Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis

It has been suggested that a heterogeneous environment enhances species richness and allows for the coexistence of species. However, there is increasing evidence that environmental heterogeneity can have no effect or even a negative effect on plant species richness and plant coexistence at a local scale. We examined whether plant species richness increases with local heterogeneity in the water table depth, microtopography, pH and light availability in a swamp forest community at three local spatial scales (grain: 0.6, 1.2 and 11.4 m). We also used the variance partitioning approach to assess the relative contributions of niche-based and other spatial processes to species occurrence. We found that heterogeneity in microtopography and light availability positively correlated with species richness, in accordance with the habitat heterogeneity hypothesis. However, we recorded different heterogeneity-diversity relationships for particular functional species groups. An increase in the richness of bryophytes and woody plant species was generally related to habitat heterogeneity at all measured spatial scales, whereas a low impact on herbaceous species richness was recorded only at the 11.4 m scale. The distribution of herbaceous plants was primarily explained by other spatial processes, such as dispersal, in contrast to the occurrence of bryophytes, which was better explained by environmental factors. Our results suggest that both niche-based and other spatial processes are important determinants of the plant composition and species turnover at local spatial scales in swamp forests.     

Phytoplankton,physical and chemical microscale variations in three brackish rock pools

Variations at the microscale in phytoplankton distribution with respect to physical and chemical variables in three brackish rock pools of different depth, size and exposure were studied during the growth season. Three hypotheses were made about the microspatial characteristics of the rock pool habitat: (i) there are distinct microscale variations in physical and chemical variables, (ii) the microscale distribution of phytoplankton is related to characteristic physical and/or chemical variables, and (iii) microscale variations are more pronounced in deep rock pools. Variations were studied at a 10 cm scale by close interval siphon sampling. Physical and chemical variations were small irrespective of rock pool. Prevailing weather, as well as the similar round basin shape of the pools contributed to the similarities. Variations in phytoplankton microscale distributions were clear and consistent throughout the study period, and less variable in the deepest pool compared to the shallower ones. The distribution of dominant phytoplankton species correlated with microscale variations in temperature, dissolved oxygen and pH, often during seasonal minima or maxima. The microscale correlations implied that short reaction times of phytoplankton enabled them to exploit, or be influenced by, brief, local variations. The results demonstrated the importance of studies at fine scales to disentangle processes even in shallow weather‐influenced ecosystems.     

Association between temporal and spatial beta diversity in phytoplankton

The rates of temporal and spatial species turnover have been compared in different organisms and scales, revealing that both are not independent but, rather, associated. However, the knowledge is limited for the association between spatial turnover and temporal turnover. Here, we performed two investigations of the phytoplankton composition in the lakes of the Yangtze River catchment in China in the spring and summer of 2012, which covered regional spatial scale and two‐season temporal scale. We analysed the association between temporal and spatial species turnover in phytoplankton. The results showed that 1) the two‐season temporal turnover of phytoplankton varied based on the mean values and the coefficient of variance of environmental variables, and pH was the most important variable negatively affecting the temporal turnover; 2) the spatial beta diversity of phytoplankton in summer was higher than that in spring, and the distance decay pattern was significant in summer, but not in spring; 3) the variation in spatial turnover in spring and summer was attributed to the primary environmental variables (nitrogen, phosphorus and underwater available light) and broader‐scale spatial variables; 4) the proportion of jointly explained variation of spatial Bray–Curtis dissimilarity by the environment and space increased from ~38% (spring) to ~55% (summer), which was mainly due to the variation in spatially structured environmental variables during the two‐season temporal turnover, such as pH and ion concentrations; 5) the community compositions in summer were more similar between the lakes with similar two‐season temporal turnover. These results indicate that the spatial turnover of phytoplankton composition in summer was partially predetermined by the variation in environmental variables and phytoplankton composition during the process of two‐season temporal turnover, and highlight the understanding of temporal variations in spatial beta diversity as well as the underlying assembly mechanisms in phytoplankton.     

Determinants of beta diversity of spiders in coastal dunes along a gradient of mediterraneity

Aim The Mediterranean Basin is recognized for its high levels of species richness, rarity and endemicity. Our main aim was to evaluate the relative effects of environmental and spatial variables and their scale‐specific importance on beta diversity patterns along a gradient of mediterraneity, using spiders as a model group. Location This study was carried out in 18 coastal dune sites along the Portuguese Atlantic coast. This area encompasses 445 km and comprises two distinct biogeographic regions, Eurosiberian (northern coast) and Mediterranean (centre and south). Methods A forward selection procedure was carried out to select environmental and spatial variables responsible for determining beta diversity patterns. Variation partitioning and principal coordinates of neighbour matrices (PCNM) were used to estimate the contribution of pure environmental and pure spatial effects and their shared influence on beta diversity patterns and to estimate the relative importance of environmental structured variation and pure spatial variation at multiple spatial scales. Results Climate, ground vegetation dune cover and area were selected by a forward selection procedure. The same procedure identified three PCNM variables, all corresponding to large and medium spatial scales. Variation partitioning revealed that 46.1% of the variation of beta diversity patterns was explained by a combination of environmental and PCNM variables. Most of this variation (42.5%) corresponded to spatial variation (environmental spatially structured and pure spatial). Climate and vegetation structure influences were predominant at the PCNM1 and PCNM3 scales, while area was more important at the intermediate PCNM2 scale. Main conclusions Our study revealed that beta diversity of spiders was primarily controlled by a broad‐scale gradient of mediterraneity. The relative importance of environmental variables on the spider assemblage composition varied with spatial scale. This study highlights the need of considering the scale‐specific influence of niche and neutral processes on beta diversity patterns.     

The importance of local and landscape-scale processes to the occupancy of wetlands by pond-breeding amphibians

Variation in the distribution and abundance of species across landscapes has traditionally been attributed to processes operating at fine spatial scales (i.e., environmental conditions at the scale of the sampling unit), but processes that operate across larger spatial scales such as seasonal migration or dispersal are also important. To determine the relative importance of these processes, we evaluated hypothesized relationships between the probability of occupancy in wetlands by two amphibians [wood frogs (Lithobates sylvaticus) and boreal chorus frogs (Pseudacris maculata)] and attributes of the landscape measured at three spatial scales in Rocky Mountain National Park, Colorado. We used cost-based buffers and least-cost distances to derive estimates of landscape attributes that may affect occupancy patterns from the broader spatial scales. The most highly ranked models provide strong support for a positive relationship between occupancy by breeding wood frogs and the amount of streamside habitat adjacent to a wetland. The model selection results for boreal chorus frogs are highly uncertain, though several of the most highly ranked models indicate a positive association between occupancy and the number of neighboring, occupied wetlands. We found little evidence that occupancy of either species was correlated with local-scale attributes measured at the scale of individual wetlands, suggesting that processes operating at broader scales may be more important in influencing occupancy patterns in amphibian populations.     

Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities

Associations between spatial distribution of ground-beetles (Carabidae) and environmental variables were studied over three hierarchical scales in deciduous forest in central Alberta, Canada We also examined the relationship between species abundance and distribution on several scales ranging from the local scale of our study to that of the North American temperate deciduous forest Understorey plant cover, tree cover, and occurrence of other carabids were associated with distribution of particular species at the smallest ecological scales within populations However, great differences in population sues of carabid species among five distinct sites several kilometres apart were not correlated with variation in the same environmental variables In central Alberta, abundance and extent of distribution were correlated positively among the 30 carabid species collected, and distributions of the ten species classified as 'core' species were generally aggregated at all spatial scales On the continental scale, there was a significant positive correlation between abundance and distribution for the 114 species of the entire data set, and the six species meeting the criteria of 'core' taxa on this scale, were also 'core' elements in central Alberta Further analysis of covariance of core elements of species assemblages across different taxa provides a sound empirical approach for understanding community organization     

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.     

Accounting for multi‐scale spatial autocorrelation improves performance of invasive species distribution modelling (iSDM)

Aim Analyses of species distributions are complicated by various origins of spatial autocorrelation (SAC) in biogeographical data. SAC may be particularly important for invasive species distribution models (iSDMs) because biological invasions are strongly influenced by dispersal and colonization processes that typically create highly structured distribution patterns. We examined the efficacy of using a multi‐scale framework to account for different origins of SAC, and compared non‐spatial models with models that accounted for SAC at multiple levels. Location We modelled the spatial distribution of an invasive forest pathogen, Phytophthora ramorum, in western USA. Methods We applied one conventional statistical method (generalized linear model, GLM) and one nonparametric technique (maximum entropy, Maxent) to a large dataset on P. ramorum occurrence (n = 3787) to develop four types of model that included environmental variables and that either ignored spatial context or incorporated it at a broad scale using trend surface analysis, a local scale using autocovariates, or multiple scales using spatial eigenvector mapping. We evaluated model accuracies and amounts of explained spatial structure, and examined the changes in predictive power of the environmental and spatial variables. Results Accounting for different scales of SAC significantly enhanced the predictive capability of iSDMs. Dramatic improvements were observed when fine‐scale SAC was included, suggesting that local range‐confining processes are important in P. ramorum spread. The importance of environmental variables was relatively consistent across all models, but the explanatory power decreased in spatial models for factors with strong spatial structure. While accounting for SAC reduced the amount of residual autocorrelation for GLM but not for Maxent, it still improved the performance of both approaches, supporting our hypothesis that dispersal and colonization processes are important factors to consider in distribution models of biological invasions. Main conclusions Spatial autocorrelation has become a paradigm in biogeography and ecological modelling. In addition to avoiding the violation of statistical assumptions, accounting for spatial patterns at multiple scales can enhance our understanding of dynamic processes that explain ecological mechanisms of invasion and improve the predictive performance of static iSDMs.     

Metacommunity versus Biogeography: A Case Study of Two Groups of Neotropical Vegetation-Dwelling Arthropods

Biogeography and metacommunity ecology provide two different perspectives on species diversity. Both are spatial in nature but their spatial scales do not necessarily match. With recent boom of metacommunity studies, we see an increasing need for clear discrimination of spatial scales relevant for both perspectives. This discrimination is a necessary prerequisite for improved understanding of ecological phenomena across scales. Here we provide a case study to illustrate some spatial scale-dependent concepts in recent metacommunity studies and identify potential pitfalls. We presented here the diversity patterns of Neotropical lepidopterans and spiders viewed both from metacommunity and biogeographical perspectives. Specifically, we investigated how the relative importance of niche- and dispersal-based processes for community assembly change at two spatial scales: metacommunity scale, i.e. within a locality, and biogeographical scale, i.e. among localities widely scattered along a macroclimatic gradient. As expected, niche-based processes dominated the community assembly at metacommunity scale, while dispersal-based processes played a major role at biogeographical scale for both taxonomical groups. However, we also observed small but significant spatial effects at metacommunity scale and environmental effects at biogeographical scale. We also observed differences in diversity patterns between the two taxonomical groups corresponding to differences in their dispersal modes. Our results thus support the idea of continuity of processes interactively shaping diversity patterns across scales and emphasize the necessity of integration of metacommunity and biogeographical perspectives.     

Spatial variance of mobile aquatic organisms: capelin and cod in Newfoundland coastal waters

Spatial variance in the distribution of aquatic mobile organisms differs from that of passive tracers such as phytoplankton or water temperature. On average, spatial variance of phytoplankton scales with sample unit as $L^2$ or equivalently with frequency as $f^{-2}$. Limited evidence suggests that spatial variance in the distribution of mobile organisms is concentrated at relatively small scales, with little increase over larger scales: spatial variance scales as $f^{-1}$ or less. We investigated whether spatial variance in distributions of a mobile predator, Atlantic cod (Gadus morhua), and a schooling prey, capelin (Mallotus villosus), also scale with frequency as $f^{-1}$. Acoustic surveys showed that at short time scales spatial variance in cod and capelin densities, as measured by spectral density, peaked at various scales ranging from 20 m to 10 km. At longer time scales, spatial variance of cod scaled as $f^{-1.08}$ at resolutions finer than 90 m, while scaling as $f^{-0.18}$ at coarser scales. Spatial variance of capelin scaled as $f^{-1.1}$ at resolutions finer than 400 m, while scaling as $f^{-0.21}$ at coarser scales. Spatial variance plots of krill and marine birds showed similar transitions from shallow to steep scaling. Shoaling, schooling and the aggregative response by predators to concentrations of prey were three processes hypothesized to influence spatial variance in distributions of mobile organisms. Numerical experiments showed that shoaling injects variance at large to intermediate scales, resulting in scalings flatter than $f^{-1}$. Additional experiments showed that schooling produces a transition from shallow to steep scaling as frequency increases. Spatial variance patterns in cod density were not due to aggregative responses by the predator to concentrations of capelin: there was no association, on average, at resolution scales from 20 m to 10 km. Exponent values for aquatic or terrestrial mobile organisms are predicted to be approximately two at the scale of an individual organism, 0.2 at scales that contain aggregations, and two at scales larger than that of populations. These findings suggest that relations between mobile organisms and large scale habitat variables will be difficult to detect, that stratified survey designs used to estimate commercial population sizes will be inefficient, and that rates of interaction between predator and prey will be underestimated if local observations are averaged over the spatial scale of the population.     

Niche partitioning at multiple scales facilitates coexistence among mosquito larvae

A theoretical dichotomy in community ecology distinguishes between mechanisms that stabilize species coexistence and those that cause neutral drift. Stable coexistence is predicted to occur in communities where competing species have niche-partitioning mechanisms that reduce interspecific competition. Neutral communities are predicted to be structured by stochastic processes that are not influenced by species identity, but that may be influenced by priority effects and dispersal limitation. Recent developments have suggested that neutral interactions may be more common at local scales, while niche structuring may be more common at larger scales. We tested for mechanisms that could promote either stable coexistence or neutral drift in a bromeliad-dwelling mosquito community by evaluating A) if a hypothesized within-bromeliad niche partitioning mechanism occurs in the community, B) if this mechanism correlates with local species co-occurrence patterns, and C) if patterns of coexistence at the larger (metacommunity) scale were consistent with those at the local scale. We found that mosquitoes in this community do partition space within containers, and that species with the strongest potential for competition co-occurred least. Species with overlapping spatial niches minimized co-occurrence by specialising in bromeliads of differing sizes, effectively changing the scale at which they coexist. In contrast, we found no evidence to support neutral dynamics in mosquito communities at either scale. In this community, a niche-based mechanism that is predicted to stabilize species coexistence explains co-occurrence patterns within and among bromeliads.     

The relative importance of spatial and environmental processes in distribution of benthic chironomid larvae within a large and shallow lake

Although chironomids are popular model organisms in ecological research and indicators of bioassessment, the relative role of dispersal and environmental filtering in their community assembly is still poorly known, especially at fine spatial scales. In this study, we applied a metacommunity framework and used various statistical tools to examine the relative role of spatial and local environmental factors in distribution of benthic chironomid taxa and their assemblages in large and shallow Lake Balaton, Hungary. Contrary to present predictions on the metacommunity organisation of aquatic insects with winged terrestrial adults, we found that dispersal limitation can considerably affect distribution of chironomids even at lake scale. However, we also revealed the predominant influence of environmental filtering, and strong taxa–environment relationships were observed especially along sediment type, sediment organic matter content and macrophyte coverage gradients. We account that identified reference conditions and assemblages along with specified optima and tolerances of the abundant taxa can contribute to our understanding of chironomid ecology and be utilised in shallow lake bioassessment. Further, we propose that predictive models of species–environment relationships should better take into account pure spatial structuring of local communities and species-specific variability of spatial processes and environmental control even at small spatial scales.     

Shared or distinct responses between intermediate and satellite stream fish species in an altered Amazonian River?

Environmental and spatial variables can distinctly influence the occupancy frequency distributions in stream fish. From a metacommunity context, we tested the following hypothesis, intermediate species are governed by dispersal and niche-based processes; in contrast, satellite species are governed by niche-based processes. To test this, we separately analyzed three data sets, the entire metacommunity, the intermediate species and the satellite species, using a forward selection of explanatory variables, and a partial Redundancy Distance Analysis. The fish and 31 variables of 52 stream reaches of a Brazilian river basin in the Western Amazon were collected during the dry period of 2012. The results for all of the data set revealed two different patterns: on one side, satellite species revealed that niche and dispersal-based processes were the most important; on the other side, for intermediate species and for all of the species set, only dispersal-based processes were the most important. For the data set including all of the species and the intermediate species, the variance was explained mainly by landscape scale variables. By contrast, the variance within the satellite species set was explained by local scale variables. Management efforts for intermediate species should be taking at larger scale, but they are usually less critical for the maintenance of aquatic biodiversity; on the other hand, management efforts for satellite species should be taken at smaller scale and based on specific biological and ecological information for the focal species.     

What drives small‐scale spatial patterns in lotic meiofauna communities?

• 1 Lotic meiofaunal communities demonstrate extremely variable dynamics, especially when viewed at small spatial scales (≤ metres). Given the limited amount of research on lotic meiofauna, we chose to organise our discussion of their small‐scale spatial patterns around the dominant factors we believe drive their spatial distributions in streams. We separate scale‐dependent effects that structure lotic meiofauna into biotic factors (e.g. predation, food quantity/quality, dispersal) and abiotic factors (e.g. local flow dynamics and substratum characteristics).
• 2 The impact of predation on the distribution of meiofauna varies with the scale over which predators forage (e.g. fish predation influences meiofauna in different ways and at broader spatial scales than do invertebrate predators), the type of streambed substrata in which the predator‐prey interactions occur, and the dispersal ability of different meiofauna. The latter is greatly influenced by predator and prey (meiofauna) interactions with the flow environment.
• 3 Organic matter influences the small‐scale distribution of meiofauna in streams. Both its quality as food (as indicated by C:N content, ATP content, or microbial biomass) and its spatial distribution on the streambed, influence meiofauna patchiness, community structure and life history characteristics. As a habitat, the structure that organic matter provides (e.g. wood or leaves) can influence predator‐prey interactions, offer materials for case‐building and offer refugia during disturbance events ‐ all of which influence the small‐scale spatial distribution of meiofauna.
• 4 Stream flow influences the distribution of meiofauna at broad scales (10s–100s of metres), primarily because of the high susceptibility of meiofauna to passive drift; small‐scale interactions between flow and substrata are also important, however, particularly at more localised (≤ metre) scales. At both scales, substratum particle size is important to interstitial‐dwelling fauna, influencing the probability of passive drift by meiofauna as well as local microhabitat conditions (e.g. dissolved oxygen; upwelling/downwelling in the hyporheic zone) and, thus, the small‐scale distribution among microhabitats.
• 5 In general, the processes governing the distribution of meiofauna at small scales cannot be separated entirely from those processes working at larger scales. A conceptual diagram is presented illustrating the relative importance of various factors in influencing the spatial patterns of meiofauna and over what scales these factors act.

     

Measuring Spider Richness: Effects of Different Sampling Methods and Spatial and Temporal Scales

Assessing the richness of invertebrate taxa to aid conservation and management requires a better understanding of the potential sources of error. Patterns of richness for heathland spiders at the species and family levels were compared across three sampling methods, four spatial scales, and monthly intervals (for 16 months). A total of 33 families and 130 species was collected: pitfall traps collected 94% of species, sweep net, 25%, and visual search, 41%. The sampling methods produced variable results. Pitfall trap and sweep net techniques identified significant, yet contrasting spatial differences in the number of families and species at one spatial scale. Pitfall trap data reflected strong temporal variation that influenced spatial patterns in richness (across one spatial scale for families and two for species). The use of broader temporal scales introduced a potential failure to detect significant differences in the richness of ground active spiders, and this risk varied spatially. The sweep net is not recommended for this habitat, although a method that targets the foliage is required for a more complete faunal assessment. Visual searches detected no significant patterns in richness, yet given its potential and increasing use for rapid biodiversity surveys, ways to improve sampling efficiency are suggested.     

Relative influence of variables at multiple spatial scales on stream macroinvertebrates in the Northern Lakes and Forest ecoregion, U.S.A.

• 1 We used 94 sites within the Northern Lakes and Forests ecoregion spanning Minnesota, Wisconsin and Michigan to identify environmental variables at the catchment, reach and riparian scales that influence stream macroinvertebrates. Redundancy analyses (RDA) found significantly influential variables within each scale and compared their relative importance in structuring macroinvertebrate assemblages.
• 2 Environmental variables included landcover, geology and groundwater delivery estimates at the catchment scale, water chemistry, channel morphology and stream habitat at the reach scale, and landcover influences at three distances perpendicular to the stream at the riparian scale. Macroinvertebrate responses were characterised with 22 assemblage attributes, and the relative abundance and presence/absence of 66 taxa.
• 3 Each scale defined macroinvertebrates along an erosional to depositional gradient. Wisconsin's macroinvertebrate index of biotic integrity, Ephemeroptera–Plecoptera–Trichoptera taxa and erosional taxa corresponded with forest streams, whereas organic pollution tolerant, Chironomidae and depositional taxa corresponded with wetland streams. Reach scale analyses defined the gradient similarly as dissolved oxygen and wide, shallow channels (erosional) opposed instream macrophytes and pool habitats (depositional). Riparian forests within 30 m of the stream coincided with an erosional assemblage and biotic integrity.
• 4 Next, we combined all significant environmental variables across scales to compare the relative influence of each spatial scale on macroinvertebrates. Partial RDA procedures described how much of the explained variance was attributable to each spatial scale and each interrelated scale combination.
• 5 Our results appeared consistent with the concept of hierarchical functioning of scale in which large‐scale variables restrict the potential for macroinvertebrate traits or taxa at smaller spatial scales. Catchment and reach variables were equally influential in defining assemblage attributes, whereas the reach scale was more influential in determining relative abundance and presence/absence.
• 6 Ultimately, comprehending the relative influence of catchment and reach scale properties in structuring stream biota will assist prioritising the scale at which to rehabilitate, manage and derive policies for stream ecosystem integrity.

     

The assembly of ecological communities inferred from taxonomic and functional composition

Among-site variation in metacommunities (beta diversity) is typically correlated with the distance separating the sites (spatial lag). This distance decay in similarity pattern has been linked to both niche-based and dispersal-based community assembly hypotheses. Here we show that beta diversity patterns in community composition, when supplemented with functional-trait information, can be used to diagnose assembly processes. First, using simulated data, we show how the relationship between distance decay patterns in taxonomic and functional measures of community composition can be used to predict the influence of a given trait on community assembly. We then use the patterns generated by the simulation as a template to show that the sorting of benthic macroinvertebrate metacommunities in headwater streams is likely influenced by different sets of functional traits at regional and local scales. We suggest that functional-trait databases and spatially referenced taxonomic surveys can be used to predict the spatial scales at which different aspects of interspecific functional variation are involved in niche-based community assembly while accounting for the influence of dispersal-based community assembly processes.     

Spatial constraint of peatland butterfly occurrences within a heterogeneous landscape

The occurrence of ten butterfly taxa (Clossiana eunomia dawsonii, Clossiana freija, Clossiana frigga, Clossiana titania, Coenonympha inornata, Erebia discoidalis, Incisalia augustinus, Lycaena dorcas, Lycaena epixanthe, Oeneis jutta) was analyzed within three acid peatland habitat types from the Lake Superior drainage basin of northwestern Wisconsin. Both first- (nearest-neighbor spatial analysis) and second-order (Ripley's K) spatial point process statistics were used to identify the extents over which each distribution pattern significantly deviated from random expectations. Versions of these tests were used that identified significant spatial pattern uncorrelated to habitat location and habitat preference. These analyses documented non-random occurrence patterns in seven species. Deviations from random were largely confined to two extents: <50 km and 70–100+ km. The majority of non-random patterns at <50 km extents were examples of aggregation, while the majority of non-random patterns noted at the 70–100+ km scale were examples of segregation. These results demonstrate that even for winged animals inside a limited landscape, spatially constrained processes can be important determinants of distribution. It is likely that metapopulation dynamics and dispersal limitation help explain why aggregation is dominant at small scales. The mechanisms underlying the predominance of segregation at large scales are less clear, but may be related to migration history and/or weak environmental gradients.     

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.