Plant diversity partitioning in Mediterranean croplands: effects of farming intensity, field edge, and landscape context

Farmland biodiversity is affected by factors acting at various spatial scales. However, most studies to date have focused on the field or farm scales that only account for local (alpha) diversity, and these may underestimate the contribution of other diversity components (beta diversity) to total (gamma) farmland diversity. In this work, we aimed to identify the most suitable management options and the scale at which they should be implemented to maximize benefits for diversity. We used a multi-scale additive partitioning approach, with data on plant diversity from 640 plots in 32 cereal crop fields from three agricultural regions of central Spain that differed in landscape configuration. We analyzed the relative contribution to overall plant diversity of different diversity components at various spatial scales and how these diversity components responded to a set of local (application of agri-environment schemes [AES] and position within the field) and landscape (field size and landscape connectivity and composition) factors. Differences in species composition among regions and then among fields within regions contributed most to overall plant diversity. Positive edge effects were found on all diversity components at both the field- and regional scales, whereas application of AES benefited all diversity components only at the field scale. Landscape factors had strong influences on plant diversity, especially length of seminatural boundaries, which increased species richness at both the field and the regional scales. In addition, positive effects of percentage of nonproductive land-uses in the landscape were found on all diversity components at the regional scale. Results showed that components that contributed most to overall plant diversity were not benefited by current AES. We conclude that agri-environmental policies should incorporate and prioritize measures aimed at the maintenance of seminatural boundaries and patches of nonproductive habitats within agricultural landscapes, through landscape planning, cross-compliance, or high nature-value farmland programs. These options will help to conserve overall plant diversity at regional scales, as well as the spillover of plant species from such seminatural elements into crops, especially in Mediterranean areas that still harbor extensive farming and relatively complex landscapes.     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.     

Habitat biodiversity as a determinant of fish community structure on coral reefs

Increased habitat diversity is often predicted to promote the diversity of animal communities because a greater variety of habitats increases the opportunities for species to specialize on different resources and coexist. Although positive correlations between the diversities of habitat and associated animals are often observed, the underlying mechanisms are only now starting to emerge, and none have been tested specifically in the marine environment. Scleractinian corals constitute the primary habitat-forming organisms on coral reefs and, as such, play an important role in structuring associated reef fish communities. Using the same field experimental design in two geographic localities differing in regional fish species composition, we tested the effects of coral species richness and composition on the diversity, abundance, and structure of the local fish community. Richness of coral species overall had a positive effect on fish species richness but had no effect on total fish abundance or evenness. At both localities, certain individual coral species supported similar levels of fish diversity and abundance as the high coral richness treatments, suggesting that particular coral species are disproportionately important in promoting high local fish diversity. Furthermore, in both localities, different microhabitats (coral species) supported very different fish communities, indicating that most reef fish species distinguish habitat at the level of coral species. Fish communities colonizing treatments of higher coral species richness represented a combination of those inhabiting the constituent coral species. These findings suggest that mechanisms underlying habitat-animal interaction in the terrestrial environment also apply to marine systems and highlight the importance of coral diversity to local fish diversity. The loss of particular key coral species is likely to have a disproportionate impact on the biodiversity of associated fish communities.     

Partitioning of \alpha and \beta diversity using hierarchical Bayesian modeling of species distribution and abundance

Diversity partitioning is becoming widely used to decompose the total number of species recorded in an area or region \((\gamma )\) into the average number of species within samples \((\alpha )\) and the average difference in species composition \((\beta )\) among samples. Single-value metrics of \(\alpha \) and \(\beta \) diversity are popular because they may be applied at multiple scales and because of their ease in computation and interpretation. Studies thus far, however, have emphasized observed diversity components or comparisons to randomized, null distributions. In addition, prediction of \(\alpha \) and \(\beta \) components using environmental or spatial variables has been limited to more extensive data sets because multiple samples are required to estimate single \(\alpha \) and \(\beta \) components. Lastly, observed diversity components do not incorporate variation in detection probabilities among species or samples. In this study, we used hierarchical Bayesian models of species abundances to provide predictions of \(\alpha \) and \(\beta \) components in species richness and composition using environmental and spatial variables. We illustrate our approach using butterfly data collected from 26 grassland remnants to predict spatially nested patterns of \(\alpha \) and \(\beta \) based on the predicted counts of butterflies. Diversity partitioning using a Bayesian hierarchical model incorporated variation in detection probabilities by butterfly species and habitat patches, and provided prediction intervals for \(\alpha \) and \(\beta \) components using environmental and spatial variables.     

Modeling the beta diversity of coral reefs

Quantifying the beta diversity (species replacement along spatiotemporal gradients) of ecosystems is important for understanding and conserving patterns of biodiversity. However, virtually all studies of beta diversity focus on one-dimensional transects orientated along a specific environmental gradient that is defined a priori. By ignoring a second spatial dimension and the associated changes in species composition and environmental gradients, this approach may provide limited insight into the full pattern of beta diversity. Here, we use remotely sensed imagery to quantify beta diversity continuously, in two dimensions, and at multiple scales across an entire tropical marine seascape. We then show that beta diversity can be modeled (0.852 > or = r2 > or = 0.590) at spatial scales between 0.5 and 5.0 km2, using the environmental variables of mean and variance of depth and wave exposure. Beta diversity, quantified within a "window" of a given size, is positively correlated to the range of environmental conditions within that window. For example, beta diversity increases with increasing variance of depth. By analyzing such relationships across seascapes, this study provides a framework for a range of disparate coral reef literature including studies of zonation, diversity, and disturbance. Using supporting evidence from soft-bottom communities, we hypothesize that depth will be an important variable for modeling beta diversity in a range of marine systems. We discuss the implications of our results for the design of marine reserves.     

Natural and anthropogenic influences on the diversity structure of reef fish communities in the Tuamotu Archipelago (French Polynesia)

Despite the rapid rate of human-induced species losses, the relative influence of natural and anthropogenic factors on the functional diversity of species assemblages remains unknown for most ecosystems. A model was previously developed to predict the diversity structure of coral reef fish assemblages in 10 atolls of low human pressure and contrasting morphology of the Tuamotu Archipelago (French Polynesia). This existing model predicted smoothed histograms (spectra) of species richness according to size classes, diet classes and life-history classes of fish assemblages using a combination of environmental characteristics at different spatial scales. The present study applied the model to Tikehau, another atoll of the same archipelago where commercial fishing is practiced and where the same sampling strategy was reproduced. Significant differences appeared between predicted and observed species richness in several size, diet and life-history classes of fish assemblages in Tikehau. Two parameters which were not accounted for in the initial model, i.e. fishing pressure and atoll position within the archipelago, explained together 63% of variance in model residuals, >60% being explained by fishing pressure only. The respective effects of fishing and atoll position on the diversity of coral reef fish assemblages are discussed, with the potential of such modelling approach to assess the relative importance of factors affecting functional diversity within communities.     

Multi-scale marine biodiversity patterns inferred efficiently from habitat image processing

Cost-effective proxies of biodiversity and species abundance, applicable across a range of spatial scales, are needed for setting conservation priorities and planning action. We outline a rapid, efficient, and low-cost measure of spectral signal from digital habitat images that, being an effective proxy for habitat complexity, correlates with species diversity and requires little image processing or interpretation. We validated this method for coral reefs of the Great Barrier Reef (GBR), Australia, across a range of spatial scales (1 m to 10 km), using digital photographs of benthic communities at the transect scale and high-resolution Landsat satellite images at the reef scale. We calculated an index of image-derived spatial heterogeneity, the mean information gain (MIG), for each scale and related it to univariate (species richness and total abundance summed across species) and multivariate (species abundance matrix) measures of fish community structure, using two techniques that account for the hierarchical structure of the data: hierarchical (mixed-effect) linear models and distance-based partial redundancy analysis. Over the length and breadth of the GBR, MIG alone explained up to 29% of deviance in fish species richness, 33% in total fish abundance, and 25% in fish community structure at multiple scales, thus demonstrating the possibility of easily and rapidly exploiting spatial information contained in digital images to complement existing methods for inferring diversity and abundance patterns among fish communities. Thus, the spectral signal of unprocessed remotely sensed images provides an efficient and low-cost way to optimize the design of surveys used in conservation planning. In data-sparse situations, this simple approach also offers a viable method for rapid assessment of potential local biodiversity, particularly where there is little local capacity in terms of skills or resources for mounting in-depth biodiversity surveys.     

Acanthochromis polyacanthus,a fish lacking larval dispersal,has genetically differentiated populations at local and regional scales on the Great Barrier Reef

Acanthochromis Gill is a monotypic genus within the damselfish family Pomacentridae, erected for an unusual species [A. polyacanthus (Bleeker)] that uniquely lacks larval dispersal. Instead, offspring are reared in the parental territory, in the manner of cichlids, and fledged into the surrounding habitat. Phenotypic and genotypic variation was surveyed on the basis of body colouration and 7 polymorphic loci in 19 populations from 5 regions of the central and southern Great Barrier Reef (GBR). Variation in both characters was found at regional and local scales. Two colour morphs were recognised: a bicoloured morph from the three northern regions and a uniform dark morph from the two southern regions. Isozyme analysis showed a similar pattern with greatest variation between the different morphs, but also with significant variation at both regional and local scales within morphotypes. Heterozygosity was maximal in the central populations, which, together with other measures of variability, suggests a mixing of separate gene pools in this region and denies species status to the two morphotypes despite numerous fixed differences in allele frequencies between the most distant populations. The presence of fixed differences in multiple alleles between populations separated by 1000 km indicates negligible gene flow over such distances and long isolation of these gene pools. These patterns may reflect recolonisation of the GBR after the last sea-level rise by fish from two stocks. Founder effects and random drift in small populations after colonisation are probably the major sources of the local and regional variations observed at smaller spatial scales. This diversity has been maintained among populations at all scales by the very low levels of gene flow possible without an effective strategy for larval dispersal between coral reefs.     

Species dispersal rates alter diversity and ecosystem stability in pond metacommunities

Metacommunity theory suggests that relationships between diversity and ecosystem stability can be determined by the rate of species dispersal among local communities. The predicted relationships, however, may depend upon the relative strength of local environmental processes and disturbance. Here we evaluate the role of dispersal frequency and local predation perturbations in affecting patterns of diversity and stability in pond plankton metacommunities. Pond metacommunities were composed of three mesocosm communities: one of the three communities maintained constant "press" predation from a selective predator, bluegill sunfish (Lepomis macrochirus); the second community maintained "press" conditions without predation; and the third community experienced recurrent "pulsed" predation from bluegill sunfish. The triads of pond communities were connected at either no, low (0.7%/d), or high (20%/d) planktonic dispersal. Richness and composition of zooplankton and stability of plankton biomass and ecosystem productivity were measured at local and regional spatial scales. Dispersal significantly affected diversity such that local and regional biotas at the low dispersal rate maintained the greatest number of species. The unimodal local dispersal-diversity relationship was predator-dependent, however, as selective press predation excluded species regardless of dispersal. Further, there was no effect of dispersal on beta diversity because predation generated local conditions that selected for distinct community assemblages. Spatial and temporal ecosystem stability responded to dispersal frequency but not predation. Low dispersal destabilized the spatial stability of producer biomass but stabilized temporal ecosystem productivity. The results indicate that selective predation can prevent species augmentation from mass effects but has no apparent influence on stability. Dispersal rates, in contrast, can have significant effects on both species diversity and ecosystem stability at multiple spatial scales in metacommunities.     

Scale-dependent responses of plant biodiversity to nitrogen enrichment

Experimental studies demonstrating that nitrogen (N) enrichment reduces plant diversity within individual plots have led to the conclusion that anthropogenic N enrichment is a threat to global biodiversity. These conclusions overlook the influence of spatial scale, however, as N enrichment may alter beta diversity (i.e., how similar plots are in their species composition), which would likely alter the degree to which N-induced changes in diversity within localities translate to changes in diversity at larger scales that are relevant to policy and management. Currently, it is unclear how N enrichment affects biodiversity at scales larger than a small plot. We synthesized data from 18 N-enrichment experiments across North America to examine the effects of N enrichment on plant species diversity at three spatial scales: small (within plots), intermediate (among plots), and large (within and among plots). We found that N enrichment reduced plant diversity within plots by an average of 25% (ranging from a reduction of 61% to an increase of 5%) and frequently enhanced beta diversity. The extent to which N enrichment altered beta diversity, however, varied substantially among sites (from a 22% increase to an 18% reduction) and was contingent on site productivity. Specifically, N enrichment enhanced beta diversity at low-productivity sites but reduced beta diversity at high-productivity sites. N-induced changes in beta diversity generally reduced the extent of species loss at larger scales to an average of 22% (ranging from a reduction of 54% to an increase of 18%). Our results demonstrate that N enrichment often reduces biodiversity at both local and regional scales, but that a focus on the effects of N enrichment on biodiversity at small spatial scales may often overestimate (and sometimes underestimate) declines in regional biodiversity by failing to recognize the effects of N on beta diversity.     

Benefits of Conservation of Plant Genetic Diversity to Arthropod Diversity

Abstract:  We argue that the genetic diversity of a dominant plant is important to the associated dependent community because dependent species such as herbivores are restricted to a subset of genotypes in the host-plant population. For plants that function as habitat, we predicted that greater genetic diversity in the plant population would be associated with greater diversity in the dependent arthropod community. Using naturally hybridizing cottonwoods (  Populus spp.) in western North America as a model system, we tested the general hypothesis that arthropod alpha (within cross-type richness) and beta (among cross-type composition) diversities are correlated with cottonwood cross types from local to regional scales. In common garden experiments and field surveys, leaf-modifying arthropod richness was significantly greater on either the F₁ (1.54 times) or backcross (1.46 times) hybrid cross types than on the pure broadleaf cross type (  P. deltoides Marshall or P. fremontii Watson). Composition was significantly different among three cross types of cottonwoods at all scales. Within a river system, cottonwood hybrid zones had 1.49 times greater richness than the broadleaf zone, and community composition was significantly different between each parental zone and the hybrid zone, demonstrating a hierarchical concentration of diversity. Overall, the habitats with the highest cottonwood cross-type diversity also had the highest arthropod diversity. These data show that the genetics of habitat is an important conservation concept and should be a component of conservation theory.     

Spatial Components of Bird Diversity in the Andes of Colombia: Implications for Designing a Regional Reserve System

Abstract:  Beta diversity, or the turnover in species composition among sampling sites in a region, is an important criterion for obtaining adequate representation of regional biodiversity in systems of protected areas. Recently, the additive model for partitioning regional (gamma) diversity (in opposition to the multiplicative model) has been proposed because it allows a direct measure of the contribution of beta diversity to gamma diversity. We determined avian beta diversity along latitudinal (among neighboring river drainages) and elevational axes in a 1347-km² region on the western slope of the Central Cordillera of the Colombian Andes, where a regional system of protected areas is being designed. We then compared avian beta diversity between sites based on rapid versus long-term (>1 year) inventories and between fragmented sites versus continuous forest. Overall, beta diversity represented 63.1% of gamma diversity among 16 sites. Elevational differences in species composition accounted for 43.3% of regional diversity, whereas differences among drainages accounted for 19.8%. A complementary cluster analysis showed that sites grouped by elevational zones. Rapid inventories overestimated beta diversity because of sampling effects, but the effect was biologically small. Estimators of species richness derived from species accumulation curves provided a useful alternative to compensate for undersampling in short-term surveys. Forest fragmentation increased beta diversity because of differential local extinction of populations. Nevertheless, in our region, forest fragments contributed to gamma diversity because they contained complementary sets of species. More importantly, they contained populations of special-interest species. Although the region is relatively small, our analyses indicate that spatial differentiation of the biota is an important factor for deciding number and location of protected areas in the Andean region.     

Hydrologic connection between ponds positively affects macrophyte alpha and gamma diversity but negatively affects beta diversity

Connections between habitat patches can positively influence the number of species in respective patches, providing a basis for preferentially conserving interconnected patches. However, from a regional perspective, it is not known whether conserving multiple sets of interconnected habitat patches would include more species (i.e., show higher gamma diversity) than conserving multiple, unconnected, solitary patches. We studied aquatic macrophytes in 15 sets of unidirectionally interconnected ponds and 19 unconnected ponds and also tested whether alpha and beta diversity, expressed as the number of species and dissimilarity in species composition, respectively, differed between connected and unconnected ponds. We found that gamma diversity was higher in connected ponds than in unconnected ponds, even after controlling for surface area. This resulted from a higher alpha diversity in connected ponds, despite lower beta diversity. These results suggest that connections between habitat patches positively influence diversity at both local and regional scales. When the total surface area available for conservation is limited, interconnected habitat patches should be preferentially conserved.     

Aggregation influences coral species richness at multiple spatial scales

The spatial dispersion of individuals across multiple spatial scales can significantly influence biodiversity patterns. Here we characterize the dispersion of corals in reef assemblages distributed across a 10000-km longitudinal biodiversity gradient from Indonesia to the Society Islands, using a multiscale sampling design. Our results indicate that most coral species were aggregated among 10-m transect samples across this vast distance. Using observed and randomized species sampling curves, we show that aggregation reduced the number of species per transect, site, and island sample on average by 13-27%. Across site, island, and regional scales, aggregation also reduced the area under species sampling curves by an average of 2.7-6.5%. The level of aggregation was relatively constant across spatial scales within regions and did not vary among habitats. However, there was significant variation among regions using transect samples across individual sites. Specifically, aggregation reduced the species richness per transect and the area under species sampling curves nearly twice as much in the Indonesian biodiversity hotspot than in the Society Islands. As a significant component of the spatial structure of coral assemblages, aggregation should be integrated into our understanding of coral community dynamics and the development of conservation strategies designed to protect these communities.     

Interpreting beta‐diversity components over time to conserve metacommunities in highly dynamic ecosystems

The concept of metacommunity (i.e., a set of local communities linked by dispersal) has gained great popularity among community ecologists. However, metacommunity research mostly addresses questions on spatial patterns of biodiversity at the regional scale, whereas conservation planning requires quantifying temporal variation in those metacommunities and the contributions that individual (local) sites make to regional dynamics. We propose that recent advances in diversity‐partitioning methods may allow for a better understanding of metacommunity dynamics and the identification of keystone sites. We used time series of the 2 components of beta diversity (richness and replacement) and the contributions of local sites to these components to examine which sites controlled source‐sink dynamics in a highly dynamic model system (an intermittent river). The relative importance of the richness and replacement components of beta diversity fluctuated over time, and sample aggregation led to underestimation of beta diversity by up to 35%. Our literature review revealed that research on intermittent rivers would benefit greatly from examination of beta‐diversity components over time. Adequately appraising spatiotemporal variability in community composition and identifying sites that are pivotal for maintaining biodiversity at the landscape scale are key needs for conservation prioritization and planning. Thus, our framework may be used to guide conservation actions in highly dynamic ecosystems when time‐series data describing biodiversity across sites connected by dispersal are available.     

Area and mammalian elevational diversity

Elevational gradients hold enormous potential for understanding general properties of biodiversity. Like latitudinal gradients, the hypotheses for diversity patterns can be grouped into historical explanations, climatic drivers, and spatial hypotheses. The spatial hypotheses include the species-area effect and spatial constraint (mid-domain effect null models). I test these two spatial hypotheses using regional diversity patterns for mammals (non-volant small mammals and bats) along 34 elevational gradients spanning 24.4 degrees S-40.4 degrees N latitude. There was high variability in the fit to the species-area hypothesis and the mid-domain effect. Both hypotheses can be eliminated as primary drivers of elevational diversity. Area and spatial constraint both represent sources of error rather than mechanisms underlying these mammalian diversity patterns. Similar results are expected for other vertebrate taxa, plants, and invertebrates since they show comparable distributions of elevational diversity patterns to mammalian patterns.     

Integrating larval connectivity into the marine conservation decision-making process across spatial scales

Larval dispersal connectivity is typically integrated into spatial conservation decisions at regional or national scales, but implementing agencies struggle with translating these methods to local scales. We used larval dispersal connectivity at regional (hundreds of kilometers) and local (tens of kilometers) scales to aid in design of networks of no-take reserves in Southeast Sulawesi, Indonesia. We used Marxan with Connectivity informed by biophysical larval dispersal models and remotely sensed coral reef habitat data to design marine reserve networks for 4 commercially important reef species across the region. We complemented regional spatial prioritization with decision trees that combined network-based connectivity metrics and habitat quality to design reserve boundaries locally. Decision trees were used in consensus-based workshops with stakeholders to qualitatively assess site desirability, and Marxan was used to identify areas for subsequent network expansion. Priority areas for protection and expected benefits differed among species, with little overlap in reserve network solutions. Because reef quality varied considerably across reefs, we suggest reef degradation must inform the interpretation of larval dispersal patterns and the conservation benefits achievable from protecting reefs. Our methods can be readily applied by conservation practitioners, in this region and elsewhere, to integrate connectivity data across multiple spatial scales.     

Extreme genetic diversity and temporal rather than spatial partitioning in a widely distributed coral reef fish

Mitochondrial control region (HVR-1) sequences were used to identify patterns of genetic structure and diversity in Naso vlamingii, a widespread coral reef fish with a long evolutionary history. We examined 113 individuals from eight locations across the Indo-Pacific Ocean. Our aims were to determine the spatial scale at which population partitioning occurred and then to evaluate the extent to which either vicariance and/or dispersal events have shaped the population structure of N. vlamingii. The analysis produced several unexpected findings. Firstly, the genetic structure of this species was temporal rather than spatial. Secondly, there was no evidence of a barrier to dispersal throughout the vast distribution range. Apparently larvae of this species traverse vicariance barriers that inhibit inter-oceanic migration of other widespread reef fish taxa. Thirdly, an unusual life history and long evolutionary history was associated with a population structure that was unique amongst coral reef fishes in terms of the magnitude and pattern of genetic diversity (haplotype diversity, h = 1.0 and nucleotide diversity π = 13.6%). In addition to these unique characteristics, there was no evidence of isolation by distance (r = 0.458, R ² = 0.210, P = 0.078) as has also been shown for some other widespread reef species. However, some reductions in gene flow were observed among and within Ocean basins [Indian–Pacific analysis of molecular variance (AMOVA), Φ _st = 0.0766, P < 0.05; West Indian–East Indian–Pacific AMOVA Φ _st = 0.079, P < 0.05]. These findings are contrasted with recent studies of coral reef fishes that imply a greater degree of spatial structuring in coral reef fish populations than would be expected from the dispersive nature of their life cycles. We conclude that increased taxon sampling of coral reef fishes for phylogeographic analysis will provide an extended view of the ecological and evolutionary processes shaping coral reef fish diversity at both ends of the life history spectrum.     

A multivariate analysis of beta diversity across organisms and environments

We examined variability in hierarchical beta diversity across ecosystems, geographical gradients, and organism groups using multivariate spatial mixed modeling analysis of two independent data sets. The larger data set comprised reported ratios of regional species richness (RSR) to local species richness (LSR) and the second data set consisted of RSR:LSR ratios derived from nested species-area relationships. There was a negative, albeit relatively weak, relationship between beta diversity and latitude. We found only relatively subtle differences in beta diversity among the realms, yet beta diversity was lower in marine systems than in terrestrial or freshwater realms. Beta diversity varied significantly among organisms' major characteristics such as body mass, trophic position, and dispersal type in the larger data set. Organisms that disperse via seeds had highest beta diversity, and passively dispersed organisms showed the lowest beta diversity. Furthermore, autotrophs had lower beta diversity than organisms higher up the food web; omnivores and carnivores had consistently higher beta diversity. This is evidence that beta diversity is simultaneously controlled by extrinsic factors related to geography and environment, and by intrinsic factors related to organism characteristics.     

Latitudinal variation in local interactions and regional enrichment shape patterns of marine community diversity

While communities are shaped by both local interactions and enrichment from the regional species pool, we propose a hypothesis that the balance of these forces shifts with latitude, with regional enrichment dominating at high latitudes and local interactions dominating at low latitudes. To test this hypothesis, we conducted a latitudinal-scale experiment with marine epifaunal communities. In four regions of the North Atlantic Ocean and Caribbean Sea, we used mimics of ecosystem engineers to manipulate biogenic structural complexity. We iteratively evaluated diversity patterns of experimental communities up to one year after deployment. Additional data were also collected from one of our tropical sites 2.5 years after initial deployment. As hypothesized, we found a reciprocal latitudinal gradient in the effects of the structurally complex mimics and regional enrichment. In the tropics, local diversity was always higher in association with the mimics than in exposed areas that were more open to predation. This effect was consistent across two spatial scales and beyond the one-year timescale of the experiment. In temperate communities, no consistent effects of the mimics on diversity were observed. However, the proportion of species from the regional species pool that were present at the local scale increased from the tropics to the temperate zone, consistent with the hypothesis that higher-latitude communities may experience greater influence from the regional species pool than communities at low latitudes. This study represents the first large-scale experimental demonstration that suggests that the relative impact of local interactions and regional enrichment on community diversity may depend on latitude.