Neighbourhood landscape effect on population kinetics of the fossorial water vole (<Emphasis Type="Italic">Arvicola terrestris scherman</Emphasis>)

This paper addresses the issue of whether landscape structure affects A. terrestris population kinetics on a neighbourhood spatial scale, and if so, at what spatial scale is that effect at its maximum. We investigated how the growth of A. terrestris populations is influenced by the landscape context of parcels used for hay production in the French Jura Mountains. Five landscape metrics (relative area of grassland, mean patch area of grassland, patch density of grassland, woodland patch density in grassland, grassland–woodland edge density) were computed over an increasing radius around each parcel (max. 3 km). Redundancy analysis showed that the extent, rate and early onset of A. terrestris population growth were favoured in open grassland areas. Landscape effects on A. terrestris populations as determined by the five metrics are scale-dependent: mean patch area of grassland, patch density of grassland and woodland patch density in grassland had an impact on a grassland parcel within a neighbourhood radius of about 800 m, while relative area of grassland and grassland–woodland edge density had an impact within a neighbourhood radius of about 400 m. Those findings corroborate earlier hypotheses about a multifactorial regulation of A. terrestris populations and a spatial hierarchy of regulating factors. They have potential implications in terms of landscape management and small mammal pest control.     

Post-storm surveys reveal large-scale spatial patterns and influences of site factors,forest structure and diversity in endemic bark-beetle populations

The storm that struck France on december 26^th and 28^th 1999 felled 140 million m³ of timber and had a high economic, social and landscape impact. This event offered the opportunity to study large-scale patterns in populations of forest insect pests that would benefit from the abundant breeding material. A large-scale survey was carried out in France in 2000 to sample the most frequently observed species developing on spruce (Ips typographus, Pityogene schalcographus) and pine (Tomicus piniperda, Ips sexdentatus) in 898 locations distributed throughout wind-damaged areas. The local abundance of each species scored on a 0 to 5 scale was analysed using geostatistical estimators to explore the extent and intensity of spatial autocorrelation, and was related to site, stand, and neighbourhood landscape metrics of the forest cover (in particular the interconnection with broadleaf forest patches) found within dispersal distance. All species but I. sexdentatus, which was much less abundant, displayed large-scale spatial dependence and regional variations in abundance. Lower infestation levels per tree (windfalls and standing trees) were observed in stands with a high proportion of wind-damaged trees, which was interpreted as the result of beetles distributing themselves among the available breeding material. More infestations were observed in wind-broken trees as compared to wind-felled trees. More importantly, populations showed significant relationships with the structure of coniferous stands (in particular with the number of coniferous patches). T. piniperda population levels were negatively correlated to the amount of coniferous edge shared with broadleaf forest patches, possibly because of the disruptive effect of non-host volatiles on host-finding processes at the landscape-scale. The differences observed between species regarding patterns and relationships to site, stand, and forest cover characteristics are discussed in relation to the ecological characteristics of each species.     

Regeneration Strategies,Disturbance and Plant Interactions as Organizers of Vegetation Spatial Patterns in a Pine Forest

To determine how vegetation pattern in early successional forests may be related to plant traits and types of disturbance, we measured percent cover of individual taxa annually in a South Carolina Pinus elliottii forest, starting one year before, and ending four years after harvest and tree girdling disturbances were applied. The 17 most important taxa surveyed were grouped into four regeneration strategies chosen a priori, and the spatial patterns of these groups and of the soil were investigated using global variability, semivariograms and kriged maps. We also examined spatial correlations across years, across taxa, and between species and soil disturbance. Seed bank taxa represented by Dichanthelium spp. increased rapidly and formed large patches, and then quickly declined. Taxa that regenerate by newly dispersed seeds, represented by Rhus copallina and Rubus spp. occurred at first in a few patches, and became widespread later. Stump sprouters, represented by Quercus spp. and Myrica cerifera, had rapid increases in cover, but their spatial patterns were largely determined by their pre-disturbance patterns. Prunus serotina, which relies on both sprouting and dispersed seed, had moderate cover and a random distribution. Within-species temporal correlation of spatial pattern was lower in girdled than in harvested plots, and was not clearly related to regeneration strategy. Forest floor disturbance was patchy and affected the pattern of Dichanthelium spp. in the harvested plots. Negative correlations between herbs and woody plants in harvested plots reflected the role of biotic (i.e., successional) filters on vegetation pattern. Surprisingly, no spatial correlations were detected between the nitrogen fixer, Myrica cerifera and other taxa in this N-limited system. In comparing the spatial and temporal patterns, we found kriged maps more informative than analysis of semivariograms alone. The maps and correlation statistics demonstrated that regeneration traits, spatial patterns of soil disturbances, and interactions among taxa influence dynamics of the spatial patterns of the plants. We also demonstrated that disturbance types affected the importance and interactions among these three factors, and caused different spatial patterns of the plant taxa.     

Assessing Rosa persica genetic diversity using amplified fragment length polymorphisms analysis

The genetic diversity among 128 Iranian Rosa persica (R. persica) accessions in the different populations was analyzed. Amplified fragment length polymorphisms (AFLP) technique was used to produce 171 polymorphic fragments. The number of polymorphic loci ranged from 101 to 147 and the polymorphism information content (PIC) varied from 0.289 to 0.073, with an average of 0.16. This shows extreme variability and genetic diversity among the studied R. persica populations. An indirect estimate of the number of migrants per generation (Nm = 0.376) indicated that gene flow was relatively low among populations of the species. Cluster analysis using the UPGMA method grouped all accessions into six clusters. The results did not show relative agreement with the genotypes’ region of origin. Based on an analysis of molecular variance, 48% of the genetic variation of R. persica was within population and 52% was among populations. The present analysis revealed that Iranian R. persica genotypes are highly variable and genetically distinct from their origins. The apparent unique nature of the R. persica genotypes revealed by our results supports the case for the implementation of more intense characterization and conservation strategies, and provides useful information to address breeding programmes and germplasm resource management in Rosa spp.     

Landscape ecology of <Emphasis Type="Italic">Phragmites australis</Emphasis> invasion in networks of linear wetlands

The interaction between landscape structure and spatial patterns of plant invasion has been little addressed by ecologists despite the new insights it can provide. Because of their spatial configuration as highly connected networks, linear wetlands such as roadside or agricultural ditches, can serve as corridors facilitating invasion at the landscape scale, but species dynamics in these important habitats are not well known. We conducted a landscape scale analysis of Phragmites australis invasion patterns (1985–2002 and 1987–2002) in two periurban areas of southern Québec (Canada) focusing on the interaction between the network of linear wetlands and the adjacent land-uses. Results show that, at the beginning of the reference period, the two landscapes were relatively non-invaded and populations occurred mostly in roadside habitats which then served as invasion foci into other parts of the landscape. The intrinsic rates of increase of P. australis populations in linear anthropogenic habitats were generally higher than those reported for natural wetlands. Riparian habitats along streams and rivers were little invaded compared to anthropogenic linear wetlands, except when they intersected transportation rights-of-way. Bivariate spatial point pattern analysis of colonization events using both Euclidian and network distances generally showed spatial dependence (association) to source populations. An autologistic regression model that included landscape and edaphic variables selected transportation rights-of-way as the best predictor of P. australis occurrence patterns in one of the landscapes. Given the high invasion rates observed, managers of linear wetlands should carefully monitor expansion patterns especially when roads intersect landscapes of conservation or economic value.     

Models based on individual level movement predict spatial patterns of genetic relatedness for two Australian forest birds

Fine-scale landscape change can alter dispersal patterns of animals, thus influencing connectivity or gene flow within a population. Furthermore, dispersal patterns of different species may be influenced by the landscape in varying ways. Our research first aimed to examine whether the spatial genetic structure within populations of closely related bird species differs in response to the same landscape. Second, we examined whether individual-level movement characteristics are a mechanistic driver of these differences. We generated a priori predictions of how landscape features will influence dispersal (particularly the response of individuals to habitat boundaries both natural and human-induced) based on a movement model developed by Fahrig (Funct Ecol 21:1003–1015, 2007). This model allowed us to predict genetic relatedness patterns in populations of two passerine bird species with different life-history traits from Queensland, Australia (yellow-throated scrubwren Sericornis citreogularis, a habitat specialist; white-browed scrubwren Sericornis frontalis, a habitat generalist). We quantified our predictions using cost-distance modelling and compared these to observed pairwise genetic distances (a _r ) between individuals as calculated from microsatellite markers. Mantel tests showed that our a priori models correlated with genetic distance. Euclidean distance was most closely correlated to genetic distance for the generalist species (r = 0.093, P = 0.002), and landscape models that included the avoidance of unsuitable habitat were best for the specialist species (r = 0.107, P = 0.001). Our study showed that predictable movement characteristics may be the mechanism driving differences in genetic relatedness patterns within populations of different bird species.     

Effects of changing spatial scale on the analysis of landscape pattern

The purpose of this study was to observe the effects of changing the grain (the first level of spatial resolution possible with a given data set) and extent (the total area of the study) of landscape data on observed spatial patterns and to identify some general rules for comparing measures obtained at different scales. Simple random maps, maps with contagion (i.e., clusters of the same land cover type), and actual landscape data from USGS land use (LUDA) data maps were used in the analyses. Landscape patterns were compared using indices measuring diversity (H), dominance (D) and contagion (C). Rare land cover types were lost as grain became coarser. This loss could be predicted analytically for random maps with two land cover types, and it was observed in actual landscapes as grain was increased experimentally. However, the rate of loss was influenced by the spatial pattern. Land cover types that were clumped disappeared slowly or were retained with increasing grain, whereas cover types that were dispersed were lost rapidly. The diversity index decreased linearly with increasing grain size, but dominance and contagion did not show a linear relationship. The indices D and C increased with increasing extent, but H exhibited a variable response. The indices were sensitive to the number (m) of cover types observed in the data set and the fraction of the landscape occupied by each cover type (P _k); both m and P _kvaried with grain and extent. Qualitative and quantitative changes in measurements across spatial scales will differ depending on how scale is defined. Characterizing the relationships between ecological measurements and the grain or extent of the data may make it possible to predict or correct for the loss of information with changes in spatial scale.     

Spatio-temporal variation in the demographic attributes of a generalist mesopredator

Human land-use practices have dramatically altered the composition and configuration of native habitats throughout many ecosystems. Within heterogeneous landscapes generalist predators often thrive, causing cascading effects on local biological communities, yet there are few data to suggest how attributes of fragmentation influence local population dynamics of these species. We monitored 25 raccoon (Procyon lotor) populations from 2004 to 2009 in a fragmented agricultural landscape to evaluate the influence of local and landscape habitat attributes on spatial and temporal variation in demography. Our results indicate that agricultural ecosystems support increased densities of raccoons relative to many other rural landscapes, but that spatial and temporal variation in demography exists that is driven by non-agricultural habitat attributes rather than the availability of crops. At the landscape scale, both density and population stability were positively associated with the size and contiguity of forest patches, while at the local scale density was positively correlated with plant diversity and the density of tree cavities. In addition, populations occupying forest patches with greater levels of plant diversity and stable water resources exhibited less temporal variability than populations with limited plant species complexity or water availability. The proportion of populations comprised of females was most strongly influenced by the availability of tree cavities and soft mast. Despite the abundance of mesopredators in heterogeneous landscapes, our results indicate that all patches do not contribute equally to the regional abundance and persistence of these species. Thus, a clear understanding of how landscape attributes contribute to variation in demography is critical to the optimization of management strategies.     

Dispersion of kangaroo rat mounds at multiple scales in New Mexico,USA

Burrowing mammals create disturbances that increase the ecological heterogeneity of landscapes. In desert systems, banner-tailed kangaroo rats (Dipodomys spectabilis) construct large mounds that greatly influence the spatial patterning of soils, plants, and animals. The overall effects of the patches generated by D. spectabilis depend on the dispersion patterns of the mounds; these patterns may be sensitive to scale and landscape position. We examined the distribution of D. spectabilis mounds across multiple scales in four 40-ha grassland plots in New Mexico, USA. We used Ripley's K-function for our point-pattern analysis. The dispersion patterns of mounds were generally scale-sensitive but depended somewhat on plot-level densities, which were related to topographic position and grazing history. Mound spacing was either regular or random at small scales (0–50 m), random or aggregated at intermediate scales (50–300 m), and aggregated at large scales (300–3000 m). This scale-dependency of pattern reflected spatial domains in which different biotic (territoriality, dispersal, grazing) and abiotic (soil texture and drainage) factors exerted strong influences. How other organisms perceive the spatial patterning of mounds will depend on the extent of their movements. Patches may appear regular to one species but aggregated to another. The dispersion of D. spectabilis mounds also has implications for the spatial structuring of desert rodent communities. D. spectabilis excludes smaller species of kangaroo rats from areas around their mounds; they create spatial heterogeneity in behavioral dominance that may influence the distribution of subordinate species at multiple scales.     

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.     

Landscape ecology: Population genetics at the metapopulation level

Distribution of genetic diversity in a landscape depends on both within and among population processes. Selective pressures within populations have traditionally been studied by population genetics, which usually assumes that populations are at equilibrium. However, when selection pressures within and among populations are different, landscape processes are required to define an equilibrium (landscape being defined as the habitat of a set of populations called a metapopulation, and populations will differ depending on their situation in the landscape, i.e. their age and the state of neighboring populations). We examine reproduction systems and life history traits, for which variation depends on landscape processes. Predictions of their states in a metapopulation are drawn from theoretical models, and confronted to observations collected in natural populations.     

The modifiable areal unit problem of spatial heterogeneity of plant community in the transitional zone between oasis and desert using semivariance analysis

The modifiable areal unit problem has significant implications for ecological research that involve investigating and analyzing the spatial heterogeneity of plant community. In this paper, semivariance analysis was used to evaluate the spatial characteristics of plant community in the␣transitional zone between oasis and desert. The spatial structures of the plant community were characterized using exponent model variogram parameters, including nugget (C ₀) range (A ₀) and sill (C ₀+C). Two methods were employed to determine the scale effect of spatial heterogeneity. (1) A constant grain size (10×10 m²) and variational plot areas have been used to analyze spatial heterogeneity of the plant community. (2) The grain size was only changed to analyze spatial heterogeneity. In addition, the plot of 500×500 m² was clustered into nested units of different shapes and different directions to analyze zoning effect. Finally, using semivariance analysis, we obtained a suitable plot area and zoning approach to weaken the scale and zoning effects. The results showed that the effects of scale on different variogram parameters had significant difference. For example, C ₀ and C ₀+C were very sensitive at small scales, A ₀ was influenced significantly by plot area at larger scales, and C ₀ and A ₀ were relatively sensitive to different zoning approaches. In order to get more representative characteristic of spatial heterogeneity of the plant community, the sampling area should be more than 200×200 m² for Nitraria sphaerocarpa populations, 100×100 m² for Reaumuria soongorica populations, and a grain size from 20×20 m² to 30×30 m² for both populations.     

Influence of fire regimes on lodgepole pine stand age and density across the Yellowstone National Park (USA) landscape

A probabilistic spatial model was created based on empirical data to examine the influence of different fire regimes on stand structure of lodgepole pine (Pinus contorta var. latifolia) forests across a >500,000-ha landscape in Yellowstone National Park, Wyoming, USA. We asked how variation in the frequency of large fire events affects (1) the mean and annual variability of age and tree density (defined by postfire sapling density and subsequent stand density) of lodgepole pine stands and (2) the spatial pattern of stand age and density across the landscape. The model incorporates spatial and temporal variation in fire and serotiny in predicting postfire sapling densities of lodgepole pine. Empirical self-thinning and in-filling curves alter initital postfire sapling densities over decades to centuries. In response to a six-fold increase in the probability of large fires (0.003 to 0.018 year⁻¹), mean stand age declined from 291 to 121 years. Mean stand density did not increase appreciably at high elevations (1,029 to 1,249 stems ha⁻¹) where serotiny was low and postfire sapling density was relatively low (1,252 to 2,203 stems ha⁻¹). At low elevations, where prefire serotiny and postfire lodgepole pine density are high, mean stand densities increased from 2,807 to 7,664 stems ha⁻¹. Spatially, the patterns of stand age became more simplified across the landscape, yet patterns of stand density became more complex. In response to more frequent stand replacing fires, very high annual variability in postfire sapling density is expected, with higher means and greater variation in stand density across lodgepole pine landscapes, especially in the few decades following large fires.     

Corridors and connectivity: when use and function do not equate

Connectivity, or the integration of populations into a single demographic unit, is an often desired, but largely untested aspect of wildlife corridors. Using a corridor system that was established at least 85 years prior, we investigated the extent of connectivity provided. This was undertaken using a combined ecological and genetic approach with connectivity estimated by gene flow. Vegetation within the corridor was found to be comparable in physical structure and species composition to that within the connected patches and the two target species (Melomys cervinipes and Uromys caudimaculatus) were shown to occur along the corridor but not within the surrounding matrix. These factors indicated that the corridor was suitable for use as a model system. The population structure (weights of individuals, sex ratios and the percentage of juveniles) of both species were also similar within the corridor and the connected patches suggesting that the corridor provided the resources necessary to sustain breeding populations along its length. Despite this, populations in patches linked by the corridor were found to show the same significant levels of genetic differentiation as those in isolated habitats. M. cervinipes, but not U. caudimaculatus, also showed population differentiation within the continuous habitat. Although based on only one corridor system, these results clearly demonstrate that connectivity between connected populations will not always be achieved by the construction or retention of a corridor and that connectivity cannot be inferred solely from the presence of individuals, or breeding populations, within the corridor. C. Wilson: deceased     

Spatial analysis of roadside Acacia populations on a road network using the network K-function

Spatial patterning of plant distributions has long been recognised as being important in understanding underlying ecological processes. Ripley’s K-function is a frequently used method for studying the spatial pattern of mapped point data in ecology. However, application of this method to point patterns on road networks is inappropriate, as the K-function assumes an infinite homogenous environment in calculating Euclidean distances. A new technique for analysing the distribution of points on a network has been developed, called the network K-function (for univariate analysis) and network cross K-function (for bivariate analysis). To investigate its applicability for ecological data-sets, this method was applied to point location data for roadside populations of three Acacia species in a fragmented agricultural landscape of south-eastern Australia. Kernel estimations of the observed density of spatial point patterns for each species showed strong spatial heterogeneity. Combined univariate and bivariate network K-function analyses confirmed significant clustering of populations at various scales, and spatial patterns of Acacia decora suggests that roadworks activities may have a stronger controlling influence than environmental determinants on population dynamics. The network K-function method will become a useful statistical tool for the analyses of ecological data along roads, field margins, streams and other networks.This revised version was published online in May 2005 with corrections to the Cover Date.     

Disentangling habitat and social drivers of nesting patterns in songbirds

Nest locations of breeding birds are often spatially clustered. This tendency to nest together has generally been related to a patchy distribution of nesting habitat in landscape studies, but behavioral studies of species with clustered breeding patterns draw attention to the importance of social and biotic factors. Indeed, it is becoming increasingly apparent that the breeding system of many territorial, migrant birds may be semi-colonial. The reasons for, and extent of, spatial clustering in their breeding systems are not well understood. Our goal was to tease apart the influence of habitat availability and social drivers of clustered breeding in a neotropical migrant species, the hooded warbler (Wilsonia citrina). To test alternative hypotheses related to clustered habitat or conspecific attraction, we combined a habitat classification based on remote sensing with point pattern analysis of nesting sites. Nest locations (n = 150, 1999–2004), collected in a 1213 ha forested area of Southern Ontario (Canada), were analyzed at multiple spatial scales. Ripley’s K and pair-correlation functions g (uni- and bivariate) were used to test whether nests were clustered merely because potential nesting habitat was also clustered, or whether nests were additionally clustered with respect to conspecifics. Nest locations tended to be significantly clustered at intermediate distances (particularly between 240 and 420 m). Nests were randomly distributed within available habitat at larger distance scales, up to 1500 m. A reasonable hypothesis to explain the detected additional clustering, and one that is consistent with the results of several behavioral studies, is that females pack their nests more tightly than the available habitat requires to be situated closer to their neighbors’ mates. Linking spatially explicit, point pattern analysis with strong inference based on Monte Carlo tests may bring us closer to understanding the generality and reasons behind conspecific attraction at different spatial scales. F. Csillag—deceased.     

Genetic structure of mulberry from different ecotypes revealed by ISSRs in China: An implications for conservation of local mulberry varieties

Mulberry is a perennial and economically important plant that has traditionally been used for feeding the silkworm. Evaluating genetic relationship is important for long-term improvement in mulberry yield, quality and resistance, and for germplasm conservation and identification. Population structure and genetic diversity of 8 mulberry populations from different ecotypes in China were analyzed by ISSR markers. Twelve ISSR primers generated a total of 83 amplification products, of which 50 were polymorphic, revealing 60.24% polymorphism among 66 mulberry local varieties, the mean PIC value was 0.1469. The total heterozygosity (H_T), heterozygosity within population (H_S), diversity between populations (D_ST) were 0.1600, 0.0851 and 0.0749, respectively. The coefficient of population differentiation (G_ST) was 0.4683, indicating that the variations among populations and those within populations contributed 46.8% and 53.2% to the total heterozygosity, respectively. The gene flow (N_m) was 0.5678, suggesting that genetic drift between populations can caused local genetic differentiation and therefore, population divergence. The mean genetic similarity coefficient was 0.8456, genetic similarity coefficient among 8 mulberry populations ranged from 0.8441 to 0.9640, indicating that genetic diversity of different populations existed variation. A dendrogram of all 66 local varieties of mulberry based on the genetic similarity using ISSR markers was generated by UPGMA cluster method. In the dendrogram, most varieties from the same ecotype clustered together.     

Disturbance increases negative spatial autocorrelation in species diversity

Context

Ecological processes that shape diversity and spatial pattern of ecological communities are often altered by disturbance. Spatial patterns (spatial autocorrelation) in species diversity are thus expected to change with disturbance.

Objective

When examining spatial patterns, ecologists traditionally lump positive and negative spatial autocorrelation into the overall spatial autocorrelation. By contrast, here we aim to understand disturbance effects on both positive and negative spatial autocorrelation of species richness and evenness, which may be related to environmental filtering and restricted dispersal, and to competition, respectively.

Methods

For 8 years, we monitored the spatial autocorrelation in species richness and evenness of riparian plant communities in both uncut control and experimentally clearcut sites in the boreal forest of Alberta, Canada. The overall spatial autocorrelation for each of these two indices of diversity was separately decomposed into the components of positive and negative spatial autocorrelations through eigendecomposition of the spatial weighting matrix.

Results

Negative spatial autocorrelation in richness and evenness were more pronounced in the clearcut than uncut sites, although positive spatial autocorrelations in all indices of diversity remained unchanged. Effect of disturbance was not detected on the overall spatial autocorrelation.

Conclusions

Disturbance increases negative spatial autocorrelation in species richness and evenness, with a stronger increase in evenness than richness, which underscores the importance of competition in structuring post-disturbance riparian communities. Our results also highlight the need for assessing positive and negative spatial autocorrelation and different aspects of diversity separately in understanding disturbance effects on the spatial pattern, or identifying processes from patterns.

     

An aggregation index (AI) to quantify spatial patterns of landscapes

There is often need to measure aggregation levels of spatial patterns within a single map class in landscape ecological studies. The contagion index (CI), shape index (SI), and probability of adjacency of the same class (Qi), all have certain limits when measuring aggregation of spatial patterns. We have developed an aggregation index (AI) that is class specific and independent of landscape composition. AI assumes that a class with the highest level of aggregation (AI =1) is comprised of pixels sharing the most possible edges. A class whose pixels share no edges (completely disaggregated) has the lowest level of aggregation (AI =0). AI is similar to SI and Qi, but it calculates aggregation more precisely than the latter two. We have evaluated the performance of AI under varied levels of (1) aggregation, (2) number of patches, (3) spatial resolutions, and (4) real species distribution maps at various spatial scales. AI was able to produce reasonable results under all these circumstances. Since it is class specific, it is more precise than CI, which measures overall landscape aggregation. Thus, AI provides a quantitative basis to correlate the spatial pattern of a class with a specific process. Since AI is a ratio variable, map units do not affect the calculation. It can be compared between classes from the same or different landscapes, or even the same classes from the same landscape under different resolutions.