Species richness of woody plants in the landscapes of Central Spain: the role of management disturbances,environment and non‐stationarity

Questions: How important is management disturbance on gamma species richness of woody plants at intermediate landscape scales? How is species richness related to other climatic and biotic factors in the study area? How does the assumption of spatial stationarity affect assessment of relationships among species richness and explanatory variables (e.g. management, biotic and climatic factors) across extensive study areas? Location: Central Spain (regions of Castilla y León, Madrid and Castilla‐La Mancha). Scale: Extent: 150 000 km². Grain: 25 km² (5 × 5‐km cells). Methods: Information from 21 064 plots from the 3SNFI was used to evaluate richness of tree and shrub species at intermediate landscape scales. In addition to variables well known to explain biodiversity, e.g. environmental and biotic factors, effect of management treatments was evaluated by assessing clearcutting, selection cutting, stand improvement treatments and agrosilvopastoral systems (dehesas). Results from GWR techniques were compared with those from OLS regression. Results: Patterns of gamma species richness, although strongly affected by both environmental and biotic variables, were also significantly modified by management factors. Species richness increased with percentage of selection cutting stands and improvement treatments but decreased with percentage of clearcutting stands. Reduced species richness of woody plants was associated with agrosilvopastoral practices. Species richness for trees was closely related to basal area, annual precipitation and topographic complexity; species richness for shrubs was closely related to topographic complexity and agrosilvopastoral systems. Most relationships between species richness and environmental or biotic factors were non‐stationary. Relationships between species richness and management effects tended to be stationary, with a few exceptions. Conclusions: Landscape models of biodiversity in Central Spain were more informative when they accounted for effects of management practices, at least at intermediate scales. In the context of current rural abandonment, silvicultural disturbances of intermediate intensity increased gamma species richness of woody plants. Exclusion of factors such as agrosilvopastoral systems from models could have led to spurious relationships with other spatially co‐varying factors (e.g. summer precipitation). Patterns of spatial variation in relationships, provided by GWR models, allowed formulating hypotheses about potential ecological processes underlying them, beyond generalizations resulting from global (OLS) models.     

Connectivity, scale-dependence, and the productivity–diversity relationship

We surveyed freshwater ponds (localities) nested within watersheds (regions) to evaluate the relationship between productivity and animal species richness at different spatial scales. In watersheds where the ponds were relatively distant from one another (likely reducing the level of interpond dispersal of many organisms), we found a scale‐dependent productivity–diversity relationship; at local scales (among ponds), diversity was a hump‐shaped function of productivity, whereas at regional scales (among watersheds), diversity monotonically increased with productivity. Furthermore, this relationship emerged because there was a strong relationship between productivity and pond‐to‐pond species compositional differences. Alternatively, in watersheds where ponds were relatively close together (likely leading to higher rates of dispersal of many organisms), we found no scale‐dependence; diversity was a hump‐shaped function of productivity at both local and regional scales. Here, the relationship between species compositional dissimilarity and productivity was much weaker. We conclude that whether or not scale‐dependence is observed in productivity–diversity relationships will depend, at least in part, on the degree of connectivity among localities within regions.     

Environmental and spatial filters of zooplankton metacommunities in shallow pools in high‐elevation peatlands in the tropical Andes

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Untangling associations between chironomid taxa in Neotropical streams using local and landscape filters

1. Analyses of species association have major implications for selecting indicators for freshwater biomonitoring and conservation, because they allow for the elimination of redundant information and focus on taxa that can be easily handled and identified. These analyses are particularly relevant in the debate about using speciose groups (such as the Chironomidae) as indicators in the tropics, because they require difficult and time‐consuming analysis, and their responses to environmental gradients, including anthropogenic stressors, are poorly known. 2. Our objective was to show whether chironomid assemblages in Neotropical streams include clear associations of taxa and, if so, how well these associations could be explained by a set of models containing information from different spatial scales. For this, we formulated a priori models that allowed for the influence of local, landscape and spatial factors on chironomid taxon associations (CTA). These models represented biological hypotheses capable of explaining associations between chironomid taxa. For instance, CTA could be best explained by local variables (e.g. pH, conductivity and water temperature) or by processes acting at wider landscape scales (e.g. percentage of forest cover). 3. Biological data were taken from 61 streams in Southeastern Brazil, 47 of which were in well‐preserved regions, and 14 of which drained areas severely affected by anthropogenic activities. We adopted a model selection procedure using Akaike’s information criterion to determine the most parsimonious models for explaining CTA. 4. Applying Kendall’s coefficient of concordance, seven genera (Tanytarsus/Caladomyia, Ablabesmyia, Parametriocnemus, Pentaneura, Nanocladius, Polypedilum and Rheotanytarsus) were identified as associated taxa. The best‐supported model explained 42.6% of the total variance in the abundance of associated taxa. This model combined local and landscape environmental filters and spatial variables (which were derived from eigenfunction analysis). However, the model with local filters and spatial variables also had a good chance of being selected as the best model. 5. Standardised partial regression coefficients of local and landscape filters, including spatial variables, derived from model averaging allowed an estimation of which variables were best correlated with the abundance of associated taxa. In general, the abundance of the associated genera tended to be lower in streams characterised by a high percentage of forest cover (landscape scale), lower proportion of muddy substrata and high values of pH and conductivity (local scale). 6. Overall, our main result adds to the increasing number of studies that have indicated the importance of local and landscape variables, as well as the spatial relationships among sampling sites, for explaining aquatic insect community patterns in streams. Furthermore, our findings open new possibilities for the elimination of redundant data in the assessment of anthropogenic impacts on tropical streams.     

Biodiversity–productivity relationship in ponds: Community and metacommunity patterns along time and environmental gradients

Primary production correlates with diversity in various ways. These patterns may result from the interaction of various mechanisms related to the environmental context and the spatial and temporal scale of analysis. However, empirical evidence on diversity‐productivity patterns typically considers single temporal and spatial scales, and does not include the effect of environmental variables. In a metacommunity of macrophytes in ephemeral ponds, we analysed the diversity‐productivity relationship patterns in the field, the importance of the environmental variables of pond size and heterogeneity on such relationship, and the variation of these patterns at local (community level) and landscape scales (metacommunity level) across 52 ponds on twelve occasions, over five years (2005–2009). Combining all sampling dates, there were 377 ponds and 1954 sample‐unit observations. Vegetation biomass was used as a proxy for productivity, and biodiversity was represented by species richness, evenness, and their interaction. Environmental variables comprised pond area, depth and internal heterogeneity. Productivity and species richness were not directly related at the metacommunity level, and were positively related at the community level. Taking environmental variables into account revealed positive species richness‐productivity relationships at the metacommunity level and positive quadratic relationships at the community level. Productivity showed both positive and negative linear and nonlinear relationships with the size and heterogeneity of ponds. We found a weak relationship between productivity and evenness. The identity of variables associated with productivity changed between spatial scales and through time. The pattern of relationships between productivity and diversity depends on spatial scale and environmental context, and changes idiosyncratically through time within the same ecosystem. Thus, the diversity‐productivity relationship is not only a property of the study system, but also a consequence of environmental variations and the temporal and spatial scale of analysis.     

The roles of landscape context, niche breadth, and range boundaries in predicting species responses to habitat alteration

Extant species in human‐dominated landscapes differ in their sensitivity to habitat loss and fragmentation, although extinctions induced by environmental alteration reduce variation and result in a surviving subset of species with some degree of ‘resistance’. Here, we test the degree to which variable responses to habitat alteration are (1) essentially an inherent property of a taxon subject to constraints imposed by its geographical range, as suggested by Swihart et al. (2003), (2) a function of the landscape in which a species occurs, or (3) a function of spatial trends occurring on large scales. We used data collected on 33 vertebrate species during 2001–04 across the upper Wabash River basin, Indiana, in 35 square ‘landscapes’, each 23 km² in size. Six species of forest rodent, six species of grassland rodents, seven species of bats, eight species of aquatic turtles, and six species of amphibians were sampled at 504, 212, 590, 228, and 625 patches, respectively. The fraction of patches of primary habitat (e.g. forests for tree squirrels, wetlands for aquatic turtles) occupied by a target species was used as a response variable. On a basin‐wide scale, 47% of variation in proportional occupancy among species could be explained by taxon‐specific variables; occupancy rates were related positively to niche breadth and negatively to the proximity of a geographical range boundary. After controlling for species effects, landscape‐level occupancy rates varied significantly for 16 of 33 species, with variation partitioned among landscape variables alone (mean = 11% of variation), spatial trend variables alone (26%), and both variable sets jointly (8%). Among landscape variables, percentage forest cover positively affected occupancy rates of three bat species and a tree squirrel. Variation in occupancy rates among landscapes was consistent with large‐scale spatial trends for 13 species. Our findings demonstrate the general importance of niche breadth as a predictor of species responses to habitat alteration and highlight the importance of viewing the effects of habitat loss and fragmentation at multiple spatial scales.     

Communities in high definition: Spatial and environmental factors shape the micro-distribution of aquatic invertebrates

1. According to metacommunity theories, the structure of natural communities is the result of both environmental filtering and spatial processes, with their relative importance depending on factors including local habitat characteristics, functional features of organisms, and the spatial scale considered. However, few studies have explored environmental and spatial processes in riverine systems at local scales, explicitly incorporating spatial coordinates into multi-taxa distribution models. To address this gap, we conducted a small-scale study to discriminate between abiotic and biotic factors affecting the distribution of aquatic macroinvertebrates, applying metacommunity concepts.
2. We studied a mountain section in each of three perennial streams within the Po River Basin (northern Italy). We sampled macroinvertebrates both in summer and winter, using specific in situ 50-point random sampling grids. Environmental factors, including benthic organic matter (BOM), flow velocity, water depth, and substrate were recorded together with spatial coordinates for each sampling point. The relationships between community metrics (taxon richness, abundance, biomass, biomass–abundance ratio, and functional feeding groups) and explanatory variables (environmental and spatial) were assessed using generalised additive models. The influence of the explanatory variables on community structure was analysed with joint species distribution models.
3. Environmental variables—primarily BOM—were the main drivers affecting community metrics, whereas the effects of spatial variables varied among metrics, streams, and seasons. During summer, community structure was strongly affected by BOM and spatial position within the riverbed, the latter probably being a proxy for mass effects mediated by biotic and stochastic processes. In contrast, community structure was mainly shaped by hydraulic variables in winter.
4. Using macroinvertebrate communities as a model group, our results demonstrate that metacommunity concepts can explain small-scale variability in community structure. We found that both environmental filtering and biotic processes shape local communities, with the strength of these drivers depending on the season. These insights provide baseline knowledge that informs our understanding of ecological responses to environmental variability in contexts including restoration ecology, habitat suitability modelling, and biomonitoring.

     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

Food Webs in Relation to Variation in the Environment and Species Assemblage: A Multivariate Approach

The abiotic environment has strong influences on the growth, survival, behavior, and ecology of aquatic organisms. Biotic interactions and species life histories interact with abiotic factors to structure the food web. One measure of food-web structure is food-chain length. Several hypotheses predict a linear relationship between one environmental variable (e.g., disturbance or ecosystem size) and food-chain length. However, many abiotic and biotic variables interact in diverse ways to structure a community, and may affect other measures of food web structure besides food-chain length. This study took a multivariate approach to test the influence of several important environmental variables on four food-web characteristics measured in nine ponds along a hydroperiod gradient over two years. This approach allowed for testing the ecosystem size and dynamic constraints hypotheses while in context of other possibly interacting environmental variables. The relationship between amphibian and invertebrate communities and pond habitat variables was assessed to understand the underlying food-web structure. Hydroperiod and pond area had a strong influence on amphibian and invertebrate communities, trophic diversity and δ¹⁵N range. The range in δ¹³C values responded strongly to dissolved oxygen. Food-chain length responded to multiple environmental variables. Invertebrate and amphibian communities were structured by pond hydroperiod which in turn influenced the trophic diversity of the food web. The results of this study suggest food-chain length is influenced by environmental variation and species assemblage and that a multivariate approach may allow us to better understand the dynamics within and across aquatic food webs.     

Metacommunity organisation,spatial extent and dispersal in aquatic systems: patterns,processes and prospects

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Spatial distribution of subfossil Chironomidae in surface sediments of a large, shallow and hypertrophic lake (Taihu, SE China)

Spatial heterogeneity of benthic communities has clear implications for estimating lake production, biodiversity as well as identifying representative sites for palaeolimnological studies. This study investigates chironomid variability and the controlling factors (i.e., environmental and spatial variables) in surface sediments from Taihu Lake (2,338 km²), a hypertrophic lake in the Yangtze delta in eastern China. The spatial distribution of chironomids shows distinct heterogeneity. Microchironomus tabarui-type and Tanypus dominate the midge communities around the estuaries, while Cricotopus sylvestris-type and Polypedilum nubifer-type are the predominant taxa in the East Bays and the East Taihu Lake. Redundancy analysis was used for exploring the relationships between chironomid variability and environmental and spatial stressors. Four variables were identified as significant factors that influence chironomid community structures. The high nutrient concentrations around the estuarial areas favor the development of nutrient-tolerant taxa. Water depth-related oxygen depletion in the open lake during algae blooms prohibits the survival of many organisms, except for a few hypoxic-resistant species. High transparency in the East Bays and the East Taihu Lake indirectly creates a favorite microhabitat for macrophyte-associated chironomid species through aquatic plants. Space per se is a significant forcing factor for organism community and distribution at scales of >1,000 km². It might be important to consider spatial variables more explicitly in future studies of chironomids in large lakes where multiple stressors make the interactions within the ecosystem more complicated. This study aims to illustrate the ecological characteristics of specific chironomid taxa related to a “microecosystem” which is contributed by the multiple environmental gradients within a large lake, and to provide empirical support for interpretation of palaeochironomid data.     

Life history determines biogeographical patterns of soil bacterial communities over multiple spatial scales

The extent to which the distribution of soil bacteria is controlled by local environment vs. spatial factors (e.g. dispersal, colonization limitation, evolutionary events) is poorly understood and widely debated. Our understanding of biogeographic controls in microbial communities is likely hampered by the enormous environmental variability encountered across spatial scales and the broad diversity of microbial life histories. Here, we constrained environmental factors (soil chemistry, climate, above‐ground plant community) to investigate the specific influence of space, by fitting all other variables first, on bacterial communities in soils over distances from m to 10² km. We found strong evidence for a spatial component to bacterial community structure that varies with scale and organism life history (dispersal and survival ability). Geographic distance had no influence over community structure for organisms known to have survival stages, but the converse was true for organisms thought to be less hardy. Community function (substrate utilization) was also shown to be highly correlated with community structure, but not to abiotic factors, suggesting nonstochastic determinants of community structure are important Our results support the view that bacterial soil communities are constrained by both edaphic factors and geographic distance and further show that the relative importance of such constraints depends critically on the taxonomic resolution used to evaluate spatio‐temporal patterns of microbial diversity, as well as life history of the groups being investigated, much as is the case for macro‐organisms.     

Hierarchy theory reveals multiscale predictors of Arkansas darter (Etheostoma cragini) abundance in a Great Plains riverscape

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Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Unifying research on the fragmentation of terrestrial and aquatic habitats: patches,connectivity and the matrix in riverscapes

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Scale-dependent relationships between landscape structure and microclimate

This paper examined the scale-dependent relationships between landscape structure (e.g., slope, elevation, and overstory canopy coverage) and microclimate (e.g., air and soil temperatures and soil moisture) at different spatial scales along a 10050 m transect in the Southeastern Missouri Ozarks, USA. The landscape structure and microclimate variables were measured every 10 m along the transect during the growing season, June to September, 1996. We used the simple correlation analysis and "moving window" technique (no overlap between two adjacent windows) to examine the correlation coefficients between landscape structure and microclimate variables at scales or window sizes from 10 m to 2000 m. Because the sample size decreased rapidly in the "moving window" method, we also used the standardized cross-variogram to investigate the relationships between landscape structure and microclimate variables at a larger range of scales from 10 m to 8000 m. We found that the relationships between landscape structure and microclimate were apparently scale-dependent along the transect, suggesting the interactions between landscape structure and microclimate were stronger at some scales than others. The landscape structure variables were poor in explaining the variation of each microclimate variable at fine scales (e.g., 10 m). The correlations between elevation and microclimate variables were, in general, significantly improved with the increase of scales, while the improvement was less significant for slope and canopy coverage. Of the landscape structure variables, elevation, in general, had a higher correlation with the microclimate variables than slope and overstory canopy coverage at most scales examined. Our results suggest that small scales (e.g., < 100 m) are not suitable to study the relationships/interactions between landscape structure and microclimate and larger scales (e.g., > 500 m) are more appropriate though the relationships vary at the larger scales. Both the simple correlation analysis and standardized cross-variogram analysis were effective and, in general, consistent in characterizing the scale-dependent relationships between landscape structure and microclimate. Meanwhile, the standardized cross-variogram had the advantage to examine the relationships at large scales over the correlation analysis because the sample size reduced rapidly in the correlation analysis.     

Environmental variables interact across spatial scales to structure trichopteran assemblages in Ouachita Mountain rivers

An important goal in aquatic ecology is to determine the interacting variables that regulate community structure; however, complex biotic and abiotic interactions coupled with the significance of scale have confounded the interpretation of community data. We evaluated stream and riparian habitat features in southeastern Oklahoma, USA at a range of spatial scales from local, in-stream variables to large-scale, regional characteristics to address the following questions: (1) How much variation in trichopteran community composition can be attributed to local, regional, and spatial variables? and (2) What environmental variables are most important in determining trichopteran community structure? We collected data on caddisfly community structure, local and regional environmental variables, and spatial location on the landscape from 25 sites in four rivers. We analyzed these data using canonical correspondence analysis (CCA) and variation partitioning. Our analysis explained approximately 60% of the variation in caddisfly community composition. We found that local and regional environmental variables were near equal in importance in governing caddisfly communities, with each accounting for approximately a quarter of the explained variation. Although pure spatial variables were less important, the amount of variation shared among spatial variables and local and regional variables was substantial, indicating that biogeographic history is also key to understanding caddisfly distributions. We also found a strong influence of human landuse (i.e., percent of land in agriculture, distance to roads) on caddisfly community composition. Our study indicated that communities are influenced by factors across scales, and that bioassessments should focus on not only local habitat conditions, but also incorporate larger-scale factors.     

Different roles of environmental variables and spatial factors in structuring stream benthic diatom and macroinvertebrate in Yangtze River Delta,China

Exploring the relative contribution of spatial factors and environmental variables in shaping communities is of widespread interest in biodiversity conservation and environmental management. Stream communities are hierarchically regulated by environmental variables over multiple spatial scales, and the reaction of different organisms to stressors are still equivocal. We sampled both macroinvertebrates and diatom at 80 sites and additional 10 sites for macroinvertebrates, field measured and laboratory analyzed environmental variables, from the tributaries of Qiantang River, Yangtze River Delta China in 2011. We used PCNM (principal coordinates of neighbor matrices) to generate spatial predictors. We applied redundancy analysis and variation partitioning procedures to identify key spatial and environmental factors, and to quantify their relative roles in shaping diatom and macroinvertebrate assemblages. Our results demonstrated the role of spatial and environmental variables differed in shaping benthic diatom and macroinvertebrate. Diatom assemblage variations were better explained by spatial factors, however macroinvertebrate assemblage variations were better explained by environmental variables. In terms of environmental variables, catchment scale variables (e.g., land use estimators, land use diversity) played the primary role in determining the patterns of both diatom and macroinvertebrate assemblages, whereas the influence of reach-scale variables (e.g., pH, substrates, and nutrients) appeared less. However, nutrients were the stronger factors influencing benthic diatom, whereas physical habitat (e.g., substrates) played more important role than water chemistry in structuring macroinvertebrates. Our results provided more evidence to the incorporation of spatial factors interpreting spatial patterns of stream organisms, and highlighted the useful of multiple organisms and environmental variables at different spatial scales in diagnosing mechanism of stream degradation and in building a sound stream conditions monitoring program for Yangtze River Delta.     

Biodiversity: towards a unifying theme for river ecology

1. A broadened concept of biodiversity, encompassing spatio‐temporal heterogeneity, functional processes and species diversity, could provide a unifying theme for river ecology. 2. The theoretical foundations of stream ecology often do not reflect fully the crucial roles of spatial complexity and fluvial dynamics in natural river ecosystems, which has hindered conceptual advances and the effectiveness of efforts at conservation and restoration. 3. Inclusion of surface waters (lotic and lentic), subsurface waters (hyporheic and phreatic), riparian systems (in both constrained and floodplain reaches), and the ecotones between them (e.g. springs) as interacting components contributing to total biodiversity, is crucial for developing a holistic framework of rivers as ecosystems. 4. Measures of species diversity, including alpha, beta and gamma diversity, are a result of disturbance history, resource partitioning, habitat fragmentation and successional phenomena across the riverine landscape. A hierarchical approach to diversity in natural and altered river‐floodplain ecosystems will enhance understanding of ecological phenomena operating at different scales along multidimensional environmental gradients. 5. Re‐establishing functional diversity (e.g. hydrologic and successional processes) across the active corridor could serve as the focus of river conservation initiatives. Once functional processes have been reconstituted, habitat heterogeneity will increase, followed by corresponding increases in species diversity of aquatic and riparian biota.