On the effects of topography on wind and the generation of currents in a large multi-basin lake

The wind field over a lake surface is the key element in driving the exchange of momentum and energy at the free surface. It is rarely uniform, having a considerable degree of spatial variability, both on a synoptic and a local scale. Topographic features are one of the most common contributors to this variability. The aerodynamic effects of topography on the windfield, and its influence on the circulation patterns and interbasin exchange rates in a large multi-basin lake are documented. The complex landscape that surrounds this lake is dominated by the presence of a mountain peak rising over 900m above the lake level. The circulation patterns in those basins of the lake located in the leeward side of the mountain reflect the spatial patterns of the wind field generated by the surrounding topography. The spatial variations in the wind field are shown to significantly alter the residence times of each basin and the exchange rates between basins, isolating one from the other two basins of the lake. The inter-basin exchange rates determined with wind field variability are consistent with annual contaminant loadings estimated for each of the three basins, which suggests a close link between chemical and hydrodynamic behavior.     

Fine-scale spatial organization of tallgrass prairie vegetation along a topographic gradient

Fine-scale spatial patterns of native tallgrass prairie vegetation were studied on Konza Prairie, Kansas, USA. Three sites, upland, slope, and lowland, were sampled in an ungrazed watershed. Presence of vascular plant species was recorded in two 25.6 m long transects of contiguous 5×5 cm micro-quadrats on each topographic position. Spatial patterns of species and functional groups were analyzed by information theory models ofJuhász-Nagy. Within-community variability of coexistence was expressed by the diversity and spatial dependence of local species combinations. Considerable diversity in the local coexistence of species was found on each site. Upland and hillside communities were richer and more diverse in species combinations than lowland. Spatial scale effected coexistence relationships. The maxima of information theory estimates varied between 15 and 30 cm. There was no trend in the variation of characteristic scales along the topographical gradient. Above 10 m, all sites tended to be homogeneous. The analysis of spatial associations revealed that variability in the local coexistence of species was strongly constrained in all topographic positions. Overall spatial association of species was the lowest on lowland. The characteristic scales of maximum association were between 1.2 m and 3 m at all sites. The maxima of information theory estimates for the functional group-based data appeared at smaller plot sizes than for the species based analyses. Only weak spatial associations were detected among the functional groups indicating that individuals of functional groups coexist well at small scales, and form combinations close to random expectations. The length of transects did not effect the relative associations. Strong positive correlations were found between the number of components (species or functional groups) and the maxima, of information theory models suggesting that richness is a good predictor of within-community coexistence relations. However, there was no relationship between richness and the characteristic scales of community patterns.     

Using hierarchical sampling to understand scales of spatial variation in early coral recruitment

Ecological patterns are created by processes acting over multiple spatial and temporal scales. By combining spatially explicit sampling with variance components models, the relative importance of spatial scale to overall variability can be determined. We used a spatially structured experimental design in the Mombasa Marine National Park in Kenya to quantify variation in coral recruitment across four spatial scales (~1–1,000 m) and to generate hypotheses about processes affecting recruitment and potential sources of post-settlement mortality during early life history. For the dominant recruiting corals (Pocillopora spp.), variation in recruitment on surfaces protected from fish grazing was greatest at the largest spatial scale examined (1,000 m). We hypothesize that recruitment on protected surfaces varies mainly with larval delivery due to different lagoonal circulation and water flow between sites. Conversely, variation on surfaces exposed to fishes was greatest at the smallest spatial scale (1 m). We hypothesize that recruitment on exposed surfaces mainly reflects local differences in the scale and intensity of fish grazing, which may obscure larval delivery patterns. Spatial variation in recruitment can affect many ecological processes and factors, including growth, survival to maturity, the distribution of habitat, and variation in species interaction strengths. This study demonstrates how spatially explicit sampling, followed by variance components modeling to partition variance across scales, can help to identify potential drivers of patterns at each relevant scale.     

Scale-dependent spatial variability of coral assemblages along the Florida Reef Tract

 Coral reef communities of the western Atlantic have changed over the past two to three decades, but the magnitude and causes of this change remain controversial. Part of the problem is that small-scale patterns observed on individual reefs have been erroneously extrapolated to landscape and geographic scales. Understanding how reef coral assemblages vary through space is an essential prerequisite to devising sampling strategies to track the dynamics of coral reefs through time. In this paper we quantify variation in the cover of hard corals in spur-and-groove habitats (13–19 m depth) at spatial scales spanning five orders of magnitude along the Florida Reef Tract. A videographic sampling program was conducted to estimate variances in coral cover at the following hierarchical levels and corresponding spatial scales: (1) among transects within sites (0.01- to 0.1-km scale), (2) among sites within reefs (0.5- to 2-km scale), (3) among reefs within sectors of the reef tract (10- to 20-km scale), and (4) among sectors of the reef tract (50- to 100-km scale). Coral cover displayed low variability among transects within sites and among sites within reefs. This means that transects from a site adequately represented the variability of the spur-and-groove habitat of the reef as a whole. Variability among reefs within sectors was highly significant, compared with marginally significant variability among sectors. Estimates from an individual reef, therefore, did not adequately characterize nearby reefs, nor did those estimates sufficiently represent variability at the scale of the sector. The structure and composition of coral reef communities is probably determined by the interaction of multiple forcing functions operating on a variety of scales. Hierarchical analyses of coral assemblages from other geographic locations have detected high variability at scales different from those in the present study. A multiscale analysis should, therefore, precede any management decisions regarding large reef systems such as the Florida Reef Tract. Accepted: 19 July 1999     

Spatial variability in prey phenology determines predator movement patterns and prey survival

Species have phenological variation among local habitats that are located at relatively small spatial scales. However, less studies have tested how this spatial variability in phenology can mediate intra-/inter-specific interactions. When predators track phenological variation of prey among local habitats, survival of prey within a local habitat strongly influenced by phenological synchrony with their conspecifics in adjacent habitats. Theory predicts that phenological synchrony among local habitats increases prey survival in local habitat within spatially structured environments because the predators have to make a habitat choice for foraging. Consequently, total survival of prey at regional scale should be higher. By using a spatially explicit field experiment, we tested above hypothesis using a prey–predator interaction between tadpole (Rhacophorus arboreus) and newt (Cynops pyrrhogaster). We established enclosures (≈regional scale) consisting of two tanks (≈local habitat scale) with different degree of prey phenological synchrony. We found that phenological synchrony of prey between tanks within each enclosure decreased the mean residence time of the predator in each tank, which resulted in higher survival of prey at a local habitat scale, supporting the theoretical prediction. Furthermore, individual-level variation in predator residence time explained the between-tank variation in prey survival in enclosures with phenological synchrony, implying that movement patterns of the predator can mediate variation in local population dynamics of their prey. However, total survival at each enclosure was not higher under phenological synchrony. These results suggest the importance of relative timing of prey phenology, not absolute timing, among local habitats in determining prey–predator interactions.     

Spatial patterns and scaling behaviors of Steller sea lion (Eumetopias jubatus) distributions and their environment

Fractal geometry and other multi-scale analyses have become popular tools for investigating spatial patterns of animal distributions in heterogeneous environments. In theory, changes in patterns of animal distributions with changes in scale reflect transitions between the controlling influences of one environmental factor or process over another. In an effort to find linkages between Steller sea lions (Eumetopias jubatus) and their environment, the objective of this study was to determine if the spatial distribution of Steller sea lions at sea displayed similar scaling properties to the variation of two environmental features, including bathymetry and sea surface temperature (SST). Additionally, distributions of Steller sea lion point patterns were examined with respect to measurements of bathymetric complexity. From February 2000 to May 2004, satellite transmitters were deployed on 10 groups of juvenile Steller sea lions (n=52) at eight different locations within the Aleutian Islands and Gulf of Alaska. Indices of fractal dimension were calculated for each group of sea lions using a unit square box-counting method, whereas indices of bathymetry and SST patchiness were derived by conducting a variance ratio analysis over the same scales. Distributions of Steller sea lions at sea displayed self-similar fractal patterns, suggesting that individuals were distributed in a continuous hierarchical set of clumps within clumps across scales, and foraging behavior was likely influenced by a scale invariant mechanism. Patterns of bathymetric variability also were self-similar, whereas patterns of SST variability were scale dependent and failed to retain self-similar spatial structure at larger scales. These results indicate that the distributions of Steller sea lions at sea were more influenced by bathymetry than SST at the scales examined, but scale-dependent patterns in the distribution of Steller sea lions at sea or linkages with SST may have been apparent if analyses were conducted at finer spatial scales.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration.     

Scale-dependent spatial variance patterns and correlations of seabirds and prey in the southeastern Bering Sea as revealed by spectral analysis

Uni- and bivariate spectral analyses of the spatial distribution of thick-billed murres Uria lomvia and acoustic estimates of prey biomass in the southeastern Bering Sea were used to examine the spatial variance patterns of a predator and its prey at multiple spatial scales. Power, phase and coherency spectra from individual transects, as well as those averaged over all transects, were examined. The average spectra, representing a temporal scale of months, showed that murres and prey had similar spatial variance patterns and were positively correlated over the range of spatial scales studied. The individual spectra, representing a temporal scale of hours, showed several patterns that were not evident in the average spectra. In particular, the transect-level analyses showed that the correlation between murres and prey was poor at spatial scales where prey variance was relatively low. This result suggests a new hypothesis to explain poor small-scale correlations between consumers and resources: resource distribution is relatively uniform at small scales resulting in only a slight increase in foraging return for consumers showing an aggregative response at these scales. The differences among spatial scales and between the average and individual spectra illustrate how ecological patterns can vary with temporal and spatial scale.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Spatial variability in intertidal macroalgal assemblages on the North Portuguese coast: consistence between species and functional group approaches

Natural assemblages are variable in space and time; therefore, quantification of their variability is imperative to identify relevant scales for investigating natural or anthropogenic processes shaping these assemblages. We studied the variability of intertidal macroalgal assemblages on the North Portuguese coast, considering three spatial scales (from metres to 10 s of kilometres) following a hierarchical design. We tested the hypotheses that (1) spatial pattern will be invariant at all the studied scales and (2) spatial variability of macroalgal assemblages obtained by using species will be consistent with that obtained using functional groups. This was done considering as univariate variables: total biomass and number of taxa as well as biomass of the most important species and functional groups and as multivariate variables the structure of macroalgal assemblages, both considering species and functional groups. Most of the univariate results confirmed the first hypothesis except for the total number of taxa and foliose macroalgae that showed significant variability at the scale of site and area, respectively. In contrast, when multivariate patterns were examined, the first hypothesis was rejected except at the scale of 10 s of kilometres. Both uni- and multivariate results indicated that variation was larger at the smallest scale, and thus, small-scale processes seem to have more effect on spatial variability patterns. Macroalgal assemblages, both considering species and functional groups as surrogate, showed consistent spatial patterns, and therefore, the second hypothesis was confirmed. Consequently, functional groups may be considered a reliable biological surrogate to study changes on macroalgal assemblages at least along the investigated Portuguese coastline.     

Variability at different spatial scales of a coralligenous assemblage at Giannutri Island (Tuscan Archipelago,northwest Mediterranean)

This study was carried out on the rocky cliffs of Giannutri Island (Tyrrhenian Sea, Italy) to test the hypothesis that coralligenous assemblages are consistent within the bathymetric range considered (25–35 m depth) over three different spatial scales (1000 m, 100 m and 10 m). A multi-factorial sampling design was used to assess patterns of vertical distribution in the studied area. Data on the percent cover of algae and invertebrates were collected at three depths (25, 30 and 35 m) using a photographic method, and percentage cover was obtained using a visual method. Analysis of the results using ANOVA indicated that the distribution and abundance of algae and some invertebrates of the sublittoral assemblages are clearly heterogeneous. Algae, sponges and bryozoans showed significant variability in distribution and abundance at different depths, but this variability was not consistent amongst transects. Ascidians did not show any variability, while the abundance of anthozoans differed significantly amongst transects. We concluded that heterogeneity in the distribution and abundance of the taxa analysed is related to the smallest spatial scale investigated (10's of m). Some of the possible causes of the observed variability are discussed.     

Contrasting spatial heterogeneity of sessile organisms within mussel (Perna perna L.) beds in relation to topographic variability

We examined the spatial heterogeneity in three sessile rocky shore organisms, the mussel Perna perna, the barnacle Octomeris angulosa (Sowerby) and the red alga Gelidium pristoides (Turn.) at a range of continuous local scales along horizontal transects within mid- and upper mussel beds of South African shores. We also examined the relationships between variability of organisms and topographic features (rock depressions, slope, aspect), and between mussel, barnacle and algal variability over the same scales. To estimate spatial heterogeneity, we analyzed scaling properties of semivariograms using a fractal approach. Relationships between different variables at the different scales were examined by cross-semivariograms. Spatial dependence of P. perna variability increased with spatial dependence of topographic variability, so that scaling regions of mussel and topographic distributions corresponded well. This relationship often improved with larger local scales (mussel cover increased with depressions, steeper slope and aspect towards waves), while at smaller spatial scales, variability in mussel cover was less well explained by variability in topography. The variability of the barnacle O. angulosa exhibited spatial dependence, even on topographically unstructured shores. In contrast, the distribution of the alga G. pristoides revealed high fractal dimensions, showing spatial independence on topographically unstructured shores. Algae also showed a very strong negative relationship with mussels at most local scales, and a negative relationship with barnacles in upper zones, especially at larger local scales. Barnacles may show clear spatial dependence because of hydrodynamics (at larger local scales) and the need to find a future mate in close proximity (at smaller local scales), while algae may show a strong negative relationship with mussels because of competition for space.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Effect of the spatial context along the invasion process: “Hierarchical spatial” or “Host-switching spatial” hypotheses?

Very little is known about how spatial effects influence invasive species throughout the invasion sequence. We propose here two mechanisms to explain the changes in spatial effects throughout the stages of invasion, using the soybean aphid (Aphis glycines) as a model. First, the “hierarchical spatial effect” hypothesis, based on a change in the relative importance of the spatial scales throughout the invasion process, with main effect at broad scale during the first years of invasion, and main effect at local scale during the subsequent years. Second, the “host-switching spatial effect” hypothesis, stating that the spatial effect is driven by a switch in the effect of the host/habitat throughout the invasion process, from effect of main summer host/habitat during the first years of invasion to effect of overwintering host/habitat during the subsequent years. Data from governmental archives and field samplings enabled to investigate the spatial effects on aphid density at three scales (regional, landscape, local) during a 7 year period (2006–2012). Our results demonstrate that the hierarchical spatial effect hypothesis is not an adequate model for the soybean aphid, aphid density being more affected by landscape-scale factors irrespective of years. In contrast, our results are in accordance with the host-switching spatial hypothesis, with positive effect of the main summer host/habitat (soybean) during the first steps of invasion (2006–2008), followed by a positive effect of overwintering habitats (buckthorn, woodland) during the subsequent years (2010–2012). Overall, investigating these hypotheses in other systems would determine whether the same tendency is observed for other invasive species.     

Consistency and variation in kelp holdfast assemblages: Spatial patterns of biodiversity for the major phyla at different taxonomic resolutions

The focus of this study was to measure natural spatial variability in the biodiversity of fauna inhabiting kelp holdfasts in northeastern New Zealand at several spatial scales: from meters up to hundreds of kilometers. We wished to test the hypothesis that multivariate variation and biodiversity would vary significantly at different spatial scales in different ways for the major phyla in the holdfast community (Arthropoda, Annelida, Mollusca and Bryozoa). Biodiversity was considered in terms of richness, total abundance, structural composition (as measured by the Bray-Curtis dissimilarity measure) and taxonomic breadth for each major phylum and for the assemblage as a whole. We also examined the effect of taxonomic resolution on multivariate patterns. Species richness and total abundance increased with increases in holdfast volume. Multivariate variation was greatest at the smallest spatial scale for all phyla, but different phyla showed different patterns of multivariate variation at different spatial scales. Variations among locations at the largest spatial scale were primarily due to differences in the composition and richness of bryozoans and molluscs. Location effects became less and less distinct with decreases in taxonomic resolution. There were very few significant differences in richness or abundance for holdfasts of a given volume, taxonomic breadth did not vary significantly across locations, nor did the proportional abundances of phyla. These consistencies across large spatial scales in the absence of environmental impacts and results from other studies suggest that holdfast communities in New Zealand systems would provide a useful model assemblage against which future impacts may be detected as changes in proportions of component phyla. In addition, high variability detected at small and large scales at the species level, especially for bryozoans and molluscs, suggest that these communities may also provide unique opportunities for studying and understanding sources and functions of marine biodiversity.     

Functional differences between summer and winter season rain assessed with MODIS‐derived phenology in a semi‐arid region

Questions: We asked several linked questions about phenology and precipitation relationships at local, landscape, and regional spatial scales within individual seasons, between seasons, and between year temporal scales. (1) How do winter and summer phenological patterns vary in response to total seasonal rainfall? (2) How are phenological rates affected by the previous season rainfall? (3) How does phenological variability differ at landscape and regional spatial scales and at season and inter‐annual temporal scales? Location: Southern Arizona, USA. Methods: We compared satellite‐derived phenological variation between 38 distinct 625‐km² landscapes distributed in the northern Sonoran Desert region from 2000 to 2007. Regression analyses were used to identify relationships between landscape phenology dynamics in response to precipitation variability across multiple spatial and temporal scales. Results: While both summer and winter seasons show increases of peak greenness and peak growth with more precipitation, the timing of peak growth was advanced with more precipitation in winter, while the timing of peak greenness was advanced with more precipitation in summer. Surprisingly, summer maximum growth was negatively affected by winter precipitation. The spatial variations between summer and winter phenology were similar in magnitude and response. Larger‐scale spatial and temporal variation showed strong differences in precipitation patterns; however the magnitudes of phenological spatial variability in these two seasons were similar. Conclusions: Vegetation patterns were clearly coupled to precipitation variability, with distinct responses at alternative spatial and temporal scales. Disaggregating vegetation into phenological variation, spanning value, timing, and integrated components revealed substantial complexity in precipitation‐phenological relationships.     

The role of recruitment in structuring patterns of small-scale spatial variability in intertidal and subtidal algal turfs

Recruitment is often important in structuring patterns of distribution and abundance of algal assemblages. Intertidal and subtidal turfing algal assemblages consistently vary on small spatial scales (tens of centimetres), and this variability may be due to patterns of recruitment varying on similar spatial scales. The validity of this model was evaluated by testing the hypothesis that the numbers and types of taxa recruiting to turfs would vary at small spatial scales within intertidal and within subtidal habitats. Abundances of algal recruits were estimated on sandstone plates that were placed at a number of spatial scales within intertidal and within subtidal habitats (centimetres to tens of metres). Significant differences in entire assemblages were found only between habitats. This was explained by abundances of individual taxa, which generally varied between intertidal and subtidal habitats or between sites within habitats. Only small proportions of the overall spatial variation (dissimilarity) could be explained at the scale of replicate recruitment plates that were centimetres apart. Results indicate that while recruitment may contribute to differences between intertidal and subtidal habitats, it cannot explain the small-scale spatial variability in established turfing algal assemblages within these habitats. There was some evidence to suggest that recruitment may contribute to variability in established turfing algal assemblages but only over longer time scales than examined here.     

Lack of spatial coherence of predators with prey: a bioenergetic explanation for Atlantic cod feeding on capelin

We tested two biologically based predictions that potentially influence scales of spatial association between Atlantic cod, Gadus morhua , and prey populations of capelin, Mallotus vilhsus . If cod aggregate in response to concentrations of prey, then spatial association (coherence) between capelin and cod was predicted to peak at the scale of maximum capelin spatial variance. If capelin-cod coherence did not match the scale of maximum prey spatial variability, then capelin-cod coherence was predicted to peak at the spatial scale that maximizes net energetic benefit to the predator. Contrary to predictions, we found no evidence of aggregative responses of cod to capelin over resolution scales of 20 m to 10 km. This result was observed consistently at the temporal scale of a single transect ( c . 1 h duration) and at the scale of averaged transects ( c . 2 weeks duration). Estimates of cod foraging energetics showed that they were not constrained by physiology to aggregate relative to capelin at any scale less than 10 km. A net energetic gain of 478 to 784 kJ would result if a 44 cm, 752 g cod consumed a ration of eight to 12 capelin over a period of 58 h. Energetic calculations included costs of egestion and excretion (317 to 476 kJ), maintenance (58 kJ), digestion (125 to 188 kJ), and continuous swimming during ration assimilation (79 kJ). During this period, a 44 cm cod could travel over 38 km swimming at 1 b.l. s⁻¹. Foraging cod are virtually certain to encounter capelin over this distance based on the abundance of pre-spawning capelin present in coastal bays during the spawning season. This study illustrates that aggregative responses of predators do no occur at all scales and possibly occur over a very limited range of scales.     

Ecosystem stability in space: α, β and γ variability

The past two decades have seen great progress in understanding the mechanisms of ecosystem stability in local ecological systems. There is, however, an urgent need to extend existing knowledge to larger spatial scales to match the scale of management and conservation. Here, we develop a general theoretical framework to study the stability and variability of ecosystems at multiple scales. Analogously to the partitioning of biodiversity, we propose the concepts of alpha, beta and gamma variability. Gamma variability at regional (metacommunity) scale can be partitioned into local alpha variability and spatial beta variability, either multiplicatively or additively. On average, variability decreases from local to regional scales, which creates a negative variability–area relationship. Our partitioning framework suggests that mechanisms of regional ecosystem stability can be understood by investigating the influence of ecological factors on alpha and beta variability. Diversity can provide insurance effects at the various levels of variability, thus generating alpha, beta and gamma diversity–stability relationships. As a consequence, the loss of biodiversity and habitat impairs ecosystem stability at the regional scale. Overall, our framework enables a synthetic understanding of ecosystem stability at multiple scales and has practical implications for landscape management.     

Local climate mediates spatial and temporal variation in carabid beetle communities in three forests in Mount Odaesan,Korea

1. Global environmental change can dramatically alter the composition of floral and faunal communities, and elucidating the mechanisms underlying this process is important for predicting its outcomes. Studies on global climate change have mostly focused on statistical summaries within wide spatial and temporal scales; less attention has been paid to variability in microclimates at narrower spatial and temporal scales. 2. The microclimate is the suite of climatic conditions measured in a local area. Environmental variables at the microclimatic scale can be critical for the ecology of organisms inhabiting each area. The effect of spatial and temporal changes in the microclimate on the ecology of carabid beetle communities in three sites on Mount Odaesan, Korea was examined. 3. Carabid beetle communities and quantified site‐specific environmental factors from measurements of air temperature, air humidity, light intensity and soil temperature over 5 years (2010–2015) were surveyed. 4. It was found that microclimatic variables and the patterns of temporal changes in carabid beetle communities differed between the three sites within the single mountain system. Microclimatic variables influencing temporal changes in beetle communities also differed between the sites. Therefore, it is suggested that variation in local microclimates affects spatial and temporal variation in carabid beetle communities at a local scale. 5. The present results demonstrate the importance of regular surveys of communities at local scales. Such surveys are expected to reveal an additional fraction of variation in communities and underlying processes that have been overlooked in studies of global community patterns and change.