Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

Use of ecoacoustics to determine biodiversity patterns across ecological gradients

The variety of local animal sounds characterizes a landscape. We used ecoacoustics to noninvasively assess the species richness of various biotopes typical of an ecofriendly forest plantation with diverse ecological gradients and both nonnative and indigenous vegetation. The reference area was an adjacent large World Heritage Site protected area (PA). All sites were in a global biodiversity hotspot. Our results showed how taxa segregated into various biotopes. We identified 65 singing species, including birds, frogs, crickets, and katydids. Large, natural, protected grassland sites in the PA had the highest mean acoustic diversity (14.1 species/site). Areas covered in nonnative timber or grass species were devoid of acoustic species. Sites grazed by native and domestic megaherbivores were fairly rich (5.1) in acoustic species but none were unique to this habitat type, where acoustic diversity was greater than in intensively managed grassland sites (0.04). Natural vegetation patches inside the plantation mosaic supported high mean acoustic diversity (indigenous forests 7.6, grasslands 8.0, wetlands 9.1), which increased as plant heterogeneity and patch size increased. Indigenous forest patches within the plantation mosaic contained a highly characteristic acoustic species assemblage, emphasizing their complementary contribution to local biodiversity. Overall, acoustic signals determined spatial biodiversity patterns and can be a useful tool for guiding conservation.     

Influence of a dominant consumer species reverses at increased diversity

Theory and experiments indicate that changes in consumer diversity affect benthic community structure and ecosystem functioning. Although the effects of consumer diversity have been tested in the laboratory and the field, little is known about effects of consumer diversity in the subtidal zone, one of the largest marine habitats. We investigated the grazing effects of sea urchins on algal abundance and benthic community structure in a natural subtidal habitat of the Galápagos Islands. Three species of urchins (Eucidaris, Lytechinus, and Tripneustes) were manipulated in inclusion cages following a replacement design with three levels of species richness (one, two, and three species) with all possible two-species urchin combinations. Identity was the main factor accounting for changes in the percentage of substrate grazed and benthic community structure. Two out of the three two-species assemblages grazed more than expected, suggesting a richness effect, but analyses revealed that this increased grazing was due to a sampling effect of the largest and commercially valued urchin species, Tripneustes. Benthic community structure in treatments with Eucidaris, Lytechinus, and Tripneustes alone was significantly different at the end of the experiment, suggesting that resource use differentiation occurred. Communities in Tripneustes enclosures were characterized by abundant crustose coralline algae and grazed substrate, while those without it contained abundant green foliose algae (Ulva sp.). An unexpected emergent property of the system was that the most species-rich urchin assemblage underyielded, grazing less than any other assemblage with Tripneustes, effectively reversing its dominant influence observed in the two-species treatments. While further experiments are needed to discern the mechanisms of underyielding, it may be related to changing interspecific interactions as richness increases from two to three species or to density-dependent Tripneustes grazing. This study highlights the general importance of evaluating consumer richness effects across the entire range of species richness considered, as the performance of the most species-rich consumer assemblage could not be predicted by manipulations of intermediate levels of consumer species richness.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

Effects of Management of Domestic Dogs and Recreation on Carnivores in Protected Areas in Northern California

Abstract: In developed countries dogs (Canis lupus familiaris) are permitted to accompany human visitors to many protected areas (e.g., >96% of protected lands in California, U.S.A.), and protected‐area management often focuses on regulating dogs due to concerns about predation, competition, or transmission of disease and conflicts with human visitors. In 2004 and 2005, we investigated whether carnivore species richness and abundance were associated with management of domestic dogs and recreational visitation in protected areas in northern California. We surveyed for mammalian carnivores and human visitors in 21 recreation areas in which dogs were allowed offleash or onleash or were excluded, and we compared our observations in the recreation areas with observations in seven reference sites that were not open to the public. Carnivore abundance and species richness did not differ among the three types of recreation areas, but native carnivore species richness was 1.7 times greater (p < 0.01) and the relative abundances of native coyotes (Canis latrans) and bobcats (Lynx rufus) were over four times greater (p < 0.01) in the reference sites. Abundances of bobcats and all carnivores declined as the number of visitors increased. The policy on domestic dogs did not appear to affect species richness and abundance of mammalian carnivores. But the number of dogs we observed was strongly associated with human visitation (R²= 0.54), so the key factors associated with recreational effects on carnivores appear to be the presence and number of human visitors to protected areas.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

Marine reserves reestablish lost predatory interactions and cause community changes in rocky reefs.

In the last decades, marine reserves have dramatically increased in number worldwide. Here I examined the potential of no-take marine reserves to reestablish lost predatory interactions and, in turn, cause community-wide changes in Mediterranean rocky reefs. Protected locations supported higher density and size of the most effective fish preying on sea urchins (the sea breams Diplodus sargus and D. vulgaris) than unprotected locations. Density of sea urchins (Paracentrotus lividus and Arbacia lixula) was lower at protected than at unprotected locations. Size structure of P. lividus was bimodal (a symptom of predation on medium-sized urchins) only at the protected locations. Coralline barrens were less extended at protected than at unprotected locations, whereas turf-forming and erect-branched algae showed an opposite pattern. Erect-unbranched and erect-calcified algae and conspicuous zoobenthic organisms did not show any pattern related to protection. Tethering experiments showed that predation impact on urchins was (1) higher at protected than at unprotected locations, (2) higher on P. lividus than on A. lixula, and (3) higher on medium-sized (2-3.5 cm test diameter) than large-sized (>3.5 cm) urchins. Sea urchins preyed on by fish in natural conditions were smaller at unprotected than at protected locations. The analysis of sea urchin remains found in Diplodus fish stomachs revealed that medium-sized P. lividus were the most frequently preyed upon urchins and that size range of consumed sea urchins expanded with increasing size of Diplodus fish. These results suggest that (1) depletion and size reduction of predatory fish caused by fishing alter patterns of predation on sea urchins, and that (2) fishing bans (e.g., within no-take marine reserves) may reestablish lost interactions among strongly interactive species in temperate rocky reefs with potential community-wide effects.     

Metropolitan Open-Space Protection with Uncertain Site Availability

Abstract:  Urban planners acquire open space to protect natural areas and provide public access to recreation opportunities. Because of limited budgets and dynamic land markets, acquisitions take place sequentially depending on available funds and sites. To address these planning features, we formulated a two-period site selection model with two objectives: maximize the expected number of species represented in protected sites and maximize the expected number of people with access to protected sites. These objectives were both maximized subject to an upper bound on area protected over two periods. The trade-off between species representation and public access was generated by the weighting method of multiobjective programming. Uncertainty was represented with a set of probabilistic scenarios of site availability in a linear-integer formulation. We used data for 27 rare species in 31 candidate sites in western Lake County, near the city of Chicago, to illustrate the model. Each trade-off curve had a concave shape in which species representation dropped at an increasing rate as public accessibility increased, with the trade-off being smaller at higher levels of the area budget. Several sites were included in optimal solutions regardless of objective function weights, and these core sites had high species richness and public access per unit area. The area protected in period one depended on current site availability and on the probabilities of sites being undeveloped and available in the second period. Although the numerical results are specific for our study, the methodology is general and applicable elsewhere.     

Stronger predation in the tropics shapes species richness patterns in marine communities

Species interactions are widely assumed to be stronger at lower latitudes, but surprisingly few experimental studies test this hypothesis, and none ties these processes to observed patterns of species richness across latitude. We report here the first experimental field test that predation is both stronger and has a disproportionate effect on species richness in the tropics relative to the temperate zone. We conducted predator-exclusion experiments on communities of sessile marine invertebrates in four regions, which span 32 degrees latitude, in the western Atlantic Ocean and Caribbean Sea. Over a three-month timescale, predation had no effect on species richness in the temperate zone. In the tropics, however, communities were from two to over ten times more species-rich in the absence of predators than when predators were present. While micro-and macro-predators likely compete for the limited prey resource in the tropics, micropredators alone were able to exert as much pressure on the invertebrate communities as the full predator community. This result highlights the extent to which exposure to even a subset of the predator guild can significantly impact species richness in the tropics. Patterns were consistent in analyses that included relative and total species abundances. Higher species richness in the absence of predators in the tropics was also observed when species occurrences were pooled across two larger spatial scales, site and region, demonstrating a consistent scaling relationship. These experimental results show that predation can both limit local species abundances and shape patterns of regional coexistence in the tropics. When preestablished diverse tropical communities were then exposed to predation for different durations, ranging from one to several days, species richness was always reduced. These findings confirmed that impacts of predation in the tropics are strong and consistent, even in more established communities. Our results offer empirical support for the long-held prediction that predation pressure is stronger at lower latitudes. Furthermore, we demonstrate the magnitude to which variation in predation pressure can contribute to the maintenance of tropical species diversity.     

Offsets and Conservation of the Species of the EU Habitats and Birds Directives

Biodiversity offsets are intended to achieve no net loss of biodiversity due to economic and human development. A variety of biodiversity components are addressed by offset policies. It is required that loss of protected species due to development be offset under the EU Habitats and Birds Directives in Europe. We call this type of offset a species‐equality offset because the offset pertains to the same species affected by the development project. Whether species equality can be achieved by offset design is unknown. We addressed this gap by reviewing derogation files (i.e., specific files that describe mitigation measures to ensure no net loss under the EU Habitats and Birds Directives) from 85 development projects in France (2009–2010). We collected information on type of effect (reversible vs. irreversible) and characteristics of affected and offset sites (i.e., types of species, total area). We analyzed how the type of effect and the affected‐site characteristics influenced the occurrence of offset measures. The proportion of species targeted by offset measures (i.e., offset species) increased with the irreversibility of the effect of development and the conservation status of the species affected by development (i.e., affected species). Not all effects on endangered species (International Union for Conservation of Nature Red List) were offset; on average, 82% of affected species would be offset. Twenty‐six percent of species of least concern were offset species. Thirty‐five percent of development projects considered all affected species in their offset measures. Species richness was much lower in offset sites than in developed sites even after offset proposals. For developed areas where species richness was relatively high before development, species richness at offset sites was 5–10 times lower. The species‐equality principle appears to have been applied only partially in offset policies, as in the EU directives. We suggest the application of this principle through offsets is highly important for the long‐term conservation of biodiversity in Europe. Compensaciones y Conservación de las Especies de las Directivas de Hábitats y Aves de la UE     

The effect of grazing on biodiversity in coastal dune heathlands

The effect of sheep grazing on species richness, higher order diversity measures, inequality, species composition, functional diversity and allometric relationships at a coastal dune heathland site was investigated. After a prescribed fire in 2002, the site was divided into two parts, where one of the parts was unmanaged and the other part was fenced and grazed by sheep. Sheep grazing had a positive effect on species richness as well as a significant positive effect on the functional diversity at the coastal dune heathland site. Generally, the cover of dwarf shrubs was negatively affected by grazing, whereas the cover of sedges and grasses was positively affected by grazing. There is a need for comparative investigations of the effect of different management methods not only on floristic biodiversity, but on all relevant kinds of biodiversity as well as on soil structure, soil chemistry and habitat micro- and macrostructure. Consequently, we advocate the initiation of an international systematic investigation of the effect of different management methods.     

Habitat as a mediator of mesopredator‐driven mammal extinction

A prevailing view in dryland systems is that mammals are constrained by the scarcity of fertile soils and primary productivity. An alternative view is that predation is a primary driver of mammal assemblages, especially in Australia, where 2 introduced mesopredators—feral cat (Felis catus) and red fox (Vulpes vulpes)—are responsible for severe declines of dryland mammals. We evaluated productivity and predation as drivers of native mammal assemblage structure in dryland Australia. We used new data from 90 sites to examine the divers of extant mammal species richness and reconstructed historic mammal assemblages to determine proportional loss of mammal species across broad habitat types (landform and vegetation communities). Predation was supported as a major driver of extant mammal richness, but its effect was strongly mediated by habitat. Areas that were rugged or had dense grass cover supported more mammal species than the more productive and topographically simple areas. Twelve species in the critical weight range (CWR) (35–5500 g) that is most vulnerable to mesopredator predation were extirpated from the continent's central region, and the severity of loss of species correlated negatively with ruggedness and positively with productivity. Based on previous studies, we expect that habitat mediates predation from red foxes and feral cats because it affects these species’ densities and foraging efficiency. Large areas of rugged terrain provided vital refuge for Australian dryland mammals, and we predict such areas will support the persistence of CWR species in the face of ongoing mammal declines elsewhere in Australia.     

Predicting Richness of Native, Rare, and Exotic Plants in Response to Habitat and Disturbance Variables across a Variegated Landscape

Species richness of native, rare native, and exotic understorey plants was recorded at 120 sites in temperate grassy vegetation in New South Wales. Linear models were used to predict the effects of environment and disturbance on the richness of each of these groups. Total native species and rare native species showed similar responses, with richness declining on sites of increasing natural fertility of parent material as well as declining under conditions of water enrichment (resulting from human-induced changes in drainage characteristics, leading to increased run-off), severe livestock grazing, and soil disturbance. The response of rare native species to water enrichment, however, was significantly greater than that of all native species. Exotic species richness varied in reverse to that of native species with positive responses to water enrichment and soil disturbance. The contrasting behaviors are attributed to differences in the evolutionary history of native and exotic assemblages and their resulting preadaptations to a landscape recently subjected to agricultural settlement. It would appear that for exogenous disturbances, the intermediate disturbance hypothesis is not supported by our data. In the sampled region, pastures represent the major land-use in terms of area, but have relatively low densities of native and rare species compared with more lightly grazed areas. However, their management is considered to be essential to the maintenance of diversity on a regional scale.     

Defensive mechanisms and ecology of some tropical holothurians

The ecology, defensive behavior and toxicity of three species of reef flat holothurians (Actinopyga mauritiana, Holothuria atra and Holothuria difficilis) were studied at Eniwetok Atoll, Marshall Islands. The average diurnal population density of H. difficilis ranged from 1.4 to 32 holothurians/900 cm²; resting respiratory rates (0.05 ml O₂/g wet wt/h) were comparable during day and night; nourishment in H. difficilis may be primarily from bacteria and foraminifera in which about 2% of the dry weight of sediment consumed is utilized, and the species probably passes at least 3 g dry wt of sediment/m²/day (>1 kg/m²/year). A fundamental difference in energy flow is suggested: considerably more energy is passed from benthic algae to grazing and browsing fishes to predatory fishes on coral reefs whereas, in extra-tropical latitudes, more energy is shunted from benthic algae to invertebrates to predators. The effects of holothurin leading to death in fishes are irreversible. Holothuria difficilis is best protected from predation. Its body wall is toxic and it can accurately eject Cuvierian tubules, which are also toxic. The discharge of tubules was regulated by a circadian rhythm in May. Studies on holothurians and sponges suggest that many exposed coral reef invertebrates have evolved effective defensive mechanisms in association with high intensity predation.Supported by A.E.C. Contract AT (29-2)-226 with the University of Hawaii.     

Incorporating geographical and evolutionary rarity into conservation prioritization

Key goals of conservation are to protect both species and the functional and genetic diversity they represent. A strictly species-based approach may underrepresent rare, threatened, or genetically distinct species and overrepresent widespread species. Although reserves are created for a number of reasons, including economic, cultural, and ecological reasons, their efficacy has been measured primarily in terms of how well species richness is protected, and it is useful to compare how well they protect other measures of diversity. We used Proteaceae species-occurrence data in the Cape Floristic Region of South Africa to illustrate differences in the spatial distribution of species and evolutionary diversity estimated from a new maximum-likelihood molecular phylogeny. We calculated species richness, phylogenetic diversity (i.e., summed phylogenetic branch lengths in a site), and a site-aggregated measure of biogeographically weighted evolutionary distinctiveness (i.e., an abundance weighted measure that captures the unique proportion of the phylogenetic tree a species represents) for sites throughout the Cape Floristic Region. Species richness and phylogenetic diversity values were highly correlated for sites in the region, but species richness was concentrated at a few sites that underrepresented the much more spatially extensive distribution of phylogenetic diversity. Biogeographically weighted evolutionary diversity produced a scheme of prioritization distinct from the other 2 metrics and highlighted southern sites as conservation priorities. In these sites, the high values of biogeographically weighted evolutionary distinctiveness were the result of a nonrandom relation between evolutionary distinctiveness and geographical rarity, where rare species also tended to have high levels of evolutionary distinctiveness. Such distinct and rare species are of particular concern, but are not captured by conservation schemes that focus on species richness or phylogenetic diversity alone.     

Representativeness and Efficiency of Bird and Insect Conservation in Norwegian Boreal Forest Reserves

I quantified local species richness of birds in different forest types and of beetles in spruce forests at different altitudes. In both cases I quantified timber production as a measure of land acquisition cost and used the ratio between the species richness and timber production as a measure of conservation cost-efficiency. I found a positive correlation between timber production and local species richness of birds as well as beetles, indicating that the forests most valuable for forestry are also the ones most valuable for biodiversity conservation. I used different selection procedures for combining sites in a reserve network to find the minimum set of sites that included all vulnerable species. The minimum set of sites for birds was 30% spruce forest, 30% pine forest, and 40% broad-leaved forest (the three main forest types). The minimum set of sites for the beetles was uniformly distributed along the altitudinal gradient. Both minimum sets were most cost-efficient for species conservation. I suggest that equal coverage of different productivity classes is more efficient for optimizing biodiversity conservation than over-representing low productivity sites. Less than 1% of Norwegian boreal forests have been protected as nature reserves. The reserve network is fairly representative with respect to altitude, but it is seriously skewed toward low productivity sites. The current network is suboptimal with respect to forest type representativeness, species protection, and cost-efficiency. This is a result of an inefficient strategy of selecting reserve sites and an unfortunate combination of selection criteria.     

Community assembly in epiphytic lichens in early stages of colonization

Colonization studies may function as natural experiments and have the potential of addressing important questions about community assembly. We studied colonization for a guild of epiphytic lichens in a former treeless heathland area of 170 km2 in southwest Norway. We investigated if epiphytic lichen species richness and composition on aspen (Populus tremula) trees corresponded to a random draw of lichen individuals from the regional species pool. We compared lichen communities of isolated young (55-120 yr) and old (140-200 yr) forest patches in the heathland area to those of aspen forest in an adjacent reference area that has been forested for a long time. All thalli (lichen bodies) of 32 selected lichen species on trunks of aspen were recorded in 35 aspen sites. When data for each site category (young, old, and reference) were pooled, we found the species richness by rarefaction to be similar for reference sites and old sites, but significantly lower for young sites. The depauperated species richness of young sites was accompanied by a skew in species composition and absence of several species that were common in the reference sites. In contrast, genetic variation screened with neutral microsatellite markers in the lichen species Lobaria pulmonaria showed no significant differences between site categories. Our null hypothesis of a neutral species assembly in young sites corresponding to a random draw from the regional species pool was rejected, whereas an alternative hypothesis based on differences in colonization capacity among species was supported. The results indicate that for the habitat configuration in the heathland area (isolated patches constituting < 0.4% of the area) lichen communities may need a colonization time of 100-150 yr for species richness to level off, but given enough time, isolation will not affect species richness. We suggest that this contradiction to expectations from classical island equilibrium theory results from low extinction rates.     

Analytical Solutions to Trade‐Offs between Size of Protected Areas and Land‐Use Intensity

Abstract: Land‐use change is affecting Earth's capacity to support both wild species and a growing human population. The question is how best to manage landscapes for both species conservation and economic output. If large areas are protected to conserve species richness, then the unprotected areas must be used more intensively. Likewise, low‐intensity use leaves less area protected but may allow wild species to persist in areas that are used for market purposes. This dilemma is present in policy debates on agriculture, housing, and forestry. Our goal was to develop a theoretical model to evaluate which land‐use strategy maximizes economic output while maintaining species richness. Our theoretical model extends previous analytical models by allowing land‐use intensity on unprotected land to influence species richness in protected areas. We devised general models in which species richness (with modified species‐area curves) and economic output (a Cobb–Douglas production function) are a function of land‐use intensity and the proportion of land protected. Economic output increased as land‐use intensity and extent increased, and species richness responded to increased intensity either negatively or following the intermediate disturbance hypothesis. We solved the model analytically to identify the combination of land‐use intensity and protected area that provided the maximum amount of economic output, given a target level of species richness. The land‐use strategy that maximized economic output while maintaining species richness depended jointly on the response of species richness to land‐use intensity and protection and the effect of land use outside protected areas on species richness within protected areas. Regardless of the land‐use strategy, species richness tended to respond to changing land‐use intensity and extent in a highly nonlinear fashion.     

Effects of Forest Fragmentation on a Dung Beetle Community in French Guiana

Abstract:  Fragmentation is the most common disturbance induced by humans in tropical forests. Some insect groups are particularly suitable for studying the effects of fragmentation on animal communities because they are taxonomically and ecologically homogenous. We investigated the effects of forest fragmentation on a dung beetle species community in the forest archipelago created in 1994–1995 by the dam of Petit Saut, French Guiana. We set and baited an equal number of pitfall traps for dung beetles on three mainland sites and seven island sites. The sites ranged from 1.1 to 38 ha. In 250 trap days, we captured 50 species in 19 genera. Diversity indices were high (2.18–4.06). The lowest diversity was on the small islands and one mainland site. Species richness and abundance were positively related to fragment area but not to distance from mainland or distance to the larger island. The islands had lower species richness and population than mainland forest, but rarefied species richness was relatively invariant across sites. There was a marked change in species composition with decreasing fragment that was not caused by the presence of a common fauna of disturbed-area species on islands. Small islands differed from larger islands, which did not differ significantly from mainland sites. Partial correlation analyses suggested that species richness and abundance of dung beetle species were positively related to the number of species of nonflying mammals and the density index of howler monkeys ( Alouatta seniculus ), two parameters positively related to fragment area.