content of chukar food items from the Cedar Mountains (59%) was significantly greater (P < 0.05) than that of birds from Box Elder (44%) and Keg-Dugway (44%). Water developments on the Cedar Mountains are likely ineffective for chukars. Spatial patterns on the other areas, however, suggest association with free water and our results demonstrate the need for site-specific considerations. Researchers should be aware of the potential to satisfy water demand with pre-formed and metabolic water for a variety of species in studies that address the effects of wildlife water developments. We encourage incorporation of spatial structure in model error components in future ecological research.     

Interactions between past land use, life-history traits and understory spatial heterogeneity

Past land use has contributed to variability in the distribution of herbaceous species by reducing plant abundance and altering species’ chances of recolonizing suitable habitat. Land use may also influence plant heterogeneity by changing environmental conditions within stands. We compared the variability of understory herb abundance in southern Appalachian forests with different land-use histories to examine how past land use influenced plant heterogeneity. The cover of eleven focal species or genera was estimated and mineral soil concentrations were determined during 2001 and 2002 in eight stands that were farmed, logged, or had no disturbance history (reference) in western North Carolina. Analysis of the coefficients of variation revealed that the abundance of understory plants was more heterogeneous in disturbed stands compared with reference stands. However, when nutrient availability differences were accounted for by detrending the plant cover data, understory variability within stands declined, and no differences between disturbed and reference stands could be distinguished. This finding suggests that nutrient availability has important effects on plant heterogeneity, which depend on past land use. Species dispersal, seed size, and phenology also explained variability in the spatial heterogeneity of plants, but generally only before soil nutrient differences were statistically controlled. In addition to demonstrating that past land use has long-term effects on plant heterogeneity, these results indicate that soil nutrients may play different roles in determining vegetation patterns in historically altered and unaltered forests.     

Ecosystem-scale spatial heterogeneity of stable isotopes of soil nitrogen in African savannas

Soil ¹⁵N is a natural tracer of nitrogen (N) cycling. Its spatial distribution is a good indicator of processes that are critical to N cycling and of their controlling factors integrated both in time and space. The spatial distribution of soil δ¹⁵N and its underlying drivers at sub-kilometer scales are rarely investigated. This study utilizes two sites (dry vs. wet) from a megatransect in southern Africa encompassing locations with similar soil substrate but different rainfall and vegetation, to explore the effects of soil moisture and vegetation distribution on ecosystem-scale patterns of soil δ¹⁵N. A 300-m long transect was set up at each site and surface soil samples were randomly collected for analyses of δ¹⁵N, %N and nitrate content. At each soil sampling location the presence of grasses, woody plants, Acacia species (potential N fixer) as well as soil moisture levels were recorded. A spatial pattern of soil δ¹⁵N existed at the dry site, but not at the wet site. Woody cover distribution determined the soil δ¹⁵N spatial pattern at ecosystem-scale; however, the two Acacia species did not contribute to the spatial pattern of soil δ¹⁵N. Grass cover was negatively correlated with soil δ¹⁵N at both sites owing to the lower foliar δ¹⁵N values of grasses. Soil moisture did not play a role in the spatial pattern of soil δ¹⁵N at either site. These results suggest that vegetation distribution, directly, and water availability, indirectly, affect the spatial patterns of soil δ¹⁵N through their effects on woody plant and grass distributions.     

Environmental drivers of community diversity in a neotropical urban landscape: a multi-scale analysis

Context

Many aquatic communities are linked by the aerial dispersal of multiple, interacting species and are thus structured by processes occurring in both the aquatic and terrestrial compartments of the ecosystem.

Objectives

To evaluate the environmental factors shaping the aquatic macroinvertebrate communities associated with tank bromeliads in an urban landscape.

Methods

Thirty-two bromeliads were georeferenced to assess the spatial distribution of the aquatic meta-habitat in one city. The relative influence of the aquatic and terrestrial habitats on the structure of macroinvertebrate communities was analyzed at four spatial scales (radius = 10, 30, 50, and 70 m) using redundancy analyses.

Results

We sorted 18,352 aquatic macroinvertebrates into 29 taxa. Water volume and the amount of organic matter explained a significant part of the taxa variance, regardless of spatial scale. The remaining variance was explained by the meta-habitat size (i.e., the water volume for all of the bromeliads within a given surface area), the distance to the nearest building at small scales, and the surface area of buildings plus ground cover at larger scales. At small scales, the meta-habitat size influenced the two most frequent mosquito species in opposite ways, suggesting spatial competition and coexistence. Greater vegetation cover favored the presence of a top predator.

Conclusions

The size of the meta-habitat and urban landscape characteristics influence the structure of aquatic communities in tank bromeliads, including mosquito larval abundance. Modifications to this landscape will affect both the terrestrial and aquatic compartments of the urban ecosystem, offering prospects for mosquito management during urban planning.

     

Spatial and temporal heterogeneity of forest site productivity drivers: a case study within the eastern boreal forests of Canada

Forest productivity is driven by a suite of direct climatic and non-climatic factors that are transient or permanent. The kind of productivity driver and the nature of their effects vary by species, and scale dependencies potentially complicate these relationships. This study explored productivity-driver relations in eastern Boreal Canada and determined spatial effects in productivity control when expressed with stand dominant height at a reference age (site index). Data from 4,217 temporary sample plots obtained from boreal mixedwood and conifer bioclimatic domains, and with varied species composition, were used in this study. A single-level global model that assumes equal sensitivities across spatial scales was calibrated and compared with three alternative models reflecting different hypotheses on possible spatial heterogeneities. Alternative models were calibrated by plot-level soil deposit types (microscale), landscape dominant deposits (mesoscale) and bioclimatic domains (macroscale). A marked difference between the global and alternative models was observed, suggesting that a single global model does not sufficiently reflect existing heterogeneity in productivity-driver relationships. A combination of macro- and microscale models provided the best explanation of site index. Results further showed that site index is mainly driven by species composition (complementarity effects of aspen and jack pine compositions) and stand diameter structural diversity effects. It is concluded that successional changes, more than direct climatic effects, drive productivity.     

Effects of spatial habitat heterogeneity on habitat selection and annual fecundity for a migratory forest songbird

Understanding how spatial habitat patterns influence abundance and dynamics of animal populations is a primary goal in landscape ecology. We used an information-theoretic approach to investigate the association between habitat patterns at multiple spatial scales and demographic patterns for black-throated blue warblers (Dendroica caerulescens) at 20 study sites in west-central Vermont, USA from 2002 to 2005. Sites were characterized by: (1) territory-scale shrub density, (2) patch-scale shrub density occurring within 25 ha of territories, and (3) landscape-scale habitat patterns occurring within 5 km radius extents of territories. We considered multiple population parameters including abundance, age ratios, and annual fecundity. Territory-scale shrub density was most important for determining abundance and age ratios, but landscape-scale habitat structure strongly influenced reproductive output. Sites with higher territory-scale shrub density had higher abundance, and were more likely to be occupied by older, more experienced individuals compared to sites with lower shrub density. However, annual fecundity was higher on sites located in contiguously forested landscapes where shrub density was lower than the fragmented sites. Further, effects of habitat pattern at one spatial scale depended on habitat conditions at different scales. For example, abundance increased with increasing territory-scale shrub density, but this effect was much stronger in fragmented landscapes than in contiguously forested landscapes. These results suggest that habitat pattern at different spatial scales affect demographic parameters in different ways, and that effects of habitat patterns at one spatial scale depends on habitat conditions at other scales.     

Interplay between physical and predator landscapes affects transferability of shorebird distribution models

Coastal landscapes with extensive intertidal mudflats provide non-breeding habitat for Arctic shorebirds. Few attempts have been made to develop and test landscape-level models predicting the intertidal distribution of these birds. We modelled the distribution of a Holarctic species, Dunlin (Calidris alpina), at a hemispherically important non-breeding site, the Fraser River Delta, British Columbia, Canada, in seasons with different predator landscapes. We trained the models during a season when nocturnal predators were common and tested temporal transferability of the models on independent datasets when nocturnal predators were absent. Snowy Owls (Nyctea scandiaca) influenced Dunlin distribution and thus model transferability. After accounting for their presence, models displayed good to excellent discrimination, i.e. prediction of the instantaneous and cumulative (over low tide period) probability of mudflat use by Dunlin, in fore- and backcasting applications. Model calibration was good or else, where over-prediction was observed, the reason for the bias was identified. The distribution models may predict mudflat use by Dunlin and possibly related species given relevant data describing the intertidal landscape. The models are amenable to GIS application, describe the amount of use per hectare of the intertidal zone and can be used to determine and visualise relative and absolute suitability of intertidal areas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Water availability is a principal driver of large-scale land cover spatial heterogeneity in sub-Saharan savannahs

Landscape Ecology - The heterogeneous mosaic nature of African savannah vegetation is a key aspect of its ecology. This study evaluates mosaic distributions and characteristics across sub-Saharan...     

Forest recovery since 1860 in a Mediterranean region: drivers and implications for land use and land cover spatial distribution

Context

Land use and land cover (LULC) change is a major part of environmental change. Understanding its long-term causes is a major issue in landscape ecology.

Objectives

Our aim was to characterise LULC transitions since 1860 and assess the respective and changing effects of biophysical and socioeconomic drivers on forest, arable land and pasture in 1860, 1958 and 2010, and of biophysical, socioeconomic and distance from pre-existing forest on forest recovery for the two time intervals.

Methods

We assessed LULC transitions by superimposing 1860, 1958 and 2010 LULCs using a regular grid of 1 × 1 km points, in a French Mediterranean landscape (195,413 ha). We tested the effects of drivers using logistic regressions, and quantified pure and joint effects by deviance partitioning.

Results

Over the whole period, the three main LULCs were spatially structured according to land accessibility and soil productivity. LULC was driven more by socioeconomic than biophysical drivers in 1860, but the pattern was reversed in 2010. A widespread forest recovery mainly occurred on steeper slopes, far from houses and close to pre-existing forest, due to traditional practice abandonment. Forest recovery was better explained by biophysical than by socioeconomic drivers and was more dependent on distance from pre-existing forest between 1958 and 2010.

Conclusions

Our results showed a shift in drivers of LULC and forest recovery over the last 150 years. Contrary to temperate regions, the set-aside of agricultural practices on difficult land has strengthened the link between biophysical drivers and LULC distribution over the last 150 years.

     

The response of elephants to the spatial heterogeneity of vegetation in a Southern African agricultural landscape

Based on the agricultural landscape of the Sebungwe in Zimbabwe, we investigated whether and how the spatial distribution of the African elephant (Loxodonta africana) responded to spatial heterogeneity of vegetation cover based on data of the early 1980s and early 1990s. We also investigated whether and how elephant distribution responded to changes in spatial heterogeneity between the early 1980s and early 1990s. Vegetation cover was estimated from a normalised difference vegetation index (NDVI). Spatial heterogeneity was estimated from a new approach based on the intensity (i.e., the maximum variance exhibited when a spatially distributed landscape property such as vegetation cover is measured with a successively increasing window size or scale) and dominant scale (i.e., the scale or window size at which the intensity is displayed). We used a variogram to quantify the dominant scale (i.e., range) and intensity (i.e., sill) of NDVI based congruent windows (i.e., 3.84 km × 3.84 km in a 61 km × 61 km landscape). The results indicated that elephants consistently responded to the dominant scale of spatial heterogeneity in a unimodal fashion with the peak elephant presence occurring in environments with dominant scales of spatial heterogeneity of around 457–734 m. Both the intensity and dominant scale of spatial heterogeneity predicted 65 and 68% of the variance in elephant presence in the early 1980s and in the early 1990s respectively. Also, changes in the intensity and dominant scale of spatial heterogeneity predicted 61% of the variance in the change in elephant distribution. The results imply that management decisions must take into consideration the influence of the levels of spatial heterogeneity on elephants in order to ensure elephant persistence in agricultural landscapes.