Estimating the spatial scales of landscape effects on abundance

Context

Spatial variation in abundance is influenced by local- and landscape-level environmental variables, but modeling landscape effects is challenging because the spatial scales of the relationships are unknown. Current approaches involve buffering survey locations with polygons of various sizes and using model selection to identify the best scale. The buffering approach does not acknowledge that the influence of surrounding landscape features should diminish with distance, and it does not yield an estimate of the unknown scale parameters.

Objectives

The purpose of this paper is to present an approach that allows for statistical inference about the scales at which landscape variables affect abundance.

Methods

Our method uses smoothing kernels to average landscape variables around focal sites and uses maximum likelihood to estimate the scale parameters of the kernels and the effects of the smoothed variables on abundance. We assessed model performance using a simulation study and an avian point count dataset.

Results

The simulation study demonstrated that estimators are unbiased and produce correct confidence interval coverage except in the rare case in which there is little spatial autocorrelation in the landscape variable. Canada warbler abundance was more highly correlated with site-level measures of NDVI than landscape-level NDVI, but the reverse was true for elevation. Canada warbler abundance was highest when elevation in the surrounding landscape, defined by an estimated Gaussian kernel, was between 1300 and 1400 m.

Conclusions

Our method provides a rigorous way of formally estimating the scales at which landscape variables affect abundance, and it can be embedded within most classes of statistical models.

     

Linking macrodetritivore distribution to desiccation resistance in small forest fragments embedded in agricultural landscapes in Europe

Purpose

Most of the agricultural landscape in Europe, and elsewhere, consists of mosaics with scattered fragments of semi-natural habitat like small forest fragments. Mutual interactions between forest fragments and agricultural areas influence ecosystem processes such as nutrient cycling, a process strongly mediated by the macrodetritivore community, which is however, poorly studied. We investigated macrodetritivore distribution patterns at local and landscape-level and used a key functional trait (desiccation resistance) to gain mechanistic insights of the putative drivers.

Methods

Macrodetritivores were sampled in forest edges-centres of 224 European forest fragments across 14 landscapes opposing in land use intensity. We used a multilevel analysis of variance to assess the relative contribution of different spatial scales in explaining activity-density and Shannon-diversity of woodlice and millipedes, together with a model-based analysis of the multivariate activity-density data testing the effect on species composition. Secondly, we tested if desiccation resistance of macrodetritivores varied across communities at different spatial scales using linear mixed effect models.

Results

Forest edge-centre and landscape use intensity determined activity-density and community composition of macrodetritivores in forest fragments, while fragment characteristics like size and continuity were relatively unimportant. Forest edges and higher intensity landscapes supported higher activity-density of macrodetritivores and determined species composition. Forest edges sustained woodlouse communities dominated by more drought tolerant species.

Conclusions

Landscape use intensity and forest edges are main drivers in macrodetritivore distribution in forest fragments with desiccation resistance a good predictor of macrodetritivore distribution. Key functional traits can help us to predict changes in community structure in changing landscapes.

     

Effects of spatial scale and taxonomic group on partitioning of butterfly and bird diversity in the Great Basin,USA

Different taxonomic groups perceive and respond to the environment at different scales. We examined the effects of spatial scale on diversity patterns of butterflies and birds in the central Great Basin of the western USA. We partitioned the landscape into three hierarchical spatial levels: mountain ranges, canyons, and sites within can yons. We evaluated the relative contribution of each level to species richness and quantified changes in species composition at each level. Using additive partitioning, we calculated the contribution of spatial level to overall species diversity. Both canyon and mountain range had significant effects on landscape-level species richness of butterflies and birds. Species composition of butterflies was more similar in space than species composition of birds, but assemblages of both groups that were closer together in space were less similar than assemblages that were further apart. These results likely reflect differences in resource specificity and the distribution of resources for each group. Additive partitioning showed that alpha diversity within canyon segments was the dominant component of overall species richness of butterflies but not of birds. As the size of a sampling unit increased, its contribution to overall species richness of birds increased monotonically, but the relationship between spatial scale and species richness of butterflies was not linear. Our work emphasizes that the most appropriate scales for studying and conserving different taxonomic groups are not the same. The ability of butterflies and birds to serve as surrogate measures of each others diversity appears to be scale-dependent.This revised version was published online in May 2005 with corrections to the Cover Date.     

Habitat and food supply across multiple spatial scales influence the distribution and abundance of a nocturnal aerial insectivore

Context

Conservation research often focuses on individual threats at a single spatial scale, but population declines can result from multiple stressors occurring at different spatial scales. Analyses incorporating alternative hypotheses across spatial scales allow more robust evaluation of the ecological processes underlying population declines.

Objectives

Populations of many aerially insectivorous birds are declining, yet conservation efforts remain focused on habitat due to an absence of data on changes in prey availability. We evaluate the potential for prey and habitat availability at multiple spatial scales to influence a population of eastern whip-poor-wills (Antrostomus vociferous).

Methods

We assess relationships between landcover (topographical map and satellite imagery) and insect abundance (moths and beetles from blacklight traps), and whip-poor-will distribution and abundance within eastern Canada using Ontario breeding bird atlas data (1980s and 2000s), acoustic recordings (regional), and point counts (local).

Results

Whip-poor-will occurrence in both atlas time periods was positively associated with forest area and fragmentation, but only a delayed effect of urban area explained reductions in detection. Contemporary regional whip-poor-will presence was positively related to moth abundance, and local whip-poor-will abundance was best predicted by area of open-canopy forest, anthropogenic linear disturbance density, and beetle abundance. Our finding that bird presence and abundance were associated with human activity and insect abundance across spatial scales suggests factors beyond habitat structure are likely driving population declines in whip-poor-wills and other aerial insectivores.

Conclusions

This study demonstrates the importance of examining multiple hypotheses, including seasonally and locally variable food availability, across a range of spatial scales to direct conservation efforts.

     

Connectivity and habitat type shape divergent dispersal behavior in a desert-dwelling fish

Context

Dispersal has important fitness consequences for individuals, populations, and species. Despite growing theoretical insights into the evolution of dispersal, its behavioral underpinnings remain empirically understudied, limiting our understanding of the extent and impact of responses to landscape-level heterogeneity of environments, and increasing the risk of inferring species-level responses from biased population sampling.

Objectives

We asked if predictable ecological variation among naturally fragmented arid waterbodies is correlated with disparate dispersal responses of populations of the desert goby Chlamydogobius eremius, which naturally inhabits two habitat “types” (permanent springs, ephemeral rivers), and different levels of hydrological connectivity (high and low) that potentially convey different costs and benefits of dispersal.

Methods

To test for possible behavioral divergence between such populations, we experimentally compared the movement behaviors (correlates of emigration and exploration) of wild-caught fish. We used two biologically relevant spatial scales to test movement relevant to different stages of the dispersal process.

Results

Behavior differed at both spatial scales, suggesting that alternative dispersal strategies enable desert gobies to exploit diverse habitat patches. However, while emigration was best predicted by the connectivity (flood risk) of fish habitats, exploration was linked to their habitat type (spring versus river).

Conclusions

Our findings demonstrate that despite a complex picture of ecological variation, key landscape factors have an overarching effect on among-population variation in dispersal traits. Implications include the maintenance of within-species variation, potentially divergent evolutionary trajectories of naturally or anthropogenically isolated populations, and the direction of future experimental studies on the ecology and evolution of dispersal behavior.

     

Implications of rescaling rules for multi-scaled habitat models

Recent work in landscape ecology suggests that organisms use environmental cues at a variety of scales to select habitat. As a result, habitat studies that evaluate environmental conditions at multiple spatial scales have become increasingly common. We examined whether the way in which data are rescaled influences inferred relationships between organisms and habitat features. Using a habitat model developed at fine scales, we systematically rescaled habitat (canopy density, slope, and cover type) and distribution maps according to a variety of different rescaling rules, including spatial averaging, thresholding, presence/absence, and majority. We found that the spatial autocorrelation of habitat data interacts with rescaling rules to alter the correspondence between species presence and habitat across scale. Different rules lead to substantially divergent and sometimes opposite correlations among the species and habitat features on the landscape. Such differences in interpretation due to variation in methodology can lead to very different interpretations of a species habitat requirements and thus have important implications for both ecology and conservation.     

Linking greenhouse gas emissions to urban landscape structure: the relevance of spatial and thematic resolutions of land use/cover data

Context

Emissions of greenhouse gases in urban areas play an important role in climate change. Increasing attention has been given to urban landscape structure–emission relationships (SERs). However, it remains unknown if and to what extent SERs are dependent on observational scale.

Objective

To assess how changing observational scales (in terms of spatial and thematic resolutions) of urban landscape structure affect SERs.

Methods

We examined correlations between 16 landscape metrics and greenhouse gas emissions across 52 European cities, through (1) systematic manipulation of spatial and thematic resolutions of the urban land use/cover (ULUC) dataset, and (2) comparison between available standard ULUC datasets with different spatial resolutions.

Results

Our analyses showed that the observed SERs significantly depend on both thematic and spatial resolutions of the ULUC data. For the 16 landscape metrics, we found diverse spatial/thematic scaling relations exhibiting monotonic, hump-shaped or scale-invariant trends. For different landscape metrics, the SERs were strongest at different spatial scales, suggesting that there is no consistent scaling relation over those observational scales.

Conclusions

SERs are highly sensitive to spatial and thematic resolutions of landscape data. To avoid the problem of ‘ecological fallacy,’ important caveats should be taken for interpretations based on single landscape metrics. Particular consideration should be paid to metrics that are easily interpretable, predictable in scaling behaviors, and important for shaping SERs, such as PLAND, ED, and LPI. Systematic investigations on scaling behaviors of SERs over well-defined scale domains are encouraged in future studies linking greenhouse gas emissions and urban landscape structure.

     

Does the response of bird assemblages to fire mosaic properties vary among spatial scales and foraging guilds?

Context

An increasing number of studies have investigated the impact of environmental heterogeneity on faunal assemblages when measured at multiple spatial scales. Few studies, however, have considered how the effects of heterogeneity on fauna vary with the spatial scale at which the response variable is characterised.

Objectives

We investigated the relationship between landscape properties in a region characterised by diverse fire mosaics, and the structure and composition of avian assemblages measured at both the site- (1 ha) and landscape-scale (100 ha).

Methods

We surveyed birds and calculated spatial landscape properties in sub-tropical woodlands of central Queensland, Australia.

Results

Environmental heterogeneity, as measured by topographic complexity, was consistently important for bird species richness and composition. However, the explanatory power of topographic complexity varied depending on the spatial scale and the component of diversity under investigation. We found different correlates of richness within particular foraging guilds depending on the scale at which richness was measured. Extent of long-unburnt habitat (>10 years since fire) was the most important variable for the landscape-scale richness of frugivores, insectivores and canopy feeders, whereas environmental heterogeneity in the surrounding landscape was more important for site-scale richness of these foraging guilds.

Conclusions

The response of species richness to landscape characteristics varies among scales, and among components of diversity. Thus, depending on the scale at which a biodiversity conservation goal is conceptualised—maximising richness at a site, or across a landscape—different landscape management approaches may be preferred.

     

Predicting across scales: Theory development and testing

Landscape ecologists deal with processes that occur at a variety of temporal and spatial scales. The ability to make predictions at more than one level of resolution requires identification of the processes of interest and parameters that affect this process at different scales, the development of rules to translate information across scales, and the ability to test these predictions at the relevant spatial and temporal scales. This paper synthesizes discussions from a workshop on Predicting Across Scales: Theory Development and Testing that was held to discuss current research on scaling and to identify key research issues.     

multifit: an R function for multi-scale analysis in landscape ecology

Context

Multi-scale analyses are a common approach in landscape ecology. Their aim is to find the appropriate spatial scale for a particular landscape attribute in order to perform a correct interpretation of results and conclusions.

Objectives

I present an R function that performs statistical analysis relating a biological response with a landscape attribute at a set of specified spatial scales and extracts the statistical strength of the models through a specified criterion index. Also, it draws a plot with the value of these indexes, allowing the user to choose the most appropriate spatial scale. This paper introduces the usage of multifit and demonstrates its functionality through a case study.

Conclusions

The spatial scale at which ecologists conduct studies may change study outcomes and conclusions. Because of this, landscape ecologists commonly conduct multi-scale studies in order to establish an appropriate spatial scale for particular biological or ecological responses. The tool presented here allows ecologists to simultaneously run several statistical models for a response variable and a specified set of spatial scales, automating the process of multi-scale analysis.

     

Scaling patterns of biomass and soil properties: an empirical analysis

We argue that studies at multiple scales must necessarilychange the extent of measurements, not just the spacing, in order toeffectivelycapture information regarding processes at multiple scales. We have implementeda multi-scale sampling scheme using transects of 10 cm, 1m, 10 m, 100 m, and 1 km ateach of four sites along an elevational gradient from dry foothills forest toalpine tundra in the Front Range of Colorado; these four sites form anadditional transect of 22 km. Along each of these transects wetookten equally spaced soil cores and measured variables important in determiningboth microbial and plant community structure: soil water content, organicmattercontent, pH, and total soil biomass. With this sampling scheme we are able totreat scale as an independent variable in our analyses, and our data show thatboth particular sites and particular variables can determine whether estimatesof mean values are scale-dependent or not. A geostatistical analysis using allof our data shows common relationships between scales across ecologicallydiverse sites; biomass shows the most complex pattern of distribution acrossscales, as measured by fractal dimension. Our analyses also reveal theinadequacy of several standard geostatistical models when applied to data frommultiple scales of measurement – we recommend the use of the boundedpowerlaw model in such cases. We hypothesize that because biological communitiesmustrespond simultaneously to multiple variables with differing patterns of spatialvariation, the spatial variation of biological communities will be at least ascomplex as the most complex environmental variable at any given site.This revised version was published online in May 2005 with corrections to the Cover Date.     

Occupancy pattern of a long-horned beetle in a variegated forest landscape: linkages between tree quality and forest cover across spatial scales

Context

Interactions between landscape-scale processes and fine-grained habitat heterogeneity are usually invoked to explain species occupancy in fragmented landscapes. In variegated landscapes, however, organisms face continuous variation in micro-habitat features, which makes necessary to consider ecologically meaningful estimates of habitat quality at different spatial scales.

Objectives

We evaluated the spatial scales at which forest cover and tree quality make the greatest contribution to the occupancy of the long-horned beetle Microplophorus magellanicus (Coleoptera: Cerambycidae) in a variegated forest landscape.

Methods

We used averaged data of tree quality (as derived from remote sensing estimates of the decay stage of single trees) and spatially independent pheromone-baited traps to model the occurrence probability as a function of multiple cross-scale combinations between forest cover and tree quality (with scales ranging between 50 and 400 m).

Results

Model support and performance increased monotonically with the increasing scale at which tree quality was measured. Forest cover was not significant, and did not exhibit scale-specific effects on the occurrence probability of M. magellanicus. The interactive effect between tree quality and forest cover was stronger than the independent (additive) effects of tree quality and particularly forest cover. Significant interactions included tree quality measured at spatial scales ≥200 m, but cross-scale interactions occurred only in four of the seven best-supported models.

Conclusions

M. magellanicus respond to the high-quality trees available in the landscape rather than to the amount of forest per se. Conservation of viable metapopulations of M. magellanicus should consider the quality of trees at spatial scales >200 m.

     

Using fractal analysis to assess how species perceive landscape structure

To develop a species-centered definition of landscapes, I suggest using a fractal analysis of movement patterns to identify the scales at which organisms are interacting with the patch structure of the landscape. Significant differences in the fractal dimensions of movement patterns of two species indicate that the species may be interacting with the patch structure at different scales. Fractal analysis therefore permits comparisons of landscape perceptions of different species within the same environment.I tested the utility of this fractal application by analyzing the movement patterns of three species of acridid grasshoppers (Orthoptera) in a grassland mosaic. The largest species moved up to 6 times faster than the two smaller species, and species exhibited different responses to microlandscape structure within 25-m² plots. Further, the largest species exhibited different responses to microlandscape structure in two pastures subjected to different intensities of cattle grazing. This species thus is able to integrate information on landscape structure at broad spatial scales. Fractal analysis of movement patterns revealed that the two small species had significantly more tortuous patterns than the larger species, which suggests that these species are interacting with patch structure at a finer scale of resolution than the large species. Fractal analysis can be used to identify the perceptive resolution of a species; that is, the spatial grain and extent at which they are able to perceive and respond to heterogeneity. Analysis of movement patterns across a range of spatial scale may reveal shifts in fractal dimension that reflect transitions in how species respond to the patch structure of the landscape at different scales.     

Effects of bird and bat exclusion on coffee pest control at multiple spatial scales

Context

Despite the key role of biological control in agricultural landscapes, we still poorly understand how landscape structure modulates pest control at different spatial scales.

Objectives

Here we take an experimental approach to explore whether bird and bat exclusion affects pest control in sun coffee plantations, and whether this service is consistent at different spatial scales.

Methods

We experimentally excluded flying vertebrates from coffee plants in 32 sites in the Brazilian Atlantic Forest, encompassing a gradient of forest cover at landscape (2 km radius) and local (300 m) spatial scales, and quantified coffee leaf loss, as an indicator of herbivory, and fruit set.

Results

Leaf loss decreased with higher landscape forest cover, but this relation was significantly different between treatment and control plants depending on local forest cover. On the other hand, fruit set responded to the interaction between treatment and local forest cover but was not affected by landscape forest cover. More specifically, fruit set increased significantly with local forest cover in exclusion treatments and showed a non-significant decrease in open controls.

Conclusions

These results suggest that services provided by flying vertebrates are modulated by processes occurring at different spatial scales. We posit that in areas with high local forest cover flying vertebrates may establish negative interactions with predaceous arthropods (i.e. intraguild predation), but this would not be the case in areas with low local forest cover. We highlight the importance of employing a multi-scale analysis in systems where multiple species, which perceive the landscape differently, are providing ecosystem services.

     

The emergence of heterogeneity in invasive-dominated grassland: a matter of the scale of detection

Context

Plant invasions of native ecosystems are one of the main causes of declines in biodiversity via system-simplification. Restoring native biodiversity can be particularly challenging in landscapes where invasive species have become dominant and where a new set of feedbacks reinforce an invaded state and preclude restoration actions. We lack an understanding of the response of invaded systems to landscape-level manipulations to restore pattern and process relationships and how to identify these relationships when they do not appear at the expected scale.

Objectives

To better understand how fire and grazing influence landscape-level heterogeneity in invaded landscapes, we assess the scale at which grazing pressure and seasonality mediate the success of re-introducing a historical disturbance regime, grazing driven by fire (termed pyric herbivory), to an invasive plant-dominated landscape.

Methods

We manipulated grazing timing and intensity in exotic grass-dominated grasslands managed for landscape heterogeneity with spring fire and grazing. In pastures under patch-burn grazing management, we evaluated the spatial and temporal variability of plant functional groups and vegetation structure among and within patches managed with separate grazing systems: season-long stocking and intensive early stocking.

Results

Warm- and cool-season grasses exhibited greater among-patch variability in invasive-plant dominated grassland under intensive early grazing than traditional season-long grazing, but landscape-level heterogeneity, as measured through vegetation structure was minimal and invariable under both levels of grazing pressure, which contrasts findings in native-dominated systems. Moreover, within-patch heterogeneity for these functional groups was detected; contrasting the prediction that among-patch heterogeneity, in mesic grasslands, manifests from within-patch homogeneity.

Conclusions

In invaded grasslands, manipulation of grazing pressure as a process that drives heterogeneous vegetation patterns influences native and non-native grass heterogeneity, but not heterogeneity of vegetation structure, within and among patches managed with fire. Fire and grazing-moderated heterogeneity patterns observed in native grass-dominated grasslands likely differ from invasive grass-dominated grasslands with implications for using pyric herbivory in invaded systems.

     

Fractal dimension estimates of a fragmented landscape: sources of variability

Although often seen as a scale-independent measure, we show that the fractal dimension of the forest cover of the Cazaville Region changes with spatial scale. Sources of variability in the estimation of fractal dimensions are multiple. First, the measured phenomenon does not always show the properties of a pure fractal for all scales, but rather exhibits local self-similarity within certain scale ranges. Moreover, some sampling components such as area of sampling unit, the use of a transect in the estimation of the variability of a plane, the location, and the orientation of a transect all affect, to different degrees, the estimation of the fractal dimension. This paper assesses the relative importance of these components in the estimation of the fractal dimension of the spatial distribution of woodlots in a fragmented landscape. Results show that different sources of variability should be considered when comparing fractal dimensions from different studies or regions.     

Scale-specific habitat relationships influence patch occupancy: defining neighborhoods to optimize the effectiveness of landscape-scale grassland bird conservation

Context

Beyond the recognized importance of protecting large areas of contiguous habitat, conservation efforts for many species are complicated by the fact that patch suitability may also be affected by characteristics of the landscape within which the patch is located. Currently, little is known about the spatial scales at which species respond to different aspects of the landscape surrounding an occupied patch.

Objectives

Using grassland bird point count data, we describe an approach to evaluating scale-specific effects of landscape composition on patch occupancy.

Methods

We used data from 793 point count surveys conducted in idle and grazed grasslands across Wisconsin, USA from 2012 to 2014 to evaluate scale-dependencies in the response of grassland birds to landscape composition. Patch occupancy models were used to evaluate the relationship between occupancy and landscape composition at scales from 100 to 3000 m.

Results

Bobolink (Dolichonyx oryzivorus) exhibited a pattern indicating selection for grassland habitats in the surrounding landscape at all spatial scales while selecting against other habitats. Eastern Meadowlark (Sturnella magna) displayed evidence of scale sensitivity for all habitat types. Grasshopper Sparrow (Ammodramus savannarum) showed a strong positive response to pasture and idle grass at all scales and negatively to cropland at large scales. Unlike other species, patch occupancy by Henslow’s Sparrow (A. henslowii) was primarily influenced by patch area.

Conclusions

Our results suggest that both working grasslands (pasture) and idle conservation grasslands can play an important role in grassland bird conservation but also highlight the importance of considering species-specific patch and landscape characteristics for effective conservation.

     

Spatial scale and pattern dependences of aboveground biomass estimation from satellite images: a case study of the Sierra National Forest,California

Context

Spatial scale and pattern play important roles in forest aboveground biomass (AGB) estimation in remote sensing. Changes in the accuracy of satellite images-estimated forest AGBs against spatial scales and pixel distribution patterns has not been evaluated, because it requires ground-truth AGBs of fine resolution over a large extent, and such data are difficult to obtain using traditional ground surveying methods.

Objectives

We intend to quantify the accuracy of AGB estimation from satellite images on changing spatial scales and varying pixel distribution patterns, in a typical mixed coniferous forest in Sierra Nevada mountains, California.

Methods

A forest AGB map of a 143 km² area was created using small-footprint light detection and ranging. Landsat Thematic Mapper images were chosen as typical examples of satellite images, and resampled to successively coarser resolutions. At each spatial scale, pixels forming random, uniform, and clustered spatial patterns were then sampled. The accuracies of the AGB estimation based on Landsat images associated with varying spatial scales and patterns were finally quantified.

Results

The changes in the accuracy of AGB estimation from Landsat images are not monotonic, but increase up to 60–90 m in spatial scale, and then decrease. Random and uniform spatial patterns of pixel distributions yield better accuracy for AGB estimation than clustered spatial patterns. The corrected NDVI (NDVIc) was the best predictor of AGB estimation.

Conclusions

A spatial scale of 60–90 m is recommended for forest AGB estimation at the Sierra Nevada mountains using Landsat images and those with similar spectral resolutions.