A wavelet‐based approach to evaluate the roles of structural and functional landscape heterogeneity in animal space use at multiple scales

The functional relationship between habitat utilization and landscape spatial heterogeneity is fundamental to understanding the spatial nature of animal distribution across scales. Although structural and functional properties of landscape spatial heterogeneity can have different consequences for animal species, few studies have explicitly considered both forms of heterogeneity, partly due to the lack of general methods for direct assessment of scale‐specific associations between variables. We present a wavelet‐based approach to evaluate the roles of structural and functional landscape spatial heterogeneity in animal space use at multiple spatial scales. As a case study, we examined scale‐specific space use patterns of American black bears Ursus americanus in response to structural and functional spatial heterogeneity as well as spatial patterns of vegetation age‐classes in a Canadian boreal forest. We found strong differences in the effects of structural and functional spatial heterogeneity and the scales at which they are associated with the patterns of habitat use by black bears. Functional heterogeneity alone affected space use at 800 and 1600‐m scales, but had significant effects when interacting with structural heterogeneity at 400, 800, and 1600‐m scales. Compared with male bears, female black bears were most sensitive to patterns of forage abundance at intermediate scales, or more specifically, in young and regenerating forests that provide abundant soft mast in boreal forests. Our study highlights the importance of accounting for scale‐dependent properties of (structural and functional) spatial heterogeneity in assessing the ecological effects of landscape components and the effectiveness of the wavelet transform technique in identifying such scale‐specific relationships.     

Linking the spatial scale of environmental variation and the evolution of phenotypic plasticity: selection favors adaptive plasticity in fine-grained environments

Adaptive phenotypic plasticity and adaptive genetic differentiation enable plant lineages to maximize their fitness in response to environmental heterogeneity. The spatial scale of environmental variation relative to the average dispersal distance of a species determines whether selection will favor plasticity, local adaptation, or an intermediate strategy. Habitats where the spatial scale of environmental variation is less than the dispersal distance of a species are fine grained and should favor the expression of adaptive plasticity, while coarse-grained habitats, where environmental variation occurs on spatial scales greater than dispersal, should favor adaptive genetic differentiation. However, there is relatively little information available characterizing the link between the spatial scale of environmental variation and patterns of selection on plasticity measured in the field. I examined patterns of spatial environmental variation within a serpentine mosaic grassland and selection on an annual plant (Erodium cicutarium) within that landscape. Results indicate that serpentine soil patches are a significantly finer-grained habitat than non-serpentine patches. Additionally, selection generally favored increased plasticity on serpentine soils and diminished plasticity on non-serpentine soils. This is the first empirical example of differential selection for phenotypic plasticity in the field as a result of strong differences in the grain of environmental heterogeneity within habitats.     

Multivariate analysis of a fine-scale breeding bird atlas using a geographical information system and partial canonical correspondence analysis: environmental and spatial effects

Aim To assess the relative roles of environment and space in driving bird species distribution and to identify relevant drivers of bird assemblage composition, in the case of a fine‐scale bird atlas data set. Location The study was carried out in southern Belgium using grid cells of 1 × 1 km, based on the distribution maps of the Oiseaux nicheurs de Famenne: Atlas de Lesse et Lomme which contains abundance for 103 bird species. Methods Species found in < 10% or > 90% of the atlas cells were omitted from the bird data set for the analysis. Each cell was characterized by 59 landscape metrics, quantifying its composition and spatial patterns, using a Geographical Information System. Partial canonical correspondence analysis was used to partition the variance of bird species matrix into independent components: (a) ‘pure’ environmental variation, (b) spatially‐structured environmental variation, (c) ‘pure’ spatial variation and (d) unexplained, non‐spatial variation. Results The variance partitioning method shows that the selected landscape metrics explain 27.5% of the variation, whilst ‘pure’ spatial and spatially‐structured environmental variables explain only a weak percentage of the variation in the bird species matrix (2.5% and 4%, respectively). Avian community composition is primarily related to the degree of urbanization and the amount and composition of forested and open areas. These variables explain more than half of the variation for three species and over one‐third of the variation for 12 species. Main conclusions The results seem to indicate that the majority of explained variation in species assemblages is attributable to local environmental factors. At such a fine spatial resolution, however, the method does not seem to be appropriated for detecting and extracting the spatial variation of assemblages. Consequently, the large amount of unexplained variation is probably because of missing spatial structures and ‘noise’ in species abundance data. Furthermore, it is possible that other relevant environmental factors, that were not taken into account in this study and which may operate at different spatial scales, can drive bird assemblage structure. As a large proportion of ecological variation can be shared by environment and space, the applied partitioning method was found to be useful when analysing multispecific atlas data, but it needs improvement to factor out all‐scale spatial components of this variation (the source of ‘false correlation’) and to bring out the ‘pure’ environmental variation for ecological interpretation.     

Multiple scales and the relationship between density and spatial aggregation in littoral zone communities

Understanding the constraints on community composition at multiple spatial scales is an immense challenge to community and ecosystem ecologists. As community composition is basically the composite result of species’ spatial patterning, studying this spatial patterning across scales may yield clues as to which scales of environmental heterogeneity influence communities. The now widely documented positive interspecific relationship between ‘regional’ range and mean ‘local’ abundance has become a generalisation describing the spatial patterning of species at coarse scales. We address some of the shortcomings of this generalisation, as well as examine the cross‐scale spatial patterning (aggregation and density levels) of littoral‐benthic invertebrates in very large lakes. Specifically, we (a) determine whether the positive range‐abundance relationship can be reinterpreted in terms of the actual spatial structure of species distributions, (b) examine the relationship between aggregation and density across different spatial scales, and (c) determine whether the spatial patterning of species (e.g. low density/aggregated distribution) is constant across scales, that is, whether our interpretation of a species spatial pattern is dependent on the scale at which we choose to observe the system. Spatial aggregation of littoral invertebrates was generally a negative function of mean density across all spatial scales and seasons (autumn and spring). This relationship may underlie positive range‐abundance relationships. Species that were uncommon and highly aggregated at coarse spatial scales can be abundant and approach random distributions at finer spatial scales. Also, the change in spatial aggregation of closely related taxa across spatial scales was idiosyncratic. The idiosyncratic cross‐scale spatial patterning of species implies that multiple scales of environmental heterogeneity may influence the assembly of littoral communities. Due to the multi‐scale, species‐specific spatial patterning of invertebrates, littoral zone communities form a complex spatial mosaic, and a ‘spatially explicit’ approach will be required by limnologists in order to link littoral‐benthic community patterns with ecosystem processes in large oligotrophic lakes.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Multilevel landscape utilization of the Siberian flying squirrel: Scale effects on species habitat use

Animals use and select habitat at multiple hierarchical levels and at different spatial scales within each level. Still, there is little knowledge on the scale effects at different spatial levels of species occupancy patterns. The objective of this study was to examine nonlinear effects and optimal‐scale landscape characteristics that affect occupancy of the Siberian flying squirrel, Pteromys volans, in South‐ and Mid‐Finland. We used presence–absence data (n = 10,032 plots of 9 ha) and novel approach to separate the effects on site‐, landscape‐, and regional‐level occupancy patterns. Our main results were: landscape variables predicted the placement of population patches at least twice as well as they predicted the occupancy of particular sites; the clear optimal value of preferred habitat cover for species landscape‐level abundance is a surprisingly low value (10% within a 4 km buffer); landscape metrics exert different effects on species occupancy and abundance in high versus low population density regions of our study area. We conclude that knowledge of regional variation in landscape utilization will be essential for successful conservation of the species. The results also support the view that large‐scale landscape variables have high predictive power in explaining species abundance. Our study demonstrates the complex response of species occurrence at different levels of population configuration on landscape structure. The study also highlights the need for data in large spatial scale to increase the precision of biodiversity mapping and prediction of future trends.     

Do species population parameters and landscape characteristics affect the relationship between local population abundance and surrounding habitat amount?

In landscape ecology, correlational approaches are typically used to analyse links between local population abundance, and the surrounding habitat amount to estimate biologically-relevant landscape size (extent) for managing endangered or pest populations. The direction, strength, and spatial extent of the correlations are then sometimes interpreted in terms of species population parameters. Here we simulated the population dynamics of generalized species across spatially explicit landscapes that included two distinct habitat types. We investigated how characteristics of a landscape (structure), including the variation in habitat quality and spatial aggregation of the habitat, and the precise population-dynamic properties of the simulated species (dispersal and growth rates) affect the correlation between population abundance and amount of surrounding favourable habitat in the landscape. To evaluate these spatial extents of correlation, proportions of favourable habitat were calculated within several circles of increasing diameter centred on sampling patches of favourable habitat where population abundance was recorded.We found that the value of the correlation coefficients between population abundance and amount of surrounding favourable habitat depended on both population dynamic parameters and landscape characteristics. Coefficients of correlation increased with the variation in habitat quality and the aggregation of favourable habitat in the landscape, but decreased with the dispersal distance. The distance at which the correlation was maximized was sensitive to an interaction between the level of aggregation of the habitat and the dispersal distance; whereas the greatest distance at which a significant correlation occurred was more sensitive to the variation in habitat quality. Our results corroborate the view that correlational analyses do provide information on the local population dynamics of a species in a given habitat type and on its dispersal rate parameters. However, even in simplified, model frameworks, direct relationships are often difficult to disentangle and global landscape characteristics should be reported in any studies intended to derive population-dynamic parameters from correlations. Where possible, replicated landscapes should be examined in order to control for the interaction between population dynamics and landscape structure. Finally, we recommend using species-specific, population-dynamic modelling in order to interpret correctly the observed patterns of correlation in the landscape.     

Scale‐Dependent Spatial Match between Fruits and Fruit‐eating Birds during the Breeding Season in Yungas Andean Forests

The multi‐scale spatial match between bird and food abundances is a main driver of the structure of fruit‐eating bird assemblages. We explored how the activity of fruit‐eating birds was influenced by the abundance of fruits at the local and landscape scales in Andean mountain forests during the breeding season, when most birds forage close to their nest. We measured: (1) the spatial scale of variation in the abundance of fruits, (2) the spatial scale of variation in the activity of fruit‐eating birds, and (3) the spatial match between both variables. The sampling design consisted of eleven 1.2‐ha sites, each subdivided into 30 cells of 20 × 20 m, where we sampled fruits and fruit‐eating birds. We found that fruit consumption, and to a lesser extent bird abundance, were associated with local spatial variation in abundance of selected fruit species. However, fruit‐eating birds did not modify their spatial distribution in the landscape following changes in availability of these fruits. Our study shows that fruit‐eating birds detect local spatial variation in fruit availability in their home breeding ranges, and exploit patches with large clusters of selected fruits. However, it may be unprofitable for breeding birds to stray too far from their nests to exploit fruit‐rich patches, accounting for the absence of fruit tracking at larger spatial scales.     

Species distribution modelling in low‐interaction environments: Insights from a terrestrial Antarctic system

It is widely acknowledged that in the terrestrial Antarctic, interspecific interactions are typically unimportant in determining species distributions and community structure. Therefore, correlative models should prove useful for predicting current and future spatial variation in species abundance patterns. However, this idea has not been formally tested, and the utility of such models, which have shown value for understanding the distribution of diversity elsewhere, for investigating biodiversity patterns in Antarctica remains unclear. Here we make a start at such tests by using generalized linear and simultaneous autoregressive models to demonstrate that simple environmental variables and information about the spatial structure of the environment can explain more than 90% of the variation in the abundance of Maudheimia wilsoni (Oribatida; Maudheimiidae), a representative of one of the most significant groups of Antarctic terrestrial arthropods, the mites. We show that a single environmental variable, maximum soil moisture content, can account for as much as 80% of the variance in the abundance of the mite, and that linear models with only a few environmental and spatial terms can be used to forecast the species abundance at the landscape scale. Given ongoing calls for better understanding of the distribution of Antarctic diversity and its likely future change, this initial test indicates that such modelling procedures, and more sophisticated versions thereof, hold much promise for the region and should be tested for other taxa with different life forms and habitat requirements.     

Environmental correlates of mammal species richness in South America: effects of spatial structure, taxonomy and geographic range

Although some consensus exists regarding the positive synergism between energy and heterogeneity in increasing species diversity, the role of environmental variability remains controversial. We examine how these factors interact to explain spatial variation in mammal species richness in South America. After taking into account the effects of spatial autocorrelation and area, elevation variability and energy mainly drive spatial variation in mammal species richness. The effect of environmental variability is less important. When different taxonomic groups of mammals are analyzed separately, three ways emerge whereby energy and heterogeneity interact to promote species richness. Heterogeneity may have no effect on species richness, habitat heterogeneity and energy availability contribute independently to species richness, or heterogeneity increases in importance with an increase in energy availability. The partition of species into range size quartiles shows that habitat heterogeneity and temporal instability in the resource supply account for the species richness pattern in the narrowest- ranging species. Habitat heterogeneity is significant also for intermediate ranging species but not for the widest-ranging species. Energy alone drives the species richness pattern in the latter species. The interplay between ecology and biogeographic history may ultimately explain these differences given that narrow- and wide-ranging species show distinct biogeographic patterns, and different taxonomic groups also unequally represent them.     

From Hutchinsonian ratios to spatial scaling theory: the interplay among limiting similarity,body size and landscape structure

Spatial scaling theory (SST) relates the physical structure of the environment to species coexistence and community assembly. Although SST is a recognized theory in ecology, few studies have evaluated its predictions, producing contradictory results and frequently failing to meet its assumptions. In addition, the ‘risk predictions’ of SST regarding an increase in species similarity with body size and the dependence of this pattern on the landscape and food fractal dimensions have not been evaluated. This study attempted to account for previous limitations, analyzing these predictions in coleopteran guilds that inhabit 18 temporary ponds. This metacommunity covers a large gradient of environmental variables, including food density, the landscape fractal dimension, the food fractal dimensions and other indicators of pond heterogeneity. Average similarity in carnivorous and herbivorous body sizes systematically increased with guild richness, fulfilling classical predictions of niche theory. Species similarity was associated with body size, but the association reverts from negative to positive as the landscape fractal dimension and heterogeneity increases, a pattern further supported by null model analyses. Several nonexclusive mechanisms may account for this pattern: 1) the body size-dependent landscape perception, through which small animals detect more heterogeneity than larger animals; 2) the reaching of landscape limits by larger species, which prevents them from accessing novel largest clusters; 3) the large differences between the landscape and food fractal dimensions; and 4) the homogenization of the landscape when an integer fractal dimension is reached. These mechanisms may dictate that smaller organisms are more able to capitalize on heterogeneity or available resources than larger organisms, thus promoting increased similarity among smaller species. The presented results support the connection between landscape spatial structure and biodiversity and a mechanistic understanding of this connection from the SST.     

Relative importance of water, energy, and heterogeneity in determining regional pteridophyte and seed plant richness in China

Environmental variables, such as ambient energy, water availability, and environmental heterogeneity have been frequently proposed to account for species diversity gradients. How taxon-specific functional traits define large-scale richness gradients is a fundamental issue in understanding spatial patterns of species diversity, but has not been well documented. Using a large dataset on the regional flora from China, we examine the contrast spatial patterns and environmental determinants between pteridophytes and seed plants which differ in dispersal capacity and environmental requirements. Pteridophyte richness shows more pronounced spatial variation and stronger environmental associations than seed plant richness. Water availability generally accounts for more spatial variance in species richness of pteridophytes and seed plants than energy and heterogeneity do, especially for pteridophytes which have high dependence on moist and shady environments. Thus, pteridophyte richness is disproportionally affected by water-related variables; this in turn results in a higher proportion of pteridophytes in regional vascular plant floras (pteridophyte proportion) in wet regions. Most of the variance in seed plant richness, pteridophyte richness, and pteridophyte proportion explained by energy is included in variation that water and heterogeneity account for, indicating the redundancy of energy in the study extent. However, heterogeneity is more important for determining seed plant distributions. Pteridophyte and seed plant richness is strongly correlated, even after the environmental effects have been removed, implying functional linkages between them. Our study highlights the importance of incorporating biological traits of different taxonomic groups into the studies of macroecology and global change biology.     

Influence of clay licks on the diversity and structure of an Amazonian forest

The spatial heterogeneity of resource availability is a major driver of biodiversity patterns. Some environmental conditions and resources are characterized by large‐scale patterns of variation within the landscape. Clumped local discontinuities or discrete elements also increase spatial heterogeneity, promoting local ‘biodiversity hot spots’ by modifying habitat characteristics and promoting plant–animal interactions. Clay licks are faunal attractors owing to their role in the nutritional ecology of the user species; nevertheless, the effect of their presence on the surrounding vegetation has been poorly quantified. Here, we use data from 100 × 10 m transects and evaluate the effects of the presence of clay licks on forest diversity and structure at local and landscape scales. In clay lick areas, there was a higher abundance of certain species, which helps to homogenize species composition between localities counteracting the natural distance‐decay of compositional similarity between transects without clay lick influence (controls). Compared to control sites, clay lick′s forests had higher palm densities, shorter but more variable individuals in the canopy and understory, a thinner canopy layer, and denser herbaceous and ground level covers. These differences were found along the whole length of transects in both sampled areas types. These results reveal that the presence of discrete elements (i.e., clay licks) may help to explain the compositional and structural heterogeneity of Amazonian forests influencing ecological processes such as seed dispersal and trampling. These considerations may be relevant for other biomes where clay licks are present and give weight to their inclusion in conservation initiatives in tropical forests.     

Temporal Beta Diversity of Bird Assemblages in Agricultural Landscapes: Land Cover Change vs. Stochastic Processes

Temporal variation in the composition of species assemblages could be the result of deterministic processes driven by environmental change and/or stochastic processes of colonization and local extinction. Here, we analyzed the relative roles of deterministic and stochastic processes on bird assemblages in an agricultural landscape of southwestern France. We first assessed the impact of land cover change that occurred between 1982 and 2007 on (i) the species composition (presence/absence) of bird assemblages and (ii) the spatial pattern of taxonomic beta diversity. We also compared the observed temporal change of bird assemblages with a null model accounting for the effect of stochastic dynamics on temporal beta diversity. Temporal assemblage dissimilarity was partitioned into two separate components, accounting for the replacement of species (i.e. turnover) and for the nested species losses (or gains) from one time to the other (i.e. nestedness-resultant dissimilarity), respectively. Neither the turnover nor the nestedness-resultant components of temporal variation were accurately explained by any of the measured variables accounting for land cover change (r²<0.06 in all cases). Additionally, the amount of spatial assemblage heterogeneity in the region did not significantly change between 1982 and 2007, and site-specific observed temporal dissimilarities were larger than null expectations in only 1% of sites for temporal turnover and 13% of sites for nestedness-resultant dissimilarity. Taken together, our results suggest that land cover change in this agricultural landscape had little impact on temporal beta diversity of bird assemblages. Although other unmeasured deterministic process could be driving the observed patterns, it is also possible that the observed changes in presence/absence species composition of local bird assemblages might be the consequence of stochastic processes in which species populations appeared and disappeared from specific localities in a random-like way. Our results might be case-specific, but if stochastic dynamics are generally dominant, the ability of correlative and mechanistic models to predict land cover change effects on species composition would be compromised.     

What's your move? Movement as a link between personality and spatial dynamics in animal populations

Recent studies have established the ecological and evolutionary importance of animal personalities. Individual differences in movement and space‐use, fundamental to many personality traits (e.g. activity, boldness and exploratory behaviour) have been documented across many species and contexts, for instance personality‐dependent dispersal syndromes. Yet, insights from the concurrently developing movement ecology paradigm are rarely considered and recent evidence for other personality‐dependent movements and space‐use lack a general unifying framework. We propose a conceptual framework for personality‐dependent spatial ecology. We link expectations derived from the movement ecology paradigm with behavioural reaction‐norms to offer specific predictions on the interactions between environmental factors, such as resource distribution or landscape structure, and intrinsic behavioural variation. We consider how environmental heterogeneity and individual consistency in movements that carry‐over across spatial scales can lead to personality‐dependent: (1) foraging search performance; (2) habitat preference; (3) home range utilization patterns; (4) social network structure and (5) emergence of assortative population structure with spatial clusters of personalities. We support our conceptual model with spatially explicit simulations of behavioural variation in space‐use, demonstrating the emergence of complex population‐level patterns from differences in simple individual‐level behaviours. Consideration of consistent individual variation in space‐use will facilitate mechanistic understanding of processes that drive social, spatial, ecological and evolutionary dynamics in heterogeneous environments.     

Importance of estimating matrix quality for modeling species distribution in complex tropical landscapes: a test with Atlantic forest small mammals

Information to guide decision making is especially urgent in human dominated landscapes in the tropics, where urban and agricultural frontiers are still expanding in an unplanned manner. Nevertheless, most studies that have investigated the influence of landscape structure on species distribution have not considered the heterogeneity of altered habitats of the matrix, which is usually high in human dominated landscapes. Using the distribution of small mammals in forest remnants and in the four main altered habitats in an Atlantic forest landscape, we investigated 1) how explanatory power of models describing species distribution in forest remnants varies between landscape structure variables that do or do not incorporate matrix quality and 2) the importance of spatial scale for analyzing the influence of landscape structure. We used standardized sampling in remnants and altered habitats to generate two indices of habitat quality, corresponding to the abundance and to the occurrence of small mammals. For each remnant, we calculated habitat quantity and connectivity in different spatial scales, considering or not the quality of surrounding habitats. The incorporation of matrix quality increased model explanatory power across all spatial scales for half the species that occurred in the matrix, but only when taking into account the distance between habitat patches (connectivity). These connectivity models were also less affected by spatial scale than habitat quantity models. The few consistent responses to the variation in spatial scales indicate that despite their small size, small mammals perceive landscape features at large spatial scales. Matrix quality index corresponding to species occurrence presented a better or similar performance compared to that of species abundance. Results indicate the importance of the matrix for the dynamics of fragmented landscapes and suggest that relatively simple indices can improve our understanding of species distribution, and could be applied in modeling, monitoring and managing complex tropical landscapes.     

Exogenous and endogenous determinants of spatial aggregation patterns in Tibetan Plateau meadow vegetation

Aims We aim to quantify the relative importance of various endogenous and exogenous processes influencing the spatial distribution of the individuals of plant species at different temporal and spatial scales in a species-rich and high-cover meadow in the eastern Tibetan Plateau.Methods We calculated Green's index of dispersion to infer the spatial distribution patterns of 73 herbaceous species at two scales (0.25 and 1.0 m 2). We constructed a series of generalized linear models to test the hypotheses that different species traits such as mean plant stem density, per capita dry biomass, maximum plant height and mean seed mass contribute to their spatial distribution. We used the first principal component of soil C, N and P to explain abundance variation across quadrats and sub-plots.Important findings The individuals of the species studied were highly spatially aggregated. At both spatial scales, biomass and stem density explained the most variation in aggregation, but there was no evidence for an effect of mean seed mass on aggregation intensity. The effects of soil carbon, nitrogen and phosphorus at different depths affected plant abundance mostly at the broader spatial scale. Our results demonstrate that self-thinning and habitat heterogeneity all contribute to determine the spatial aggregation patterns of plant individuals in alpine meadow vegetation in the eastern Tibetan Plateau.     

Spatial variation in insect community and species responses to habitat loss and plant community composition

Several experimental studies have examined species responses to manipulations of habitat area and spatial arrangement, but plant composition and spatial variation in species distributions also affect animal responses to habitat alteration. We used an experimental approach to study the combined effects of habitat area, edge, and plant community composition on the spatial structure of insect species richness and composition. The abundance of three guilds (herbivores, predators and parasitoids) and individual species were also analyzed. Habitat patches were created that differed in area and edge by selectively mowing portions of 15 m×15 m plots in a 1.7-ha old field. Spatial and environmental variables were used to predict insect responses in separate multiple regression and ordination models. The variation in species responses due to spatial and environmental variables was then partitioned by combining these variables into an overall regression or ordination. Spatial and environmental variables contributed similar percentages to the total variance in insect species richness, abundance or composition. No significant effects of habitat area were observed in any response variable. Herbivore abundance showed positive responses to legume or grass cover, as well as spatial variation that was unrelated to environmental variables. Predators and parasitoids had greater effects of plant species richness and habitat edge, and less unexplained spatial variation. Individual species differed in their responses to plant variables, depending on host specialization or intraspecific aggregation. Our study highlights the importance of plant community composition and spatial variation apart from environmental variables. Spatial variation stems both from species responses to environmental features as well as species differences in habitat specialization and intraspecific aggregation.     

Moth assemblages in Costa Rica rain forest mirror small-scale topographic heterogeneity

In many tropical lowland rain forests, topographic variation increases environmental heterogeneity, thus contributing to the extraordinary biodiversity of tropical lowland forests. While a growing number of studies have addressed effects of topographic differences on tropical insect communities at regional scales (e.g., along extensive elevational gradients), surprisingly little is known about topographic effects at smaller spatial scales. The present study investigates moth assemblages in a topographically heterogeneous lowland rain forest landscape, at distances of less than a few hundred meters, in the Golfo Dulce region (SW Costa Rica). Three moth lineages—Erebidae–Arctiinae (tiger and lichen moths), the bombycoid complex, and Geometridae (inchworm moths)—were examined by means of automatic light traps in three different forest types: creek forest, slope forest, and ridge forest. Altogether, 6,543 individuals of 419 species were observed. Moth assemblages differed significantly between the three forest types regarding species richness, total abundance, and species composition. Moth richness and abundance increased more than fourfold and eightfold from creek over slope to ridge forest sites. All three taxonomic units showed identical biodiversity patterns, notwithstanding their strong differences in multiple eco-morphological traits. An indicator species analysis revealed that most species identified as characteristic were associated either with the ridge forest alone or with ridge plus slope forests, but very few with the creek forest. Despite their mobility, local moth assemblages are highly differentially filtered from the same regional species pool. Hence, variation in environmental factors significantly affects assemblages of tropical moth species at small spatial scales.     

Environmental heterogeneity,species diversity and co‐existence at different spatial scales

The positive relationship between spatial environmental heterogeneity and species diversity is a widely accepted concept, generally associated with niche limitation. However, niche limitation cannot account for negative heterogeneity–diversity relationships (HDR) revealed in several case studies. Here we explore how HDR varies at different spatial scales and provide novel theories for small‐scale species co‐existence that explain both positive and negative HDR. At large spatial scales of heterogeneity (e.g. landscape level), different communities co‐exist, promoting large regional species pool size and resulting in positive HDR. At smaller scales within communities, species co‐existence can be enhanced by increasing the number of different patches, as predicted by the niche limitation theory, or alternatively, restrained by heterogeneity. We conducted meta‐regressions for experimental and observational HDR studies, and found that negative HDRs are significantly more common at smaller spatial scales. We propose three theories to account for niche limitation at small spatial scales. (1) Microfragmentation theory: with increasing spatial heterogeneity, large homogeneous patches lose area and become isolated, which in turn restrains the establishment of new plant individuals and populations, thus reducing species richness. (2) Heterogeneity confounded by mean: when heterogeneity occurs at spatial scales smaller than the size of individual plants, which forage through the patches, species diversity can be either positively or negatively affected by a change in the mean of an environmental factor. (3) Heterogeneity as a separate niche axis: the ability of species to tolerate heterogeneity at spatial scales smaller than plant size varies, affecting HDR. We conclude that processes other than niche limitation can affect the relationship between heterogeneity and diversity.