Incorporating phylogenetic information for the definition of floristic districts in hyperdiverse Amazon forests: Implications for conservation

Using complementary metrics to evaluate phylogenetic diversity can facilitate the delimitation of floristic units and conservation priority areas. In this study, we describe the spatial patterns of phylogenetic alpha and beta diversity, phylogenetic endemism, and evolutionary distinctiveness of the hyperdiverse Ecuador Amazon forests and define priority areas for conservation. We established a network of 62 one‐hectare plots in terra firme forests of Ecuadorian Amazon. In these plots, we tagged, collected, and identified every single adult tree with dbh ≥10 cm. These data were combined with a regional community phylogenetic tree to calculate different phylogenetic diversity (PD) metrics in order to create spatial models. We used Loess regression to estimate the spatial variation of taxonomic and phylogenetic beta diversity as well as phylogenetic endemism and evolutionary distinctiveness. We found evidence for the definition of three floristic districts in the Ecuadorian Amazon, supported by both taxonomic and phylogenetic diversity data. Areas with high levels of phylogenetic endemism and evolutionary distinctiveness in Ecuadorian Amazon forests are unprotected. Furthermore, these areas are severely threatened by proposed plans of oil and mining extraction at large scales and should be prioritized in conservation planning for this region.     

Geographical and environmental controls of palm beta diversity in paleo-riverine terrace forests in Amazonian Peru

The palm (Arecaceae) community on low paleo-riverine terraces (terrace forest) in the north-western Amazon, is described, and we assessed the importance of environmental differences and geographic distance as drivers of its local (25² grain and 0–500 extent) and regional scale (500² grain and 0.3–143 km extent) beta diversity using ordination, multiple regressions on distance matrices and Indicator Species Analysis. A total of 15,869 individuals and 37 species of palm were sampled in 10 terrace forest transects, while 3758 individuals and 21 species were sampled in two adjacent floodplain forest transects for comparison. The terrace and floodplain forest were clearly different in their diversity and floristic composition. The relative importance of geographical distance and environmental difference as controls of terrace forest beta diversity was scale dependent, with environmental differences, notably in soil moisture, dominating at local scales and geographical distance dominating at regional scales. In fact, none of the environmental factors had a significant influence on regional-scale beta diversity. The geographical distance decay in floristic similarity was markedly steeper at local scale ( −0.25 km ⁻¹) than at regional scale ( −0.003 km⁻¹). Such a nonlinear decay is expected if simple dispersal limitation controls beta diversity. However, the absent flattening of the distance decay at the largest distances and the sub-Andean affinities of the westernmost palm communities suggest that large-scale biogeographical processes also contribute to the regional-scale beta diversity. Hereby our results indicate that not only local environment, but also dispersal limitation and biogeographical history can be important controls of the diversity and composition of local plant communities.     

Beta diversity and oligarchic dominance in the tropical forests of Southern Costa Rica

Recent studies have reported a consistent pattern of strong dominance of a small subset of tree species in neotropical forests. These species have been called “hyperdominant” at large geographical scales and “oligarchs” at regional‐landscape scales when being abundant and frequent. Forest community assembly is shaped by environmental factors and stochastic processes, but so far the contribution of oligarchic species to the variation of community composition (i.e., beta diversity) remains poorly known. To that end, we established 20.1‐ha plots, that is, five sites with four forest types (ridge, slope and ravine primary forest, and secondary forest) per site, in humid lowland tropical forests of southwestern Costa Rica to (a) investigate how community composition responds to differences in topography, successional stage, and distance among plots for different groups of species (all, oligarch, common and rare/very rare species) and (b) identify oligarch species characterizing changes in community composition among forest types. From a total of 485 species of trees, lianas and palms recorded in this study only 27 species (i.e., 6%) were nominated as oligarch species. Oligarch species accounted for 37% of all recorded individuals and were present in at least half of the plots. Plant community composition significantly differed among forest types, thus contributing to beta diversity at the landscape scale. Oligarch species was the component best explained by geographical and topographic variables, allowing a confident characterization of the beta diversity among tropical lowland forest stands. Abstract in Spanish is available with online material.     

Biodiversity assessment in incomplete inventories: leaf litter ant communities in several types of Bornean rain forest

Biodiversity assessment of tropical taxa is hampered by their tremendous richness, which leads to large numbers of singletons and incomplete inventories in survey studies. Species estimators can be used for assessment of alpha diversity, but calculation of beta diversity is hampered by pseudo-turnover of species in undersampled plots. To assess the impact of unseen species, we investigated different methods, including an unbiased estimator of Shannon beta diversity that was compared to biased calculations. We studied alpha and beta diversity of a diverse ground ant assemblage from the Southeast Asian island of Borneo in different types of tropical forest: diperocarp forest, alluvial forest, limestone forest and heath forests. Forests varied in plant composition, geology, flooding regimes and other environmental parameters. We tested whether forest types differed in species composition and if species turnover was a function of the distance between plots at different spatial scales. As pseudo-turnover may bias beta diversity we hypothesized a large effect of unseen species reducing beta diversity. We sampled 206 ant species (25% singletons) from ten subfamilies and 55 genera. Diversity partitioning among the four forest types revealed that whereas alpha species richness and alpha Shannon diversity were significantly smaller than expected, beta-diversity for both measurements was significantly higher than expected by chance. This result was confirmed when we used the unbiased estimation of Shannon diversity: while alpha diversity was much higher, beta diversity differed only slightly from biased calculations. Beta diversity as measured with the Chao-Sørensen or Morisita-Horn Index correlated with distance between transects and between sample points, indicating a distance decay of similarity between communities. We conclude that habitat heterogeneity has a high influence on ant diversity and species turnover in tropical sites and that unseen species may have only little impact on calculation of Shannon beta diversity when sampling effort has been high.     

Mountain metacommunities: climate and spatial connectivity shape ant diversity in a complex landscape

Understanding what drives biodiversity patterns across scales is a central goal of ecology. Both environmental gradients and spatial landscape structure have been found to be important factors influencing species distributions and community composition, and partly reflect the balance of underlying deterministic and stochastic community processes. In some systems, environmental gradients and spatial connectivity are intertwined in that steep environmental gradients serve as boundaries on species movements and impose environment‐derived complex spatial structure to metacommunities. Mountainous landscapes are prime examples of this, and recent theory has linked principles of geomorphology, environmental gradients, and spatial structure to make predictions for resulting community patterns. In this context, we examine variation in taxonomic and phylogenetic ant diversity patterns along a geographic transect spanning > 5000 m in elevational range in the Hengduan mountains of southern China. We found that environmental gradients dominate variation in both alpha and beta diversity in this landscape, with alpha diversity strongly declining with elevation and beta diversity driven by elevational differences. However, within an elevational band spatial connectivity predicts beta diversity better than geographic distance. Our findings deviate from theoretical predictions in several ways, notably alpha diversity is monotonically declining and within‐band beta diversity is invariant with increasing elevation. The discrepancies between theory and observation may be explained by differences in the Hengduan landscape from idealized fluvial landscapes, such as a lack of a mid‐elevation peak in connectivity, as well as evolutionary limits on the source pool of species available to populate metacommunities at different elevations. The latter is supported by variation in phylogenetic community structure with elevation. Our results demonstrate the power of conceptual, statistical, and theoretical frameworks that integrate the roles of environment and spatial structure in metacommunities, but that additional work is needed to bridge the gap between abstract theory and real systems.     

Long distance dispersal and connectivity in amphi-Atlantic corals at regional and basin scales

Among Atlantic scleractinian corals, species diversity is highest in the Caribbean, but low diversity and high endemism are observed in various peripheral populations in central and eastern Atlantic islands and along the coasts of Brazil and West Africa. The degree of connectivity between these distantly separated populations is of interest because it provides insight into processes at both evolutionary and ecological time scales, such as speciation, recruitment dynamics and the persistence of coral populations. To assess connectivity in broadly distributed coral species of the Atlantic, DNA sequence data from two nuclear markers were obtained for six coral species spanning their distributional ranges. At basin-wide scales, significant differentiation was generally observed among populations in the Caribbean, Brazil and West Africa. Concordance of patterns in connectivity among co-distributed taxa indicates that extrinsic barriers, such as the Amazon freshwater plume or long stretches of open ocean, restrict dispersal of coral larvae from region to region. Within regions, dispersal ability appears to be influenced by aspects of reproduction and life history. Two broadcasting species, Siderastrea siderea and Montastraea cavernosa, were able to maintain gene flow among populations separated by as much as 1,200 km along the coast of Brazil. In contrast, brooding species, such as Favia gravida and Siderastrea radians, had more restricted gene flow along the Brazilian coast.     

Connectivity and gene flow among Eastern Tiger Salamander (Ambystoma tigrinum) populations in highly modified anthropogenic landscapes

Fragmented landscapes resulting from anthropogenic habitat modification can have significant impacts on dispersal, gene flow, and persistence of wildlife populations. Therefore, quantifying population connectivity across a mosaic of habitats in highly modified landscapes is critical for the development of conservation management plans for threatened populations. Endangered populations of the eastern tiger salamander (Ambystoma tigrinum) in New York and New Jersey are at the northern edge of the species’ range and remaining populations persist in highly developed landscapes in both states. We used landscape genetic approaches to examine regional genetic population structure and potential barriers to migration among remaining populations. Despite the post-glacial demographic processes that have shaped genetic diversity in tiger salamander populations at the northern extent of their range, we found that populations in each state belong to distinct genetic clusters, consistent with the large geographic distance that separates them. We detected overall low genetic diversity and high relatedness within populations, likely due to recent range expansion, isolation, and relatively small population sizes. Nonetheless, landscape connectivity analyses reveal habitat corridors among remaining breeding ponds. Furthermore, molecular estimates of population connectivity among ponds indicate that gene flow still occurs at regional scales. Further fragmentation of remaining habitat will potentially restrict dispersal among breeding ponds, cause the erosion of genetic diversity, and exacerbate already high levels of inbreeding. We recommend the continued management and maintenance of habitat corridors to ensure long-term viability of these endangered populations.     

Phylogenetic beta diversity: linking ecological and evolutionary processes across space in time

A key challenge in ecological research is to integrate data from different scales to evaluate the ecological and evolutionary mechanisms that influence current patterns of biological diversity. We build on recent attempts to incorporate phylogenetic information into traditional diversity analyses and on existing research on beta diversity and phylogenetic community ecology. Phylogenetic beta diversity (phylobetadiversity) measures the phylogenetic distance among communities and as such allows us to connect local processes, such as biotic interactions and environmental filtering, with more regional processes including trait evolution and speciation. When combined with traditional measures of beta diversity, environmental gradient analyses or ecological niche modelling, phylobetadiversity can provide significant and novel insights into the mechanisms underlying current patterns of biological diversity.     

Conservation biological control and enemy diversity on a landscape scale

Conservation biological control in agroecosystems requires a landscape management perspective, because most arthropod species experience their habitat at spatial scales beyond the plot level, and there is spillover of natural enemies across the crop–noncrop interface. The species pool in the surrounding landscape and the distance of crop from natural habitat are important for the conservation of enemy diversity and, in particular, the conservation of poorly-dispersing and specialized enemies. Hence, structurally complex landscapes with high habitat connectivity may enhance the probability of pest regulation. In contrast, generalist and highly vagile enemies may even profit from the high primary productivity of crops at a landscape scale and their abundance may partly compensate for losses in enemy diversity. Conservation biological control also needs a multitrophic perspective. For example, entomopathogenic fungi, plant pathogens and endophytes as well as below- and above-ground microorganisms are known to influence pest-enemy interactions in ways that vary across spatiotemporal scales. Enemy distribution in agricultural landscapes is determined by beta diversity among patches. The diversity needed for conservation biological control may occur where patch heterogeneity at larger spatial scales is high. However, enemy communities in managed systems are more similar across space and time than those in natural systems, emphasizing the importance of natural habitat for a spillover of diverse enemies. According to the insurance hypothesis, species richness can buffer against spatiotemporal disturbances, thereby insuring functioning in changing environments. Seemingly redundant enemy species may become important under global change. Complex landscapes characterized by highly connected crop–noncrop mosaics may be best for long-term conservation biological control and sustainable crop production, but experimental evidence for detailed recommendations to design the composition and configuration of agricultural landscapes that maintain a diversity of generalist and specialist natural enemies is still needed.     

Biotic homogenisation and differentiation as directional change in beta diversity: synthesising driver–response relationships to develop conceptual models across ecosystems

Biotic homogenisation is defined as decreasing dissimilarity among ecological assemblages sampled within a given spatial area over time. Biotic differentiation, in turn, is defined as increasing dissimilarity over time. Overall, changes in the spatial dissimilarities among assemblages (termed ‘beta diversity’) is an increasingly recognised feature of broader biodiversity change in the Anthropocene. Empirical evidence of biotic homogenisation and biotic differentiation remains scattered across different ecosystems. Most meta-analyses quantify the prevalence and direction of change in beta diversity, rather than attempting to identify underlying ecological drivers of such changes. By conceptualising the mechanisms that contribute to decreasing or increasing dissimilarity in the composition of ecological assemblages across space, environmental managers and conservation practitioners can make informed decisions about what interventions may be required to sustain biodiversity and can predict potential biodiversity outcomes of future disturbances. We systematically reviewed and synthesised published empirical evidence for ecological drivers of biotic homogenisation and differentiation across terrestrial, marine, and freshwater realms to derive conceptual models that explain changes in spatial beta diversity. We pursued five key themes in our review: (i) temporal environmental change; (ii) disturbance regime; (iii) connectivity alteration and species redistribution; (iv) habitat change; and (v) biotic and trophic interactions. Our first conceptual model highlights how biotic homogenisation and differentiation can occur as a function of changes in local (alpha) diversity or regional (gamma) diversity, independently of species invasions and losses due to changes in species occurrence among assemblages. Second, the direction and magnitude of change in beta diversity depends on the interaction between spatial variation (patchiness) and temporal variation (synchronicity) of disturbance events. Third, in the context of connectivity and species redistribution, divergent beta diversity outcomes occur as different species have different dispersal characteristics, and the magnitude of beta diversity change associated with species invasions also depends strongly on alpha and gamma diversity prior to species invasion. Fourth, beta diversity is positively linked with spatial environmental variability, such that biotic homogenisation and differentiation occur when environmental heterogeneity decreases or increases, respectively. Fifth, species interactions can influence beta diversity via habitat modification, disease, consumption (trophic dynamics), competition, and by altering ecosystem productivity. Our synthesis highlights the multitude of mechanisms that cause assemblages to be more or less spatially similar in composition (taxonomically, functionally, phylogenetically) through time. We consider that future studies should aim to enhance our collective understanding of ecological systems by clarifying the underlying mechanisms driving homogenisation or differentiation, rather than focusing only on reporting the prevalence and direction of change in beta diversity, per se.     

How pervasive is biotic homogenization in human‐modified tropical forest landscapes?

Land‐cover change and ecosystem degradation may lead to biotic homogenization, yet our understanding of this phenomenon over large spatial scales and different biotic groups remains weak. We used a multi‐taxa dataset from 335 sites and 36 heterogeneous landscapes in the Brazilian Amazon to examine the potential for landscape‐scale processes to modulate the cumulative effects of local disturbances. Biotic homogenization was high in production areas but much less in disturbed and regenerating forests, where high levels of among‐site and among‐landscape β‐diversity appeared to attenuate species loss at larger scales. We found consistently high levels of β‐diversity among landscapes for all land cover classes, providing support for landscape‐scale divergence in species composition. Our findings support concerns that β‐diversity has been underestimated as a driver of biodiversity change and underscore the importance of maintaining a distributed network of reserves, including remaining areas of undisturbed primary forest, but also disturbed and regenerating forests, to conserve regional biota.     

Disentangling environmental correlates of vascular plant biodiversity in a Mediterranean hotspot

We determined the environmental correlates of vascular plant biodiversity in the Baetic‐Rifan region, a plant biodiversity hotspot in the western Mediterranean. A catalog of the whole flora of Andalusia and northern Morocco, the region that includes most of the Baetic‐Rifan complex, was compiled using recent comprehensive floristic catalogs. Hierarchical cluster analysis (HCA) and detrended correspondence analysis (DCA) of the different ecoregions of Andalusia and northern Morocco were conducted to determine their floristic affinities. Diversity patterns were studied further by focusing on regional endemic taxa. Endemic and nonendemic alpha diversities were regressed to several environmental variables. Finally, semi‐partial regressions on distance matrices were conducted to extract the respective contributions of climatic, altitudinal, lithological, and geographical distance matrices to beta diversity in endemic and nonendemic taxa. We found that West Rifan plant assemblages had more similarities with Andalusian ecoregions than with other nearby northern Morocco ecoregions. The endemic alpha diversity was explained relatively well by the environmental variables related to summer drought and extreme temperature values. Of all the variables, geographical distance contributed by far the most to spatial turnover in species diversity in the Baetic‐Rifan hotspot. In the Baetic range, climate was the most significant driver of nonendemic species beta diversity, while lithology and climate were the main drivers of endemic beta diversity. Despite the fact that Andalusia and northern Morocco are presently separated by the Atlantic Ocean and the Mediterranean Sea, the Baetic and Rifan mountain ranges have many floristic similarities – especially in their western ranges – due to past migration of species across the Strait of Gibraltar. Climatic variables could be shaping the spatial distribution of endemic species richness throughout the Baetic‐Rifan hotspot. Determinants of spatial turnover in biodiversity in the Baetic‐Rifan hotspot vary in importance between endemic and nonendemic species.     

Contrasting demographic and genetic estimates of dispersal in the endangered Coahuilan box turtle: a contemporary approach to conservation

The evolutionary viability of an endangered species depends upon gene flow among subpopulations and the degree of habitat patch connectivity. Contrasting population connectivity over ecological and evolutionary timescales may provide novel insight into what maintains genetic diversity within threatened species. We employed this integrative approach to evaluating dispersal in the critically endangered Coahuilan box turtle (Terrapene coahuila) that inhabits isolated wetlands in the desert‐spring ecosystem of Cuatro Ciénegas, Mexico. Recent wetland habitat loss has altered the spatial distribution and connectivity of habitat patches; and we therefore predicted that T. coahuila would exhibit limited movement relative to estimates of historic gene flow. To evaluate contemporary dispersal patterns, we employed mark–recapture techniques at both local (wetland complex) and regional (intercomplex) spatial scales. Gene flow estimates were obtained by surveying genetic variation at nine microsatellite loci in seven subpopulations located across the species’ geographical range. The mark–recapture results at the local spatial scale reveal frequent movement among wetlands that was unaffected by interwetland distance. At the regional spatial scale, dispersal events were relatively less frequent between wetland complexes. The complementary analysis of population genetic substructure indicates strong historic gene flow (global F_ST = 0.01). However, a relationship of genetic isolation by distance across the geographical range suggests that dispersal limitation exists at the regional scale. Our approach of contrasting direct and indirect estimates of dispersal at multiple spatial scales in T. coahuila conveys a sustainable evolutionary trajectory of the species pending preservation of threatened wetland habitats and a range‐wide network of corridors.     

Additive partitioning of butterfly diversity in a fragmented landscape: importance of scale and implications for conservation

Aim Most of the Atlantic Forest in Brazil occurs in fragments of various sizes. Previous studies indicate that forest fragmentation affects fruit‐feeding butterflies. Conservation strategies that seek to preserve organisms that are distributed in high‐fragmented biomes need to understand the spatial distribution of these organisms across the landscape. In view of the importance of understanding the fauna of these forest remnants, the objective of the present work is to investigate the extent to which the diversity of this group varies across spatial scales ranging from within‐forest patches to between landscapes. Location South America, south‐eastern Brazil, São Paulo State. Methods We used bait traps to sample fruit feeding butterflies at 50 points in 10 fragments in two different landscapes during a period of 12 months. Total species richness and Shannon index were partitioned additively in diversity at trap level, and beta diversity was calculated among traps, among forest patches, and between landscapes. We used permutation tests to compare these values to the expected ones under the null hypothesis that beta diversity is only a random sampling effect. Results There was significant beta diversity at the smallest scale examined; however, the significance at higher scales depends on the diversity measurement used. Beta diversity with Shannon index was smaller than expected by chance among fragments, whereas species richness was not. Among landscapes, only beta diversity in richness was higher than expected by chance. Main conclusions The results observed occur because there is great variability in species composition among forest patches in the same landscape, changing this diversity even though the communities are formed from the same pool of species. At the largest scale evaluated (between landscapes), these pattern changes and differences in beta diversity in richness were detectable. This difference is probably caused by the presence of rare species. Thus, a conservation strategy that seeks to preserve as many species as possible per unit of area in high‐fragmented biomes should give priority to protecting fragments in different landscapes, rather than more fragments in the same landscape.     

Assessing endemism at multiple spatial scales, with an example from the Australian vascular flora

Aim  To develop an approach for assessing the spatial scale of centres of endemism among species level data.
Location Australia.
Methods  Endemism is inherently scale dependent. Therefore, the Corrected Weighted Endemism (CWE) index used by Crisp et al. [ J. Biogeogr. (2001)28:183] is extended to account for species samples in local neighbourhoods as a Spatial CWE index. This then allows an analysis of how the degree of endemism of a location (cell) changes with spatial scale. The quality of the Spatial CWE index results are assessed using three spatial randomizations at the species level with and without preserving species richness and distributional patterns. We show that CWE is equivalent to beta diversity and predict that it should show high rates of change around centres of endemism.
Results  Similar patterns to those found by Crisp et al. using a data set of vascular flora from Australia are retrieved, but the extent to which they are scale dependent is more easily identified. For example, the Central Australian centre discounted by Crisp et al. is identified when a three-cell radius neighbourhood is used. However, the level of endemism in this centre is no greater than in the margins of many of the coastal centres of endemism. Most of the identified centres of endemism are better than random at all scales and are increasingly so as the spatial scale increases. As predicted, the highest rate of change in Spatial CWE (beta diversity) is most often between zero- and one-cell radius neighbours in most centres of endemism.
Main conclusions  The explicit incorporation of geographical space in analyses allows for a greater understanding of the scale-dependence of phenomena, in this case endemism and beta diversity.     

The effects of forest conversion to oil palm on ground‐foraging ant communities depend on beta diversity and sampling grain

Beta diversity – the variation in species composition among spatially discrete communities – and sampling grain – the size of samples being compared – may alter our perspectives of diversity within and between landscapes before and after agricultural conversion. Such assumptions are usually based on point comparisons, which do not accurately capture actual differences in total diversity. Beta diversity is often not rigorously examined. We investigated the beta diversity of ground‐foraging ant communities in fragmented oil palm and forest landscapes in Sabah, Malaysia, using diversity metrics transformed from Hill number equivalents to remove dependences on alpha diversity. We compared the beta diversities of oil palm and forest, across three hierarchically nested sampling grains. We found that oil palm and forest communities had a greater percentage of total shared species when larger samples were compared. Across all grains and disregarding relative abundances, there was higher beta diversity of all species among forest communities. However, there were higher beta diversities of common and very abundant (dominant) species in oil palm as compared to forests. Differences in beta diversities between oil palm and forest were greatest at the largest sampling grain. Larger sampling grains in oil palm may generate bigger species pools, increasing the probability of shared species with forest samples. Greater beta diversity of all species in forest may be attributed to rare species. Oil palm communities may be more heterogeneous in common and dominant species because of variable community assembly events. Rare and also common species are better captured at larger grains, boosting differences in beta diversity between larger samples of forest and oil palm communities. Although agricultural landscapes support a lower total diversity than natural forests, diversity especially of abundant species is still important for maintaining ecosystem stability. Diversity in agricultural landscapes may be greater than expected when beta diversity is accounted for at large spatial scales.     

Contrasting effects of mosaic structure on alpha and beta diversity of bird assemblages in a human‐modified landscape

Habitat loss and fragmentation are key processes causing biodiversity loss in human‐modified landscapes. Knowledge of these processes has largely been derived from measuring biodiversity at the scale of ‘within‐habitat’ fragments with the surrounding landscape considered as matrix. Yet, the loss of variation in species assemblages ‘among’ habitat fragments (landscape‐scale) may be as important a driver of biodiversity loss as the loss of diversity ‘within’ habitat fragments (local‐scale). We tested the hypothesis that heterogeneity in vegetation cover is important for maintaining alpha and beta diversity in human‐modified landscapes. We surveyed bird assemblages in eighty 300‐m‐long transects nested within twenty 1‐km² vegetation ‘mosaics’, with mosaics assigned to four categories defined by the cover extent and configuration of native eucalypt forest and exotic pine plantation. We examined bird assemblages at two spatial scales: 1) within and among transects, and 2) within and among mosaics. Alpha diversity was the mean species diversity within‐transects or within‐mosaics and beta diversity quantified the effective number of compositionally distinct transects or mosaics. We found that within‐transect alpha diversity was highest in vegetation mosaics defined by continuous eucalypt forest, lowest in mosaics of continuous pine plantation, and at intermediate levels in mosaics containing eucalypt patches in a pine matrix. We found that eucalypt mosaics had lower beta diversity than other mosaic types when ignoring relative abundances, but had similar or higher beta diversity when weighting with species abundances. Mosaics containing both pine and eucalypt forest differed in their bird compositional variation among transects, despite sharing a similar suite of species. This configuration effect at the mosaic scale reflected differences in vegetation composition among transects. Maintaining heterogeneity in vegetation cover could help to maintain variation among bird assemblages across landscapes, thus partially offsetting local‐scale diversity losses due to fragmentation. Critical to this is the retention of remnant native vegetation.     

Cover Image

1. Understanding the role of environmental filtering and spatial processes along environmental gradients in assembling and maintaining aquatic communities in rare habitats is crucial for land management and biological conservation. We investigated the relative roles of environmental and spatial factors influencing beta (β) diversity of aquatic macroinvertebrate assemblages in 36 interdunal wetlands from five freshwater sand dune areas across two ecoregions spanning the latitudinal gradient of Lake Michigan. We hypothesised that aquatic macroinvertebrate diversity and composition would vary along the coastline because of an underlying environmental gradient. We predicted high species replacement correlated with environmental (local and regional) conditions and increasing species diversity with decreasing latitude.
2. We calculated sample completeness, obtained diversity estimates based on Hill numbers and used abundance-based β partitioning and multivariate analysis to examine β diversity, and its replacement and nestedness components in relation to local and regional predictors.
3. Despite a short latitudinal gradient, we detected a significant increase in species richness with decreasing latitude, underpinned by a strong temperature and precipitation gradient. Species replacement (balanced variation in abundance) was high at all spatial scales (wetland, dune area, ecoregion and coastline), and correlated with local and regional environmental variables.
4. Community dissimilarity showed no marked increase with spatial extent, which suggests a system where local-scale environmental filtering prevents dispersal driven homogenisation. Both local and bioclimatic factors were correlated with aquatic macroinvertebrate dissimilarity, but local factors played a larger role.
5. This study simultaneously examined the response of alpha and β diversity to geographical and environmental variables. Our results indicate that macroinvertebrates respond to abiotic factors by tracking suitable environmental conditions among locally variable interdunal wetlands. Thus, high dispersal along the coastline conveys resiliency to a hydrologically dynamic wetland system, which allows aquatic macroinvertebrates to contribute an integral portion of sand dune biodiversity within the Laurentian Great Lakes region. The high species turnover found suggests that conservation strategies should consider coastline connectivity among dune areas to maintain freshwater biodiversity.

     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Partitioning species diversity across landscapes and regions: a hierarchical analysis of alpha, beta, and gamma diversity

Species diversity may be additively partitioned within and among samples (alpha and beta diversity) from hierarchically scaled studies to assess the proportion of the total diversity (gamma) found in different habitats, landscapes, or regions. We developed a statistical approach for testing null hypotheses that observed partitions of species richness or diversity indices differed from those expected by chance, and we illustrate these tests using data from a hierarchical study of forest-canopy beetles. Two null hypotheses were implemented using individual- and sample-based randomization tests to generate null distributions for alpha and beta components of diversity at multiple sampling scales. The two tests differed in their null distributions and power to detect statistically significant diversity components. Individual-based randomization was more powerful at all hierarchical levels and was sensitive to departures between observed and null partitions due to intraspecific aggregation of individuals. Sample-based randomization had less power but still may be useful for determining whether different habitats show a higher degree of differentiation in species diversity compared with random samples from the landscape. Null hypothesis tests provide a basis for inferences on partitions of species richness or diversity indices at multiple sampling levels, thereby increasing our understanding of how alpha and beta diversity change across spatial scales.