Effect of Human Disturbance on Bee Communities in a Forested Ecosystem

Abstract:  It is important for conservation biologists to understand how well species persist in human-dominated ecosystems because protected areas constitute a small fraction of the Earth's surface and because anthropogenic habitats may offer more opportunities for conservation than has been previously thought. We investigated how an important functional group, pollinators (bees; Hymenoptera: Apiformes), are affected by human land use at the landscape and local scales in southern New Jersey (U.S.A.). We established 40 sites that differed in surrounding landscape cover or local habitat type and collected 2551 bees of 130 species. The natural habitat in this ecosystem is a forested, ericaceous heath. Bee abundance and species richness within forest habitat decreased, not increased, with increasing forest cover in the surrounding landscape. Similarly, bee abundance was greater in agricultural fields and suburban and urban developments than in extensive forests, and the same trend was found for species richness. Particular species groups that might be expected to show greater sensitivity to habitat loss, such as floral specialists and bees of small or large body size, did not show strong positive associations with forest habitat. Nevertheless, 18 of the 130 bee species studied were positively associated with extensive forest. One of these species is a narrow endemic that was last seen in 1939. Our results suggest that at least in this system, moderate anthropogenic land use may be compatible with the conservation of many, but not all, bee species.     

Distinguishing stressors acting on land bird communities in an urbanizing environment

Urbanization has profound influences on ecological communities, but our understanding of causal mechanisms is limited by a lack of attention to its component stressors. Published research suggests that at landscape scales, habitat loss and fragmentation are the major drivers of community change, whereas at local scales, human activity and vegetation management are the primary stressors. Little research has focused on whether urbanization stressors may supplant natural factors as dominant forces structuring communities. We used model selection to determine the relative importance of urban development, human activity, local and landscape vegetation, topography, and geographical location in explaining land bird species richness, abundance, and dominance. We analyzed the entire community and groups of species based on ecological characteristics, using data collected in remnant forests along a gradient of urban development in the Lake Tahoe basin, California and Nevada, USA. Urbanization stressors were consistently among the principal forces structuring the land bird community. Strikingly, disturbance from human activity was the most important factor for richness in many cases, surpassing even habitat loss from development. Landscape-scale factors were consistently more important than local-scale factors for abundance. In demonstrating considerable changes in land bird community structure, our results suggest that ecosystem function in urban areas may be severely compromised. Such changes compel local- and landscape-scale management, focused research, and long-term monitoring to retain biodiversity in urban areas to the extent possible.     

Bird‐community responses to habitat creation in a long‐term,large‐scale natural experiment

Ecosystem function and resilience are compromised when habitats become fragmented due to land‐use change. This has led to national and international conservation strategies aimed at restoring habitat extent and improving functional connectivity (i.e., maintaining dispersal processes). However, biodiversity responses to landscape‐scale habitat creation and the relative importance of spatial and temporal scales are poorly understood, and there is disagreement over which conservation strategies should be prioritized. We used 160 years of historic post‐agricultural woodland creation as a natural experiment to evaluate biodiversity responses to habitat creation in a landscape context. Birds were surveyed in 101 secondary, broadleaf woodlands aged 10–160 years with ≥80% canopy cover and in landscapes with 0‐17% broadleaf woodland cover within 3000 m. We used piecewise structural equation modeling to examine the direct and indirect relationships between bird abundance and diversity, ecological continuity, patch characteristics, and landscape structure and quantified the relative conservation value of local and landscape scales for bird communities. Ecological continuity indirectly affected overall bird abundance and species richness through its effects on stand structure, but had a weaker influence (effect size near 0) on the abundance and diversity of species most closely associated with woodland habitats. This was probably because woodlands were rapidly colonized by woodland generalists in ≤10 years (minimum patch age) but were on average too young (median 50 years) to be colonized by woodland specialists. Local patch characteristics were relatively more important than landscape characteristics for bird communities. Based on our results, biodiversity responses to habitat creation depended on local‐ and landscape‐scale factors that interacted across time and space. We suggest that there is a need for further studies that focus on habitat creation in a landscape context and that knowledge gained from studies of habitat fragmentation and loss should be used to inform habitat creation with caution because the outcomes are not necessarily reciprocal.     

Climate, habitat, and species interactions at different scales determine the structure of a Neotropical bat community

Climate, habitat, and species interactions are factors that control community properties (e.g., species richness, abundance) across various spatial scales. Usually, researchers study how a few properties are affected by one factor in isolation and at one scale. Hence, there are few multi-scale studies testing how multiple controlling factors simultaneously affect community properties at different scales. We ask whether climate, habitat structure, or insect resources at each of three spatial scales explains most of the variation in six community properties and which theory best explains the distribution of selected community properties across a rainfall gradient. We studied a Neotropical insectivorous bat ensemble in the Isthmus of Panama with acoustic monitoring techniques. Using climatological data, habitat surveys, and insect captures in a hierarchical sampling design we determined how much variation of the community properties was explained by the three factors employing two approaches for variance partitioning. Our results revealed that most of the variation in species richness, total abundance, and feeding activity occurred at the smallest spatial scale and was explained by habitat structure. In contrast, climate at large scales explained most of the variation in individual species' abundances. Although each species had an idiosyncratic response to the gradient, species richness peaked at intermediate levels of precipitation, whereas total abundance was very similar across sites, suggesting density compensation. All community properties responded in a different manner to the factor and scale under consideration.     

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.     

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.     

A multi-scale examination of stopover habitat use by birds

Most of our understanding of habitat use by migrating land birds comes from studies conducted at single, small spatial scales, which may overemphasize the importance of intrinsic habitat factors, such as food availability, in shaping migrant distributions. We believe that a multi-scale approach is essential to assess the influence of factors that control en route habitat use. We determined the relative importance of eight variables, each operating at a habitat-patch, landscape, or regional spatial scale, in explaining the differential use of hardwood forests by Nearctic-Neotropical land birds during migration. We estimated bird densities through transect surveys at sites near the Mississippi coast during spring and autumn migration within landscapes with variable amounts of hardwood forest cover. At a regional scale, migrant density increased with proximity to the coast, which was of moderate importance in explaining bird densities, probably due to constraints imposed on migrants when negotiating the Gulf of Mexico. The amount of hardwood forest cover at a landscape scale was positively correlated with arthropod abundance and had the greatest importance in explaining densities of all migrants, as a group, during spring, and of insectivorous migrants during autumn. Among landscape scales ranging from 500 m to 10 km radius, the densities of migrants were, on average, most strongly and positively related to the amount of hardwood forest cover within a 5 km radius. We suggest that hardwood forest cover at this scale may be an indicator of habitat quality that migrants use as a cue when landing at the end of a migratory flight. At the patch scale, direct measures of arthropod abundance and plant community composition were also important in explaining migrant densities, whereas habitat structure was of little importance. The relative amount of fleshy-fruited trees was positively related and was the most important variable explaining frugivorous migrant density during autumn. Although constraints extrinsic to habitat had a moderate role in explaining migrant distributions, our results are consistent with the view that food availability is the ultimate factor shaping the distributions of birds during stopover.     

Using Gall Wasps on Oaks to Test Broad Ecological Concepts

Abstract:  Planning conservation of insect herbivores requires knowing what needs to be conserved and developing a set of predictor variables that aid management. We conducted a state-wide survey to examine the species richness of gall wasps (Hymenoptera: Cynipidae) on six oak species dominant in the threatened scrub-oak vegetation in peninsular Florida. Eighty-eight cynipid species were recorded; 23 were new species to Florida (a 35% increase), including 17 species new to science and 6 species newly recorded in the state. The cynipid species represented 68% of cynipids of Florida, on only 24% of oak species sampled. This fauna represents a hotspot of richness, justifying conservation initiatives in scrub-oak habitat and throughout the state. We derived predictor variables from general ecological concepts: (1) the theory of island biogeography that insect species richness increases as host plant geographic area increases and as local abundance increases, (2) the plant-architecture hypothesis that insect species richness increases with increased plant size, and (3) phytochemical patterns in leaves, including nutrients and digestibility reducers predicting suitability for insect herbivores. Concepts 1 and 2, developed for large scales and species numbers, were tested at smaller scales relevant to much conservation research and management. A stepwise multiple regression including all predictor variables accounted for 99% of the variance in cynipid species richness with three variables: foliar hemicellulose concentration (81%), host geographic area (16%), and tree height (2%). The trends were negative, however, and opposite to those predicted by concepts 1 and 2. Ecological theory was not applicable to discovery of predictors of cynipid species richness on six oak species. Thus, we promote caution in applying ecological theory to a narrow set of species without specific testing of how patterns conform to theoretical predictions.     

Local Species Richness of Leaf-Chewing Insects Feeding on Woody Plants from One Hectare of a Lowland Rainforest

Abstract:  Local species diversity of insect herbivores feeding on rainforest vegetation remains poorly known. This ignorance limits evaluation of species extinction patterns following various deforestation scenarios. We studied leaf-chewing insects feeding on 59 species of woody plants from 39 genera and 18 families in a lowland rainforest in Papua New Guinea and surveyed all plants with a stem diameter at breast height of ≥5 cm in a 1-ha plot within the same area. We used two extrapolation methods, based on randomized species-accumulation curves, to combine these two data sets and estimate the number of species of leaf-chewing herbivores feeding on woody plants from the 1-ha area. We recorded 58,483 feeding individuals from 940 species of leaf-chewing insects. The extrapolation estimated that there were 1567–2559 species of leaf-chewing herbivores feeding on the 152 plant species from 97 genera and 45 families found in 1 ha of the forest. Most of the herbivore diversity was associated with plant diversity on the familial and generic levels. We predicted that, on average, the selection of 45 plant species each representing a different family supported 39% of all herbivore species, the 52 plant species each representing a different additional genus from these families supported another 39% of herbivore species, and the remaining 55 plant species from these genera supported 22% of herbivore species. Lepidoptera was the most speciose taxon in the local fauna, followed by Coleoptera and orthopteroids (Orthoptera and Phasmatodea). The ratio of herbivore to plant species and the estimated relative species richness of the Lepidoptera, Coleoptera, and orthopteroids remained constant on the spatial scale from 0.25 to 1 ha. However, the utility of local taxon-to-taxon species ratios for extrapolations to geographic scales requires further study.     

Birds in agricultural mosaics: the influence of landscape pattern and countryside heterogeneity.

Agricultural environments are critical to the conservation of biota throughout the world. Efforts to identify key influences on the conservation status of fauna in such environments have taken complementary approaches. Many studies have focused on the role of remnant or seminatural vegetation and emphasized the influence on biota of spatial patterns in the landscape. Others have recognized that many species use diverse "countryside" elements within farmland, and emphasize the benefits of landscape heterogeneity for conservation. Here, we investigated the effect of independent measures of both the spatial pattern (extent and configuration) and heterogeneity of elements (i.e., land uses/vegetation types) on bird occurrence in farm-scale agricultural mosaics in southeastern Australia. Birds were sampled in all types of elements in 27 mosaics (each 1 x 1 km) selected to incorporate variation in cover of native vegetation and the number of different element types in the mosaic. We used an information-theoretic approach to identify the mosaic properties that most strongly influenced bird species richness. Subgroups of birds based on habitat requirements responded most strongly to the extent of preferred elements in mosaics. Woodland birds were richer in mosaics with higher cover of native vegetation while open-tolerant species responded to the extent of scattered trees. In contrast, for total species richness, mosaic heterogeneity (richness of element types) and landscape context (cover of native vegetation in surrounding area) had the greatest influence. These results showed that up to 76% of landscape-level variation in richness of bird groups is attributable to mosaic properties directly amenable to management by landowners. Key implications include (1) conservation goals for farm landscapes must be carefully defined because the richness of different faunal components is influenced by different mosaic properties; (2) the extent of native vegetation is a critical influence in agricultural environments because it drives the farm-scale richness of woodland birds and has a broader context effect on total bird richness in mosaics; (3) land-use practices that enhance the heterogeneity of farmland mosaics are beneficial for native birds; and (4) the cumulative effect of even small elements in farm mosaics contribute to the structural properties of entire landscapes.     

Latitudinal variation in abundance of herbivorous fishes: a comparison of temperate and tropical reefs

The aim of the study was to provide comparable estimates of abundance of herbivorous reef fishes at temperate and tropical localities using a standardized methodology. Faunas of herbivorous fish were sampled on the rocky reefs of temperate northern New Zealand and on the coral reefs of the northern Great Barrier Reef (GBR), Australia, and the San Blas Archipelago in the Caribbean. A pilot study established the most appropriate habitat setting and the scale and magnitude of replication for the sampling program in temperate waters. Herbivorous fishes, including members of families endemic to the southern hemisphere (Odacidae and Aplodactylidae), were most abundant in turbulent, shallow water (0 to 6 m) and had patchy distributions within this habitat. A hierarchical sampling program using 10-min transect counts within the 0 to 6 m depth stratum examined abundance patterns at a range of spatial scales including mainland and island coasts, localities separated by up to 100 km and sites separated by up to 10 km. This program identified a characteristic fauna of seven species of herbivorous fishes with mean total abundances ranging from 23 to 30 individuals per 10-min transect. Species composition of the fauna varied between islands and coasts. A similar methodology was used to sample the major families of herbivorous fish in a number of sites in each of the tropical regions. These sampling programs revealed a fauna dominated by acanthurids and scarids in both the GBR and Caribbean localities. Estimates of abundance from these regions were similar, with a mean of 108 individuals recorded on the GBR and 129 per 10-min transect in the Caribbean. Species richness varied between each region, with 44 taxa recorded from the GBR and 11 from the Caribbean. Abundances of temperate water herbivores in New Zealand were found to be 75 to 80% lower than those recorded from shallow water habitats sampled on coral reefs. This was not related to species richness, since both New Zealand and the Caribbean locality had patterns of low richness. We suggest that the differences in abundance found by our study between temperate and tropical regions are not restricted to herbivorous fishes, but are representative of general latitudinal trends in reef fish faunas. Received: 4 November 1996 / Accepted: 15 December 1996     

Spatial scale and species identity influence the indigenous-alien diversity relationship in springtails

Although theory underlying the invasion paradox, or the change in the relationship between the richness of alien and indigenous species from negative to positive with increasing spatial scale, is well developed and much empirical work on the subject has been undertaken, most of the latter has concerned plants and to a lesser extent marine invertebrates. Here we therefore examine the extent to which the relationships between indigenous and alien species richness change from the local metacommunity to the interaction neighborhood scales, and the influences of abundance, species identity, and environmental favorability thereon, in springtails, a significant component of the soil fauna. Using a suite of modeling techniques, including generalized least squares and geographically weighted regressions to account for spatial autocorrelation or nonstationarity of the data, we show that the abundance and species richness of both indigenous and alien species at the metacommunity scale respond strongly to declining environmental favorability, represented here by altitude. Consequently, alien and indigenous diversity covary positively at this scale. By contrast, relationships are more complex at the interaction neighborhood scale, with the relationship among alien species richness and/or density and the density of indigenous species varying between habitats, being negative in some, but positive in others. Additional analyses demonstrated a strong influence of species identity, with negative relationships identified at the interaction neighborhood scale involving alien hypogastrurid springtails, a group known from elsewhere to have negative effects on indigenous species in areas where they have been introduced. By contrast, diversity relationships were positive with the other alien species. These results are consistent with both theory and previous empirical findings for other taxa, that interactions among indigenous and alien species change substantially with spatial scale and that environmental favorability may play a key role in explaining the larger scale patterns. However, they also suggest that the interactions may be affected by the identity of the species concerned, especially at the interaction neighborhood scale.     

Spatial location dominates over host plant genotype in structuring an herbivore community

Recent work has shown a potential role for both host plant genotype and spatial context in structuring insect communities. In this study, we use three separate data sets on herbivorous insects on oak (Quercus robur) to estimate the relative effects of host plant genotype (G), location (E), and the G x E interaction on herbivore community structure: a common garden experiment replicated at the landscape scale (approximately 5 km2); two common gardens separated at the regional scale (approximately 10 000 km2); and survey data on wild trees in various spatial settings. Our experiments and survey reveal that, at the landscape scale, the insect community is strongly affected by the spatial setting, with 32% of the variation in species richness explained by spatial connectivity. In contrast, G and G x E play minor roles in structuring the insect community. Results remained similar when extending the spatial scale of the study from the more local (landscape) level to the regional level. We conclude that in our study system, spatial processes play a major role in structuring these insect communities at both the landscape and regional scales, whereas host plant genotype seems of secondary importance.     

Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Spatial clustering of habitat structure effects patterns of community composition and diversity

Natural ecosystems often show highly productive habitats that are clustered in space. Environmental disturbances are also often nonrandomly distributed in space and are either intrinsically linked to habitat quality or independent in occurrence. Theoretical studies predict that configuration and aggregation of habitat patch quality and disturbances can affect metacommunity composition and diversity, but experimental evidence is largely lacking. In a metacommunity experiment, we tested the effects of spatially autocorrelated disturbance and spatial aggregation of patch quality on regional and local richness, among-community dissimilarity, and community composition. We found that spatial aggregation of patch quality generally increased among-community dissimilarity (based on two measures of beta diversity) of communities containing protozoa and rotifers in microcosms. There were significant interacting effects of landscape structure and location of disturbances on beta diversity, which depended in part on the specific beta diversity measures used. Effects of disturbance on composition and richness in aggregated landscapes were generally dependent on distance and connectivity among habitat patches of different types. Our results also show that effects of disturbances in single patches cannot directly be extrapolated to the landscape scale: the predictions may be correct when only species richness is considered, but important changes in beta diversity may be overlooked. There is a need for biodiversity and conservation studies to consider the spatial aggregation of habitat quality and disturbance, as well as connectivity among spatial aggregations.     

Proximity to forests drives bird conservation value of coffee plantations: implications for certification

Widespread loss of primary habitat in the tropics has led to increased interest in production landscapes for biodiversity conservation. In the Western Ghats biodiversity hotspot in India, shade coffee plantations are located in close proximity to sites of high conservation value: protected and unprotected forests. Coffee is grown here under a tree canopy that may be dominated by native tree species or by nonnative species, particularly silver oak (Grevillea robusta). We investigated the influence of properties at the local scale and the landscape scale in determining bird communities in coffee plantations, with particular emphasis on species of conservation priority. We used systematic point counts in 11 coffee plantation sites and analyzed data in a randomized linear modeling framework that addressed spatial autocorrelation. Greater proportion of silver oak at the local scale and distance to contiguous forests at the landscape scale were implicated as factors most strongly driving declines in bird species richness and abundance, while increased basal area of native tree species, a local-scale variable, was frequently related to increased bird species richness and abundance. The influence of local-scale variables increased at greater distances from the forest. Distance to forests emerged as the strongest predictor of declines in restricted-range species, with 92% reduction in the abundance of two commonly encountered restricted-range species (Pompadour Green Pigeon and Yellow-browed Bulbul) and a 43% reduction in richness of bird species restricted to Indian hill forests within 8 km of forests. Increase in proportion of silver oak from 33% to 55% was associated with 91% reduction in the abundance of one commonly encountered restricted-range species (Crimson-fronted Barbet). One conservation strategy is providing incentives to grow coffee in a biodiversity-friendly manner. One implication of our study is that plantations located at varying distances to the forest cannot be compared fairly for biodiversity friendliness by existing certification methodology. Another is that conservation of existing forests at the landscape scale is essential for maintaining higher biodiversity in coffee plantations. Incentive schemes that promote conservation of remnant forests at the landscape scale and biodiversity-friendly practices locally and that relate to coffee communities as a whole rather than individual planters are likely to be more effective.     

Regional variation in the hierarchical partitioning of diversity in coral-dwelling fishes

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.     

Elephants as agents of habitat creation for small vertebrates at the patch scale

Ecologists increasingly recognize the ability of certain species to influence ecological processes by engineering the physical environment, but efforts to develop a predictive understanding of this phenomenon are in their early stages. While many believe that the landscape-scale effects of ecosystem engineers will be to increase habitat diversity and therefore the abundance and richness of other species, few generalities exist about the effects of engineering at the scale of the engineered patch. According to one hypothesis, activities that increase structural habitat complexity within engineered patches will have positive effects on the abundance or diversity of other organisms. Here I show that, by damaging trees and increasing their structural complexity, browsing elephants create refuges used by a common arboreal lizard. Observational surveys and a lizard transplant experiment revealed that lizards preferentially occupy trees with real or simulated elephant damage. A second experiment showed that lizards vacate trees when elephant-engineered refuges are removed. Furthermore, local lizard densities increased with (and may be constrained by) local densities of elephant-damaged trees. This facilitative effect of elephants upon lizards via patch-scale habitat modification runs contrary to previously documented negative effects of the entire ungulate guild on lizards at the landscape scale, suggesting that net indirect effects of large herbivores comprise opposing trophic and engineering interactions operating at different spatial scales. Such powerful megaherbivore-initiated interactions suggest that anthropogenic changes in large-mammal densities will have important cascading consequences for ecological communities.     

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.