The importance of spatial scale for trait–abundance relations

The concept of community assembly through trait‐based environmental filtering has played a key role in our understanding of how communities change over space and time, however, the importance of spatial scale in the filtering process remains unclear. We propose that different environmental filters may operate at different spatial scales, and that filters at finer scales would be nested within those acting at coarser scales. We tested for the existence of spatially nested sets of trait‐based filters in a temperate native grassland by applying the recently proposed maximum entropy (MaxEnt) approach to trait‐based community assembly, which we extend through a trait selection procedure. We found that different traits were important in influencing the abundances of species at the three different spatial scales examined (micro‐habitat, habitat, landscape), supporting the idea that trait based filtering processes operating at coarse spatial scales can be quite distinct from those operating at fine scales. Despite this result, we identified several traits which were frequently related to abundance at all spatial scales. Taken together, our results support the proposition that trait‐based environmental filters at finer spatial scales are nested within those operating at coarser scales. We compared our results to those obtained using a simpler trait‐by‐trait analytical approach (correlation analysis and MaxEnt on individual traits). The capacity for MaxEnt to incorporate multiple traits simultaneously provided unique insights into the important traits at each spatial scale and presents significant advantages over existing univariate and multivariate approaches.     

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

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

Integrating over uncertainty in spatial scale of response within multispecies occupancy models yields more accurate assessments of community composition

Species abundance and community composition are affected not only by the local environment, but also by broader landscape and regional context. Yet, determining the spatial scales at which landscapes affect species remains a persistent challenge, hindering our ability to understand how environmental gradients shape communities. This problem is amplified by rare species and imperfect species detection. Here, we present a Bayesian framework that allows uncertainty surrounding the ‘true’ spatial scale of species’ responses (i.e. changes in presence/absence) to be integrated directly into a community hierarchical model. This scale‐selecting multispecies occupancy model (ssMSOM) estimates the scale of response, and shows high accuracy and correct levels of uncertainty in parameter estimates across a broad range of simulation conditions. An ssMSOM can be run in a matter of minutes, as opposed to the many hours required to run normal multispecies occupancy models at all queried spatial scales, and then conduct model selection – a problem that up to now has prohibited scale of response from being rigorously evaluated in an occupancy framework. Alternatives to the ssMSOM, such as GLM‐based approaches frequently fail to detect the correct spatial scale and magnitude of response, and are often falsely confident by favoring the incorrect parameter estimates, especially as species’ detection probabilities deviate from perfect. We further show how trait information can be leveraged to understand how individual species’ scales of response vary within communities. Integrating spatial scale selection directly into hierarchical community models provides a means of formally testing hypotheses regarding spatial scales of response, and more accurately determining the environmental drivers that shape communities.     

Species turnover in lake littorals: spatial and temporal variation of benthic macroinvertebrate diversity and community composition

Aims Despite wide consensus that ecological patterns and processes should be studied at multiple spatial scales, the temporal component of diversity variation has remained poorly examined. Specifically, rare species may exhibit patterns of diversity variation profoundly different from those of dominant taxa. Location Southern Finland. Methods We used multiplicative partitioning of true diversities (species richness, Shannon diversity) to identify the most important scale(s) of variation of benthic macroinvertebrate communities across several hierarchical scales, from individual samples to multiple littorals, lakes and years. We also assessed the among‐scale variability of benthic macroinvertebrate community composition by using measures of between‐ and within‐group distances at hierarchical scales. Results On average, a single benthic sample contained 23% of the total regional macroinvertebrate species pool. For both species richness and Shannon diversity, beta‐diversity was clearly the major component of regional diversity, with within‐littoral beta‐diversity (β₁) being the largest component of gamma‐diversity. The interannual component of total diversity was small, being almost negligible for Shannon index. Among‐sample (within‐littoral) diversity was related to variation of substratum heterogeneity at the same scale. By contrast, only a small proportion of rare taxa was found in an average benthic sample. Thus, dominant species among lakes and years were about the same, whereas rare species were mostly detected in a few benthic samples in one lake (or year). For rare species, the temporal component of diversity was more important than spatial turnover at most scales. Main conclusions While individual species occurrences and abundances, particularly those of rare taxa, may vary strongly through space and time, patterns of dominance in lake littoral benthic communities are highly predictable. Consequently, many rare species will be missed in temporally restricted samples of lake littorals. In comprehensive biodiversity surveys, interannual sampling of littoral macroinvertebrate communities is therefore needed.     

A phylogenetic and trait‐based analysis of community assembly in a subtropical forest in central China

Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co‐occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait‐based tests to gain insights into community processes at four spatial scales in a large stem‐mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait‐based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co‐occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait‐based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.     

Stability and synchrony across ecological hierarchies in heterogeneous metacommunities: linking theory to data

Understanding stability across ecological hierarchies is critical for landscape management in a changing world. Recent studies showed that synchrony among lower‐level components is key to scaling temporal stability across two hierarchical levels, whether spatial or organizational. But an extended framework that integrates both spatial scale and organizational level simultaneously is required to clarify the sources of ecosystem stability at large scales. However, such an extension is far from trivial when taking into account the spatial heterogeneities in real‐world ecosystems. In this paper, we develop a partitioning framework that bridges variability and synchrony measures across spatial scales and organizational levels in heterogeneous metacommunities. In this framework, metacommunity variability is expressed as the product of local‐scale population variability and two synchrony indices that capture the temporal coherence across species and space, respectively. We develop an R function ‘var.partition’ and apply it to five types of desert plant communities to illustrate our framework and test how diversity shapes synchrony and variability at different hierarchical levels. As the observation scale increased from local populations to metacommunities, the temporal variability of plant productivity was reduced mainly by factors that decreased species synchrony. Species synchrony decreased from local to regional scales, and spatial synchrony decreased from species to community levels. Local and regional species diversity were key factors that reduced species synchrony at the two scales. Moreover, beta diversity contributed to decreasing spatial synchrony among communities. We conclude that our new framework offers a valuable toolbox for future empirical studies to disentangle the mechanisms and pathways by which ecological factors influence stability at large scales.     

Linking community assembly and structure across scales in a wild mouse parasite community

Understanding what processes drive community structure is fundamental to ecology. Many wild animals are simultaneously infected by multiple parasite species, so host–parasite communities can be valuable tools for investigating connections between community structures at multiple scales, as each host can be considered a replicate parasite community. Like free‐living communities, within‐host–parasite communities are hierarchical; ecological interactions between hosts and parasites can occur at multiple scales (e.g., host community, host population, parasite community within the host), therefore, both extrinsic and intrinsic processes can determine parasite community structure. We combine analyses of community structure and assembly at both the host population and individual scales using extensive datasets on wild wood mice (Apodemus sylvaticus) and their parasite community. An analysis of parasite community nestedness at the host population scale provided predictions about the order of infection at the individual scale, which were then tested using parasite community assembly data from individual hosts from the same populations. Nestedness analyses revealed parasite communities were significantly more structured than random. However, observed nestedness did not differ from null models in which parasite species abundance was kept constant. We did not find consistency between observed community structure at the host population scale and within‐host order of infection. Multi‐state Markov models of parasite community assembly showed that a host's likelihood of infection with one parasite did not consistently follow previous infection by a different parasite species, suggesting there is not a deterministic order of infection among the species we investigated in wild wood mice. Our results demonstrate that patterns at one scale (i.e., host population) do not reliably predict processes at another scale (i.e., individual host), and that neutral or stochastic processes may be driving the patterns of nestedness observed in these communities. We suggest that experimental approaches that manipulate parasite communities are needed to better link processes at multiple ecological scales.     

温带落叶阔叶林地表鞘翅目成虫小尺度空间格局动态分析

土壤动物空间格局是格局—过程以及生物多样性维持机制研究的重要基础,目前小尺度空间土壤动物空间格局动态特征仍不清楚。基于地统计空间分析方法,以鞘翅目成虫为研究对象,研究帽儿山温带落叶阔叶林小尺度空间(5m)地表鞘翅目成虫群落及类群的空间格局动态特征。结果表明:4次调查共捕获鞘翅目成虫11科、29类、1021只个体,调查月份鞘翅目成虫群落具有较强的时空变异性;Moran'sⅠ系数表明鞘翅目成虫群落和类群具有复杂的正的空间自相关性,其空间异质性可用球状、指数、高斯和线性模型进行拟合。这种空间异质性具有一定的时间变化特征,且这种空间分异是由随机性因素单一调控或结构性因素和随机性因素共同调控的结果;类群之间在多种尺度上表现为复杂的以负相关居多的空间关联性,这种空间关联性的形成主要是结构性因素或随机性因素单一调控的结果。本实验表明地表鞘翅目成虫群落在小尺度空间具有明显的空间异质性特征,这种空间异质性时间变异性较明显。     

Association of Host and Microbial Species Diversity across Spatial Scales in Desert Rodent Communities

Relationships between host and microbial diversity have important ecological and applied implications. Theory predicts that these relationships will depend on the spatio-temporal scale of the analysis and the niche breadth of the organisms in question, but representative data on host-microbial community assemblage in nature is lacking. We employed a natural gradient of rodent species richness and quantified bacterial communities in rodent blood at several hierarchical spatial scales to test the hypothesis that associations between host and microbial species diversity will be positive in communities dominated by organisms with broad niches sampled at large scales. Following pyrosequencing of rodent blood samples, bacterial communities were found to be comprised primarily of broad niche lineages. These communities exhibited positive correlations between host diversity, microbial diversity and the likelihood for rare pathogens at the regional scale but not at finer scales. These findings demonstrate how microbial diversity is affected by host diversity at different spatial scales and suggest that the relationships between host diversity and overall disease risk are not always negative, as the dilution hypothesis predicts.     

Scaling coexistence and assemblage patterns of desert small mammals

Scaling biodiversity patterns has been recognized lately as a very important issue in the search of global processes; however coexistence and assemblage patterns are typically approached at a single spatial scale. Here, we examined coexistence and co-occurrence patterns of desert small mammal communities across different spatial scales in the search of general community patterns. We sampled small mammals in Monte desert (Argentina, South America) for small spatial scales and reviewed published papers from other worldwide deserts for large spatial scale analyses. We used classic community estimators (Shannon, Richness), rank abundance curves and fitting distributions to analyze species coexistence and co-occurrence patterns. Assemblage patterns were analyzed evaluating nestedness across spatial scales and among deserts. Worldwide desert small mammal assemblages are characterized mainly by low species richness and high variation in species composition. The central Monte desert of Argentina showed a consistent assemblage pattern across spatial scales, with a generalist species being the most abundant and widely distributed, accompanied by other subordinate and more narrowly distributed species. All Monte desert communities were significantly nested, with nestedness increasing with scale from patch to regional. Assemblage and coexistence patterns were similar when comparing worldwide deserts despite differences in total richness and faunal singularity. The degree of nestedness varied among worldwide deserts; however all of them showed a consistent nested pattern. Differences in the degree of nestedness could be a result of different regulating factors depending on the desert and scale. These results highlight the importance of including multiscale approaches when dealing with processes that structure desert communities.     

Scale specific determinants of tree diversity in an old growth temperate forest in China

The major processes generating pattern in plant community composition depend upon the spatial scale and resolution of observation, therefore understanding the role played by spatial scale on species patterns is of major concern. In this study, we investigate how well environmental (topography and soil variables) and spatial variables explain variation in species composition in a 25-ha temperate forest in northeastern China. We used new variation partitioning approaches to discover the spatial scale (using multi-scale spatial PCNM variables) at which environmental heterogeneity and other spatially structured processes influence tree community composition. We also test the effect of changing grain of the study (i.e. quadrat size) on the variation partitioning results. Our results indicate that (1) species composition in the Changbai mixed broadleaf-conifer forest was controlled mainly by spatially structured soil variation at broad scales, while at finer scales most of the explained variation was of a spatial nature, pointing to the importance of biotic processes. (2) These results held at all sampling grains. However, reducing quadrat size progressively reduced both spatially and environmentally explained variance. This probably partly reflects increasing stochasticity in species abundances, and the smaller proportion of quadrats occupied by each species, when quadrat size is reduced. The results suggest that environmental heterogeneity (i.e. niche processes) and other biotic processes such as dispersal work together, but at different spatial scales, to build up diversity patterns.     

The combined effects of exogenous and endogenous variability on the spatial distribution of ant communities in a forested ecosystem (Hymenoptera: Formicidae)

Spatial patterns observed in ecosystems have traditionally been attributed to exogenous processes. Recently, ecologists have found that endogenous processes also have the potential to create spatial patterns. Yet, relatively few studies have attempted to examine the combined effects of exogenous and endogenous processes on the distribution of organisms across spatial and temporal scales. Here we aim to do this, by investigating whether spatial patterns of under-story tree species at a large spatial scale (18 ha) influences the spatial patterns of ground foraging ant species at a much smaller spatial scale (20 m by 20 m). At the regional scale, exogenous processes (under-story tree community) had a strong effect on the spatial patterns in the ground-foraging ant community. We found significantly more Camponotus noveboracensis, Formica subsericae, and Lasius alienus species in black cherry (Prunis serotine Ehrh.) habitats. In witch-hazel (Hamamelis virginiana L.) habitats, we similarly found significantly more Myrmica americana, Formica fusca, and Formica subsericae. At smaller spatial scales, we observed the emergence of mosaic ant patches changing rapidly in space and time. Our study reveals that spatial patterns are the result of both exogenous and endogenous forces, operating at distinct scales.     

A multiscale analysis of herbaceous species richness in a Mediterranean ecosystem

Aims Studies of species distribution patterns traditionally have been conducted at a single scale, often overlooking species–environment relationships operating at finer or coarser scales. Testing diversity-related hypotheses at multiple scales requires a robust sampling design that is nested across scales. Our chief motivation in this study was to quantify the contributions of different predictors of herbaceous species richness at a range of local scales.Methods Here, we develop a hierarchically nested sampling design that is balanced across scales, in order to study the role of several environmental factors in determining herbaceous species distribution at various scales simultaneously. We focus on the impact of woody vegetation, a relatively unexplored factor, as well as that of soil and topography. Light detection and ranging (LiDAR) imaging enabled precise characterization of the 3D structure of the woody vegetation, while acoustic spectrophotometry allowed a particularly high-resolution mapping of soil CaCO 3 and organic matter contents.Important findings We found that woody vegetation was the dominant explanatory variable at all three scales (10, 100 and 1000 m 2), accounting for more than 60% of the total explained variance. In addition, we found that the species richness–environment relationship was scale dependent. Many studies that explicitly address the issue of scale do so by comparing local and regional scales. Our results show that efforts to conserve plant communities should take into account scale dependence when analyzing species richness–environment relationships, even at much finer resolutions than local vs. regional. In addition, conserving heterogeneity in woody vegetation structure at multiple scales is a key to conserving diverse herbaceous communities.     

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.     

Spatial scale and benthic community organisation in the littoral zones of large oligotrophic lakes: potential for cross-scale interactions

1. We studied the spatial organisation of littoral, benthic invertebrate communities in two large oligotrophic lakes with very extensive heterogeneous littoral zones, and report that littoral communities show significant variation in their structure at multiple spatial scales. 2. At coarser scales, littoral zone topography, riparian inputs and exposure may be important drivers of community spatial structure. At finer scales, the composition of substrates, particularly macrophytes and inorganic sediments, may be drivers of community spatial structure. 3. The effects of substrates on community spatial patterns were non‐additive across scales; community‐habitat correspondence became very noisy when we attempted to extrapolate results throughout the littoral. We present some evidence to suggest that this non‐additivity is because of the interaction among structuring processes that operate at different scales.     

Geographical ecology of the palms (Arecaceae): determinants of diversity and distributions across spatial scales

Background

The palm family occurs in all tropical and sub-tropical regions of the world. Palms are of high ecological and economical importance, and display complex spatial patterns of species distributions and diversity.

Scope

This review summarizes empirical evidence for factors that determine palm species distributions, community composition and species richness such as the abiotic environment (climate, soil chemistry, hydrology and topography), the biotic environment (vegetation structure and species interactions) and dispersal. The importance of contemporary vs. historical impacts of these factors and the scale at which they function is discussed. Finally a hierarchical scale framework is developed to guide predictor selection for future studies.

Conclusions

Determinants of palm distributions, composition and richness vary with spatial scale. For species distributions, climate appears to be important at landscape and broader scales, soil, topography and vegetation at landscape and local scales, hydrology at local scales, and dispersal at all scales. For community composition, soil appears important at regional and finer scales, hydrology, topography and vegetation at landscape and local scales, and dispersal again at all scales. For species richness, climate and dispersal appear to be important at continental to global scales, soil at landscape and broader scales, and topography at landscape and finer scales. Some scale–predictor combinations have not been studied or deserve further attention, e.g. climate on regional to finer scales, and hydrology and topography on landscape and broader scales. The importance of biotic interactions – apart from general vegetation structure effects – for the geographic ecology of palms is generally underexplored. Future studies should target scale–predictor combinations and geographic domains not studied yet. To avoid biased inference, one should ideally include at least all predictors previously found important at the spatial scale of investigation.     

Hierarchical Dynamics of Ecological Communities: Do Scales of Space and Time Match?

Theory posits that community dynamics organize at distinct hierarchical scales of space and time, and that the spatial and temporal patterns at each scale are commensurate. Here we use time series modeling to investigate fluctuation frequencies of species groups within invertebrate metacommunities in 26 boreal lakes over a 20-year period, and variance partitioning analysis to study whether species groups with different fluctuation patterns show spatial signals that are commensurate with the scale-specific fluctuation patterns identified. We identified two groups of invertebrates representing hierarchically organized temporal dynamics: one species group showed temporal variability at decadal scales (slow patterns of change), whilst another group showed fluctuations at 3 to 5-year intervals (faster change). This pattern was consistently found across all lakes studied. A spatial signal was evident in the slow but not faster-changing species groups. As expected, the spatial signal for the slow-changing group coincided with broad-scale spatial patterns that could be explained with historical biogeography (ecoregion delineation, and dispersal limitation assessed through a dispersal trait analysis). In addition to spatial factors, the slow-changing groups correlated with environmental variables, supporting the conjecture that boreal lakes are undergoing environmental change. Taken together our results suggest that regionally distinct sets of taxa, separated by biogeographical boundaries, responded similarly to broad-scale environmental change. Not only does our approach allow testing theory about hierarchically structured space-time patterns; more generally, it allows assessing the relative role of the ability of communities to track environmental change and dispersal constraints limiting community structure and biodiversity at macroecological scales.     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

Tropical paradox: a multi-scale analysis of the invasion paradox within Miami Rock Ridge tropical hardwood hammocks

The invasion paradox describes the scale dependence of native-exotic richness relationships (NERRs), where NERRs are negative at neighborhood scales and positive at landscape scales. However, a lack of tropical surveys and past failures to isolate potential confounding variables contribute to significant gaps in our understanding of the processes producing these patterns. We surveyed the vascular flora of 13 tropical hardwood hammocks for community characteristics (e.g., native and exotic species richness, vegetative cover) with a hierarchical sampling design. Using model selection, we determined which variables best predicted patterns of exotic species richness at each spatial scale of consideration. We found that native and exotic species richness were positively correlated at neighborhood scales, but negatively correlated at landscape scales. The latter result stands in stark opposition to the patterns published in the literature thus far. We found that natural disturbance history (as approximated by vegetative cover) was positively correlated with exotic species richness at intermediate and landscape scales only. Overall, hammock identity was the most important factor driving exotic species richness patterns at all spatial scales. Hammocks with highly-disturbed hydrologies, brought about by water management, had fewer native species and more exotic species than hammocks with more natural hydrological conditions. Our results are among the first from examination of subtropical communities, and may support the hypothesis that tropical and subtropical communities are subject to more intense biotic interactions. However, given our unique sampling design, our results do not reject the hypothesis that environmental heterogeneity drives the relationship between native and exotic species richness patterns.     

Spatial variability,structure and composition of crustose algal communities in Diadema africanum barrens

Crustose algal communities were studied in Diadema africanum urchin barrens around Tenerife (Canary Islands, NE Atlantic). A hierarchical nested sampling design was used to study patterns of community variability at different spatial scales (sectors, three sides of the island; sites within each sector, 5–10 km apart; stations within each site, 50–100 m apart). Although noncrustose species contributed the most to community richness, cover was dominated by crustose forms, like the coralline algae Hydrolithon farinosum, H. samoënse, H. onkodes, Neogoniolithon orotavicum and N. hirtum, and the phaeophycean Pseudolithoderma adriaticum. The structure of these communities showed high spatial variability, and we found differences in the structure of urchin barrens when compared across different spatial scales. Multivariate analysis showed that variability in community structure was related to the five environmental variables studied (wave exposure, urchin density, substrate roughness, productivity and depth). Wave exposure was the variable that contributed most to community variability, followed by urchin density and substrate roughness. Productivity and depth had limited influence. The effects of these variables differed depending on the spatial scale; wave exposure and productivity were the main variables influencing community changes at the largest scale (between different sectors of the island), while D. africanum density, roughness and depth were the most influential at medium and small scales.