Implications of scale for patterns and processes in stream ecology

Abstract Although the scale-dependence of ecological patterns and processes is recognized by freshwater ecologists, current knowledge of scale effects is rudimentary and non-quantitative. We review issues of spatial and temporal scale in this paper to highlight conceptual problems relating to scale and some potential solutions. We present examples of how the spatial scale of a study influences observed patterns and their interpretation, and discuss how the size of an experimental arena influences the degree to which the dynamics of studied populations are influenced by exchange processes (immigration and emigration). The results of small-scale field experiments in streams will often be strongly influenced by the per capita exchange rates of organisms and differences in exchange rates may explain differences in the perceived effects of stream manipulations across scales. Spatial extent also influences the amount of spatial heterogeneity within a study site or arena, with important consequences for the outcome of predator-prey interactions. We suggest that changes in the availability of prey refuges may help explain why predator manipulations in streams appear to weaken as arena size increases. We also recommend that new techniques for decomposing and quantifying spatial heterogeneity be applied to characterize scale-dependent variation in freshwater systems. Lastly, we discuss the pitfalls of mismatching the temporal scale of experiments and models. Models incorporating spatial heterogeneity and the behaviour of organisms are needed to predict the short-term outcome of perturbations in streams, whereas models predicting long-term dynamics will need to integrate the impacts of episodic disturbance and all life history stages of organisms. In general, we recommend that freshwater ecologists undertake more multi-scale sampling and experimentation to examine patterns and processes at multiple scales, and make greater attempts to match the scales of their observations and experiments to the characteristic scales of the phenomena that they investigate.     

Testing for temporal coherence across spatial extents: the roles of climate and local factors in regulating stream macroinvertebrate community dynamics

Temporal coherence or spatial synchrony refers to the tendency of population, community or ecosystem dynamics to behave similarly among locations through time as a result of spatially‐correlated environmental stochasticity (Moran effect), dispersal or trophic interactions. While terrestrial studies have treated synchrony mainly as a population‐level concept, the majority of freshwater studies have focused on community‐level patterns, particularly in lake planktonic communities. We used spatially and temporally hierarchical data on benthic stream invertebrates across six years, with three seasonal samples a year, in 11 boreal streams to assess temporal coherence at three spatial extents: 1) among regions (watersheds), 2) among streams within a region, and 3) among riffles within a stream, using the average of correlation coefficients for stream/riffle pairs across years. Our results revealed the primacy of strongly synchronized climatic factors (precipitation, air temperature) in inducing temporal coherence of macroinvertebrate assemblages across geographically distinct sites (i.e. Moran effect). Coherence tended to decrease with increasing spatial extent, but positive coherence was detected for most biological variables even at the largest extent (about 350 km). The generally high level of coherence reflected the strong seasonality of boreal freshwater communities. A hydrologically exceptional year enhanced the synchrony of biological variables, particularly total macroinvertebrate abundance. Regionally low precipitation in that year led to a substantial decrease in benthic densities across a broad spatial extent, followed by a rapid post‐drought recovery. Coherence at the among‐riffle (within‐stream) extent was lower than expected, implying that local‐scale habitat filters determine community dynamics at smaller spatial extents. Thus, temporal coherence of stream benthic communities appears to be controlled by partly different processes at different spatial scales.     

Design and analysis of biodiversity field experiments

The design and analysis of field experiments have figured prominently in the current debate about biodiversity and ecosystem function. These debates have identified important issues about species traits, functional groups, and community assembly, but a broader debate needs to include discussions about the scale (grain and extent) of experiments relative to natural spatial and temporal heterogeneity. In addition, alternative statistical analyses need to be explored that focus on comparison among several statistical models rather than simple hypothesis testing. Analyses of the first 2 years of data from a new biodiversity field experiment are used to illustrate these concepts. A traditional one-way ANOVA demonstrates the expected increase in aboveground biomass with higher levels of vascular plant diversity. Further analysis demonstrates that this relationship is absent when the community contains either Arrhenatherum elatius or Holcus lanatus, two dominant species of grass. Variance in biomass is also a function of diversity and both spatial and temporal heterogeneity are significant factors in the analysis despite precautions taken to minimize them. These examples illustrate the degree to which the analysis of a field experiment influences the interpretation of the observed results. Ultimately, results from field experiments must be validated through continued comparisons among field experiments, mathematical models, laboratory trials, and mesocosm experiments.     

A cross-system synthesis of consumer and nutrient resource control on producer biomass

Nutrient availability and herbivory control the biomass of primary producer communities to varying degrees across ecosystems. Ecological theory, individual experiments in many different systems, and system-specific quantitative reviews have suggested that (i) bottom-up control is pervasive but top-down control is more influential in aquatic habitats relative to terrestrial systems and (ii) bottom-up and top-down forces are interdependent, with statistical interactions that synergize or dampen relative influences on producer biomass. We used simple dynamic models to review ecological mechanisms that generate independent vs. interactive responses of community-level biomass. We calibrated these mechanistic predictions with the metrics of factorial meta-analysis and tested their prevalence across freshwater, marine and terrestrial ecosystems with a comprehensive meta-analysis of 191 factorial manipulations of herbivores and nutrients. Our analysis showed that producer community biomass increased with fertilization across all systems, although increases were greatest in freshwater habitats. Herbivore removal generally increased producer biomass in both freshwater and marine systems, but effects were inconsistent on land. With the exception of marine temperate rocky reef systems that showed positive synergism of nutrient enrichment and herbivore removal, experimental studies showed limited support for statistical interactions between nutrient and herbivory treatments on producer biomass. Top-down control of herbivores, compensatory behaviour of multiple herbivore guilds, spatial and temporal heterogeneity of interactions, and herbivore-mediated nutrient recycling may lower the probability of consistent interactive effects on producer biomass. Continuing studies should expand the temporal and spatial scales of experiments, particularly in understudied terrestrial systems; broaden factorial designs to manipulate independently multiple producer resources (e.g. nitrogen, phosphorus, light), multiple herbivore taxa or guilds (e.g. vertebrates and invertebrates) and multiple trophic levels; and - in addition to measuring producer biomass - assess the responses of species diversity, community composition and nutrient status.     

Genetic structure of freshwater Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) populations from the lakes Onega and Ladoga of northwest Russia and implications for conservation

Freshwater Atlantic salmon (Salmo salar L.) populations are a living example of adaptation to the changing conditions caused by glacial cycles. The uniqueness of these populations is emphasized by almost complete resistance to the dangerous parasite Gyrodactylus salaris. In Europe, freshwater salmon populations occur primarily in north-western Russia in the republic of Karelia. These systems include Lakes Ladoga and Onega, the two largest lakes in Europe, each of which harbours a number of freshwater salmon spawning rivers. We used microsatellite markers to study the genetic structure and temporal stability of 11 freshwater salmon populations in Russian Karelia. Populations clustered according their region of origin. Although temporal variation in allele frequencies was observed in the majority of temporal comparisons, various lines of evidence demonstrated that this influence was relatively minor compared to spatial variation that explained eight times more of the variability than temporal variation. Temporal stability tended to occur in populations from rivers with a higher linear lake coefficient. The high level of genetic structuring observed in both lake systems and the apparent low level of migration between populations suggests that treating each river as a separate management unit is recommended. In addition, as the number of populations is large, the best strategy for such fine scale management would be to ensure that the level of natural reproduction in each river is sufficient to sustain the population. A prioritization strategy for population conservation based on estimating the relative roles of different evolutionary forces shaping the gene pools highlighted a number of populations where further monitoring is warranted and also identified populations which could be prioritized for conservation as living gene banks in the event that conservation resources are limited. This prioritization agreed well with the occurrence of temporal (in)stability.     

Effects of Land Use on Lake Nutrients: The Importance of Scale,Hydrologic Connectivity,and Region

Catchment land uses, particularly agriculture and urban uses, have long been recognized as major drivers of nutrient concentrations in surface waters. However, few simple models have been developed that relate the amount of catchment land use to downstream freshwater nutrients. Nor are existing models applicable to large numbers of freshwaters across broad spatial extents such as regions or continents. This research aims to increase model performance by exploring three factors that affect the relationship between land use and downstream nutrients in freshwater: the spatial extent for measuring land use, hydrologic connectivity, and the regional differences in both the amount of nutrients and effects of land use on them. We quantified the effects of these three factors that relate land use to lake total phosphorus (TP) and total nitrogen (TN) in 346 north temperate lakes in 7 regions in Michigan, USA. We used a linear mixed modeling framework to examine the importance of spatial extent, lake hydrologic class, and region on models with individual lake nutrients as the response variable, and individual land use types as the predictor variables. Our modeling approach was chosen to avoid problems of multi-collinearity among predictor variables and a lack of independence of lakes within regions, both of which are common problems in broad-scale analyses of freshwaters. We found that all three factors influence land use-lake nutrient relationships. The strongest evidence was for the effect of lake hydrologic connectivity, followed by region, and finally, the spatial extent of land use measurements. Incorporating these three factors into relatively simple models of land use effects on lake nutrients should help to improve predictions and understanding of land use-lake nutrient interactions at broad scales.     

Influence of spatial and temporal variations on organic pollutant biodegradation rates in an estuarine environment.

The influence of spatial and temporal environmental variations on rates of organic pollutant biodegradation were assessed by using heterotrophic uptake kinetics. These studies were conducted at three sites, representing the gradient from freshwater to estuarine to marine systems. Of the compounds tested, total uptake Vmax rates decreased in the order of nitrilotriacetic acid, m-cresol, chlorobenzene, and 1,2,4-trichlorobenzene. In general, the freshwater site exhibited the highest uptake rates, with somewhat lower rates at the estuarine site. Rates at the marine site were much lower than at the other sites, except during the winter. Metabolic rates at both the freshwater and estuarine areas were significantly decreased during periods of low water temperature. Rates at the marine site were relatively uniform throughout the year. Linear regression analysis was used to compare m-cresol biodegradation rates to characteristics of the microbial community, which included direct microscopic counts, CFU counts, and cellular incorporation of amino acids. The observed rates did not consistently correlate well with any of the measured characteristics of the microbial community.     

Modeling denitrification in aquatic sediments

Sediment denitrification is a major pathway of fixed nitrogen loss from aquatic systems. Due to technical difficulties in measuring this process and its spatial and temporal variability, estimates of local, regional and global denitrification have to rely on a combination of measurements and models. Here we review approaches to describing denitrification in aquatic sediments, ranging from mechanistic diagenetic models to empirical parameterizations of nitrogen fluxes across the sediment-water interface. We also present a compilation of denitrification measurements and ancillary data for different aquatic systems, ranging from freshwater to marine. Based on this data compilation we reevaluate published parameterizations of denitrification. We recommend that future models of denitrification use (1) a combination of mechanistic diagenetic models and measurements where bottom-waters are temporally hypoxic or anoxic, and (2) the much simpler correlations between denitrification and sediment oxygen consumption for oxic bottom waters. For our data set, inclusion of bottom water oxygen and nitrate concentrations in a multivariate regression did not improve the statistical fit.     

Abundance of sea kraits correlates with precipitation

Recent studies have shown that sea kraits (Laticauda spp.)--amphibious sea snakes--dehydrate without a source of fresh water, drink only fresh water or very dilute brackish water, and have a spatial distribution of abundance that correlates with freshwater sites in Taiwan. The spatial distribution correlates with sites where there is a source of fresh water in addition to local precipitation. Here we report six years of longitudinal data on the abundance of sea kraits related to precipitation at sites where these snakes are normally abundant in the coastal waters of Lanyu (Orchid Island), Taiwan. The number of observed sea kraits varies from year-to-year and correlates positively with previous 6-mo cumulative rainfall, which serves as an inverse index of drought. Grouped data for snake counts indicate that mean abundance in wet years is nearly 3-fold greater than in dry years, and this difference is significant. These data corroborate previous findings and suggest that freshwater dependence influences the abundance or activity of sea kraits on both spatial and temporal scales. The increasing evidence for freshwater dependence in these and other marine species have important implications for the possible impact of climate change on sea snake distributions.     

Mechanisms influencing competition between hatchery and wild juvenile anadromous Pacific salmonids in fresh water and their relative competitive abilities

Avoiding negative effects of competition from released hatchery salmonids on wild fish is a primary concern for recovery efforts and fisheries management. Several factors affect competition among juvenile salmonids including: (1) whether competition is intra- or interspecific, (2) duration of freshwater cohabitation of hatchery and wild fish, (3) relative body size, (4) prior residence, (5) environmentally induced developmental differences, and (6) fish density. Intraspecific competition is expected to be greater than interspecific because of greater niche overlap between conspecific hatchery and wild fish. Competition is expected to increase with prolonged freshwater cohabitation. Hatchery smolts are often larger than wild, and larger fish are usually superior competitors. However, wild fish have the advantage of prior residence when defending territories and resources in natural streams. Hatchery-induced developmental differences are variable and can favor both hatchery and wild fish. Although all these factors influence competitive interactions, fish density of the composite population (wild + hatchery fish) in relation to habitat carrying capacity likely exerts the greatest influence. The extent of competition and relative competitive ability of wild and hatchery fish can be determined by additive and substitutive experimental designs, respectively, and the limited body of substitutive experiments suggests that the relative competitive ability of hatchery and wild fish is approximately equal when measured as growth. Conducting substitutive experiments becomes difficult as the spatial and temporal scales increase. Large-scale experiments comparing supplemented and control reaches or streams hold some promise for quantifying the effects of released hatchery fish on wild fish behavior, growth and survival.     

Spatial scale and the aggregation of stream macroinvertebrates associated with leaf packs

1. An experimental field study examined the aggregation of stream macroinvertebrates associated with leaf packs over different spatial scales (several metres–km) (extent), at different patch sizes (grain) and temporal scales (2 and 4 weeks). 2. Standardized leaf packs were constructed and set in eighteen blocks of nine equally spaced packs in glide areas over a 2 km stretch of a wooded stream. The distribution of macroinvertebrates colonizing the artificial leaf packs was investigated to examine the extent of both intraspecific and interspecific aggregation across leaf packs. 3. All major colonizing taxa were intraspecifically aggregated across the leaf packs. Aggregation decreased with increasing patch size (grain) (from pack to block), and also decreased with decreasing spatial extent (from 2 km stretch to within-block scale) with patch size held constant. Interspecific associations among all major taxa were not common on most occasions at the short temporal scale, although the proportion of significant associations tended to increase somewhat over time and with spatial scale, but did not exceed 42% of all possible associations. The vast majority of significant associations were positive rather than negative. 4. The influence of heterogeneity in a number of environmental variables measured for each leaf pack (accumulated detritus and sediment, leaf mass, flow and depth) on the distribution of invertebrates was considered, but this could only partially explain the variation in macroinvertebrate abundance across leaf packs. 5. The roles of intrinsic aggregation and stochastic processes were examined as alternative explanations for the distribution patterns observed. It is apparent from this study that intrinsic aggregation, in concert with resource partitioning, influences the community structure of stream macroinvertebrates associated with leaf packs. These findings may also have implications for the distribution of taxa in the benthos as a whole.     

The spatial scaling of beta diversity

Beta diversity is an important concept used to describe turnover in species composition across a wide range of spatial and temporal scales, and it underpins much of conservation theory and practice. Although substantial progress has been made in the mathematical and terminological treatment of different measures of beta diversity, there has been little conceptual synthesis of potential scale dependence of beta diversity with increasing spatial grain and geographic extent of sampling. Here, we evaluate different conceptual approaches to the spatial scaling of beta diversity, interpreted from ‘fixed’ and ‘varying’ perspectives of spatial grain and extent. We argue that a ‘sliding window’ perspective, in which spatial grain and extent covary, is an informative way to conceptualize community differentiation across scales. This concept more realistically reflects the varying empirical approaches that researchers adopt in field sampling and the varying scales of landscape perception by different organisms. Scale dependence in beta diversity has broad implications for emerging fields in ecology and biogeography, such as the integration of fine‐resolution ecogenomic data with large‐scale macroecological studies, as well as for guiding appropriate management responses to threats to biodiversity operating at different spatial scales.     

Overview: Spatial information and indicators for sustainable management of natural resources

Natural resource management (NRM) is becoming increasingly important at all scales, local, regional, national and global, because of an increasing human population and increasing per capita use of resources and space. Conflicts are intensifying between different interest groups. Production and conservation aspects are particularly debated because conservation often conflicts with economic and social sustainability. There is public demand for objective decision based NRM but limitations are all pervasive due to the spatial and temporal complexity and interdisciplinary nature.This special issue explores the use of spatial data and models to overcome some limitations of NRM decision making. The papers in this issue show modern approaches of natural resources management with a particular focus on spatial data collection, analysis and the development of spatial indicators. This issue presents a balanced mix of review and research papers that give examples of how to find or improve the spatial information base for evidence-based decision making.This overview makes the argument that understanding complex spatial pattern and processes, and the development of spatial indicators, is an essential aspect of evidence-based NRM. If spatial and temporal patterns are complex, ecological evidence from field data or experiments may have limited value for NRM and observational study designs become more appropriate for understanding complex spatial pattern and processes. Data quality should be documented as a combination of accuracy and spatio-temporal representativeness in order to be useful in the NRM decision process.     

Spatio‐temporal scaling of biodiversity and the species–time relationship in a stream fish assemblage

1. The increase of species richness with the area of the habitat sampled, that is the species–area relationship, and its temporal analogue, the species–time relationship (STR), are among the few general laws in ecology with strong conservation implications. However, these two scale‐dependent phenomena have rarely been considered together in biodiversity assessment, especially in freshwater systems. 2. We examined how the spatial scale of sampling influences STRs for a Central‐European stream fish assemblage (second‐order Bernecei stream, Hungary) using field survey data in two simulation‐based experiments. 3. In experiment one, we examined how increasing the number of channel units, such as riffles and pools (13 altogether), and the number of field surveys involved in the analyses (12 sampling occasions during 3 years), influence species richness. Complete nested curves were constructed to quantify how many species one observes in the community on average for a given number of sampling occasions at a given spatial scale. 4. In experiment two, we examined STRs for the Bernecei fish assemblage from a landscape perspective. Here, we evaluated a 10‐year reach level data set (2000–09) for the Bernecei stream and its recipient watercourse (third‐order Kemence stream) to complement results on experiment one and to explore the mechanisms behind the observed patterns in more detail. 5. Experiment one indicated the strong influence of the spatial scale of sampling on the accumulation of species richness, although time clearly had an additional effect. The simulation methodology advocated here helped to estimate the number of species in a diverse combination of spatial and temporal scale and, therefore, to determine how different scale combinations influence sampling sufficiency. 6. Experiment two revealed differences in STRs between the upstream (Bernecei) and downstream (Kemence) sites, with steeper curves for the downstream site. Equations of STR curves were within the range observed in other studies, predominantly from terrestrial systems. Assemblage composition data suggested that extinction–colonisation dynamics of rare, non‐resident (i.e. satellite) species influenced patterns in STRs. 7. Our results highlight that the determination of species richness can benefit from the joint consideration of spatial and temporal scales in biodiversity inventory surveys. Additionally, we reveal how our randomisation‐based methodology may help to quantify the scale dependency of diversity components (α, β, γ) in both space and time, which have critical importance in the applied context.     

环境影响评价的尺度约束性及技术框架

尺度约束是地表复杂系统的基本规律,环评尺度约束常隐存于主观经验或零存于环评导则中,环评尺度约束较少被明确关注。讨论了环境影响尺度约束、环评尺度约束和环评技术框架尺度约束。研究发现,环境影响的尺度约束性内在原因在于环境影响主体、客体和影响内容存在等级结构;环评受空间、时间和分析三类尺度约束,空间尺度约束体现于空间范围和空间信息分辨率对环评影响,时间尺度约束体现于环评的时长和时频,分析尺度约束表现为环评技术方法和环境影响主观认知水平对评价的影响,三类尺度约束同时贯穿于环评过程,任何环评都可以在三类尺度空间中定位;环评技术框架的关键环节都受空间、时间和分析尺度约束,且以环境影响主体的空间、时间尺度为核心,具有一定弹性,一般空间或时间范围先放大再缩小、分辨率由粗到细。     

Spatial species-richness gradients across scales: a meta-analysis

Aim We surveyed the empirical literature to determine how well six diversity hypotheses account for spatial patterns in species richness across varying scales of grain and extent. Location Worldwide. Methods We identified 393 analyses (‘cases’) in 297 publications meeting our criteria. These criteria included the requirement that more than one diversity hypothesis was tested for its relationship with species richness. We grouped variables representing the hypotheses into the following ‘correlate types’: climate/productivity, environmental heterogeneity, edaphics/nutrients, area, biotic interactions and dispersal/history (colonization limitation or other historical or evolutionary effect). For each case we determined the ‘primary’ variable: the one most strongly correlated with taxon richness. We defined ‘primacy’ as the proportion of cases in which each correlate type was represented by the primary variable, relative to the number of times it was studied. We tested for differences in both primacy and mean coefficient of determination of the primary variable between the hypotheses and between categories of five grouping variables: grain, extent, taxon (animal vs. plant), habitat medium (land vs. water) and insularity (insular vs. connected). Results Climate/productivity had the highest overall primacy, and environmental heterogeneity and dispersal/history had the lowest. Primacy of climate/productivity was much higher in large‐grain and large‐extent studies than at smaller scales. It was also higher on land than in water, and much higher in connected systems than in insular ones. For other hypotheses, differences were less pronounced. Throughout, studies on plants and animals showed similar patterns. Coefficients of determination of the primary variables differed little between hypotheses and across the grouping variables, the strongest effects being low means in the smallest grain class and for edaphics/nutrients variables, and a higher mean for water than for land in connected systems but vice versa in insular systems. We highlight areas of data deficiency. Main conclusions Our results support the notion that climate and productivity play an important role in determining species richness at large scales, particularly for non‐insular, terrestrial habitats. At smaller extents and grain sizes, the primacy of the different types of correlates appears to differ little from null expectation. In our analysis, dispersal/history is rarely the best correlate of species richness, but this may reflect the difficulty of incorporating historical factors into regression models, and the collinearity between past and current climates. Our findings are consistent with the view that climate determines the capacity for species richness. However, its influence is less evident at smaller spatial scales, probably because (1) studies small in extent tend to sample little climatic range, and (2) at large grains some other influences on richness tend to vary mainly within the sampling unit.     

Predator Assemblage Structure and Temporal Variability of Species Richness and Abundance in Forests of High Tree Diversity

Predators significantly affect ecosystem functions, but our understanding of to what extent findings can be transferred from experiments and low‐diversity systems to highly diverse, natural ecosystems is limited. With a particular threat of biodiversity loss at higher trophic levels, however, knowledge of spatial and temporal patterns in predator assemblages and their interrelations with lower trophic levels is essential for assessing effects of trophic interactions and advancing biodiversity conservation in these ecosystems. We analyzed spatial and temporal variability of spider assemblages in tree species‐rich subtropical forests in China, across 27 study plots varying in woody plant diversity and stand age. Despite effects of woody plant richness on spider assemblage structure, neither habitat specificity nor temporal variability of spider richness and abundance were influenced. Rather, variability increased with forest age, probably related to successional changes in spider assemblages. Our results indicate that woody plant richness and theory predicting increasing predator diversity with increasing plant diversity do not necessarily play a major role for spatial and temporal dynamics of predator assemblages in such plant species‐rich forests. Diversity effects on biotic or abiotic habitat conditions might be less pronounced across our gradient from medium to high plant diversity than in previously studied less diverse systems, and bottom‐up effects might level out at high plant diversity. Instead, our study highlights the importance of overall (diversity‐independent) environmental heterogeneity in shaping spider assemblages and, as indicated by a high species turnover between plots, as a crucial factor for biodiversity conservation at a regional scale in these subtropical forests.     

Scale,disturbance and productivity control the native‐exotic richness relationship

The relationship between native and exotic species richness may be highly context‐dependent. Spatial scale, including both plot size (grain) and study area (extent), is likely to influence this relationship, as are environmental conditions such as resource availability and disturbance intensity. We used experimental manipulations of soil fertility and disturbance in a California coastal grassland to directly examine the sensitivity of the native–exotic richness relationship (NERR) to these factors across five grain sizes and two spatial extents. The slope of the NERR was a function of grain size, extent and treatment. Over small spatial extents, native and exotic richness were usually uncorrelated. Across a larger extent, NERRs were negative in control plots, neutral in disturbance plots, and positive in plots with experimentally reduced soil fertility. These patterns were strongest for small grain sizes. We verify the importance of spatial grain in determining the NERR, and emphasize the role of spatial extent.     

Scale for resource selection functions

Resource selection functions (RSFs) are statistical models defined to be proportional to the probability of use of a resource unit. My objective with this review is to identify how RSFs can be used to unravel the influence of scale in habitat selection. In wildlife habitat studies, including radiotelemetry, RSFs can be estimated using a variety of statistical methods, all of which can be used to explore the role of scale. All RSFs are bounded by the resolution of data and the spatial extent of the study area, but also allow predictor covariates to be measured at a variety of scales. Conditional logistic regression permits designs (e.g. matched case) that relate the process of habitat selection to a limited domain of resource units that might better characterize what is truly ‘available’ to the animal. Scale influences the process of habitat selection, e.g. food resources are often selected at fine spatial scales, whereas landscape patterns at much larger scales typically influence the location of home ranges. Scale also influences appropriate sampling in many ways: (1) heterogeneity might be obliterated (transmutation) if resolution or grain size is too large, (2) variance of habitat characteristics might be undersampled if extent or domain is too small, (3) timing and duration of observations can influence RSF models, and (d) both spatial and temporal autocorrelations can vary directly with the intensity of sampling. Using RSFs, researchers can examine habitat selection at multiple scales, and predictive models that bridge scales can be estimated. Using Geographical Information Systems, predictor covariates in RSF models can be measured at different scales easily so that the predictive ability of models at alternative spatial and temporal domains can be explored by the investigator. Identification of the scale that best explains the data can be evaluated by comparing alternative models using information‐theoretic metrics such as Akaike Information Criteria, and predictive capability of the models can be assessed using k‐fold cross validation.