Effects of land use,cover, and protection on stream and riparian ecosystem services and biodiversity

Protected areas are an important part of broader landscapes that are often used to preserve biodiversity or natural features. Some argue that protected areas may also help ensure provision of ecosystem services. However, the effect of protection on ecosystem services and whether protection affects the provision of ecosystem services is known only for a few services in a few types of landscapes. We sought to fill this gap by investigating the effect of watershed protection status and land use and land cover on biodiversity and the provision of ecosystem services. We compared the ecosystem services provided in and around streams in 4 watershed types: International Union for Conservation of Nature category II protected forests, unprotected forests, unprotected forests with recent timber harvesting, and unprotected areas with agriculture. We surveyed 28 streams distributed across these watershed types in Quebec, Canada, to quantify provisioning of clean water, carbon storage, recreation, wild foods, habitat quality, and terrestrial and aquatic biodiversity richness and abundance. The quantity and quality of ecosystem services and biodiversity were generally higher in sites with intact forest—whether protected or not—relative to those embedded in production landscapes with forestry or agriculture. Clean-water provision, carbon storage, habitat quality, and tree diversity were significantly higher in and around streams surrounded by forest. Recreation, wild foods, and aquatic biodiversity did not vary among watershed types. Although some services can be provided by both protected and unprotected areas, protection status may help secure the continued supply of services sensitive to changes in land use or land cover. Our findings provide needed information about the ecosystem service and biodiversity trade-offs and synergies that result from developing a watershed or from protecting it.     

The Significance of Protected Areas for the Land Birds of Southern Finland

Protected areas (nature reserves) cover 0.7% of the land area in southern Finland (162,000 km²), which belongs to the south- and mid-boreal coniferous forest zones. The value of these areas in preserving land birds was studied on the basis of quantitative censuses. The numbers of bird pairs were estimated in nature reserves and in the whole of southern Finland, and the significance of the reserves was evaluated based on how large a proportion of the total population in southern Finland was found in them. Forest habitat generalists and species of coniferous forests occurred in protected areas as expected by the proportion of these areas in southern Finland. Specimens of these species groups comprise two thirds of all the country's land birds. Species of old-growth forests and open peatlands clearly preferred protected areas, whereas species of bushes, and lush and deciduous forest were scarcer in protected areas than elsewhere in southern Finland. The latter fertile habitats are poorly represented in protected areas, even though due to their disappearance or alteration elsewhere several species confined to these habitats have declined and are even threatened. Drainage of open peatlands and clearcutting of old-growth forests has caused a decrease in the area of these habitats in southern Finland during the past decades. Therefore, protected areas have a high significance in preserving bird species preferring these habitats. Protected areas are particularly important for decreased species of old-growth forests, such as the Three-toed Woodpecker Picoides tridactylus and the Siberian Jay Perisoreus infaustus , as about 10% of their total population in southern Finland was estimated to breed in protected areas. The future protection of both old-growth and deciduous forests is important if we are to preserve bio-diversity of land birds in Finland.     

How expert insight into alpine peatland conservation complements global scientific evidence

Experts can provide valuable information to fill knowledge gaps in published research on management effectiveness, particularly for threatened ecosystems, for which there is often limited evidence and the need for prompt intervention to ensure their persistence. One such ecosystem, alpine peatland, is threatened by climate change and other pressures, provides vital ecosystem services, and supports unique biodiversity. In a workshop, we gathered and synthesized into an accessible format information from experts on interventions used, threat context, and intervention effectiveness for Australian alpine peatland and used this knowledge to evaluate local relevance of the global literature for this threatened ecosystem. Experts identified 15 interventions used to conserve Australian peatlands, most of which enhanced or restored peatland condition and effectively addressed diverse threats. Experts’ perspectives and global studies were strongly aligned, suggesting that research on peatland management may be broadly relevant across contexts, despite the distinct characteristics of Australian systems. Our workshop-based expert elicitation approach provided insights into current management practices unavailable in the literature.     

Why We Need Megareserves in Amazonia

Abstract:  Brazilian Amazonia faces one of the greatest threats and opportunities for tropical biodiversity conservation of our times. I consider several large-scale issues in biodiversity conservation planning (e.g., resource extraction, large areas needed for top predators, species migration, fire, and carbon sequestration) in light of our severely deficient understanding of basinwide patterns of species distribution and little-known Amazonian biota. The long-term persistence of this biota is best served by strictly protected and sustainable development forest reserves that are both embedded in a benign forest matrix and sufficiently large to support a full complement of species and landscape-scale ecological processes. Given rapidly accelerating trends in agricultural frontier expansion into previously unclaimed public lands, protection and controlled development of forests is urgent.     

Identifying conservation priorities for threatened Eastern Himalayan mammals

To augment mammal conservation in the Eastern Himalayan region, we assessed the resident 255 terrestrial mammal species and identified the 50 most threatened species based on conservation status, endemism, range size, and evolutionary distinctiveness. By using the spatial analysis package letsR and the complementarity core‐area method in the conservation planning software Zonation, we assessed the current efficacy of their protection and identified priority conservation areas by comparing protected areas (PAs), land cover, and global ecoregion 2017 maps at a 100 × 100 m spatial scale. The 50 species that were most threatened, geographically restricted, and evolutionarily distinct faced a greater extinction risk than globally nonthreatened and wide‐ranging species and species with several close relatives. Small, medium‐sized, and data‐deficient species faced extinction from inadequate protection in PAs relative to wide‐ranging charismatic species. There was a mismatch between current PA distribution and priority areas for conservation of the 50 most endangered species. To protect these species, the skewed regional PA distribution would require expansion. Where possible, new PAs and transboundary reserves in the 35 priority areas we identified should be established. There are adequate remaining natural areas in which to expand current Eastern Himalayan PAs. Consolidation and expansion of PAs in the EH requires strengthening national and regional transboundary collaboration, formulating comprehensive regional land‐use plans, diversifying conservation funding, and enhancing information sharing through a consolidated regional database.     

Conservation of the Brazilian Cerrado

Abstract:  The Cerrado is one of the world's biodiversity hotspots. In the last 35 years, more than 50% of its approximately 2 million km² has been transformed into pasture and agricultural lands planted in cash crops. The Cerrado has the richest flora among the world's savannas (>7000 species) and high levels of endemism. Species richness of birds, fishes, reptiles, amphibians, and insects is equally high, whereas mammal diversity is relatively low. Deforestation rates have been higher in the Cerrado than in the Amazon rainforest, and conservation efforts have been modest: only 2.2% of its area is under legal protection. Numerous animal and plant species are threatened with extinction, and an estimated 20% of threatened and endemic species do not occur in protected areas. Soil erosion, the degradation of the diverse Cerrado vegetation formations, and the spread of exotic grasses are widespread and major threats. The use of fire for clearing land and to encourage new growth for pasture has also caused damage, even though the Cerrado is a fire-adapted ecosystem. Ecosystem experiments and modeling show that change in land cover is altering the hydrology and affecting carbon stocks and fluxes. Cerrado agriculture is lucrative, and agricultural expansion is expected to continue, requiring improvements in and extension of the transportation infrastructure, which will affect not only the Cerrado but also the Amazon forest. Large-scale landscape modification and threats to numerous species have led to renewed interest from various sectors in promoting the conservation of the Cerrado, particularly through strengthening and enlarging the system of protected areas and improving farming practices and thus the livelihoods of local communities.     

Conversion or conservation? Understanding wetland change in northwest Costa Rica.

Wetlands are more threatened than any other ecosystem type, with losses exceeding 50% of their original extent worldwide. Despite the small portion of the Earth's surface that they comprise, wetlands contribute significantly to global ecosystem services. In this study, we tested the hypothesis that the location and rate of change in wetland amount in the Tempisque Basin of northwest Costa Rica is predictable from landscape setting. Our results demonstrate that a strong potential exists for developing predictive models of wetland conversion based on an understanding of wetland location and surrounding trends of land use. We found that topography was the single most important predictor of wetland conversion in this area, entraining other conversion processes, and that spatial patterns of wetland loss could consistently be predicted from landscape-level variables. Areas with highest probabilities of conversion were found in the most accessible, non-protected regions of the landscape. While Palo Verde National Park made a substantial contribution to wetland conservation, our results highlight the dependence of lower-lying protected areas on upland processes, adding a little-addressed dimension of complexity to the dialogue about protected area management. Conservation strategies aimed at reducing wetland loss in tropical habitats will benefit from careful analysis of the dominant land use system(s) at a relatively broad scale, and the subsequent development of management and policy responses that take into account dynamic opportunities and constraints in the landscape.     

Ecological‐Niche Modeling and Prioritization of Conservation‐Area Networks for Mexican Herpetofauna

Abstract: One of the most important tools in conservation biology is information on the geographic distribution of species and the variables determining those patterns. We used maximum‐entropy niche modeling to run distribution models for 222 amphibian and 371 reptile species (49% endemics and 27% threatened) for which we had 34,619 single geographic records. The planning region is in southeastern Mexico, is 20% of the country's area, includes 80% of the country's herpetofauna, and lacks an adequate protected‐area system. We used probabilistic data to build distribution models of herpetofauna for use in prioritizing conservation areas for three target groups (all species and threatened and endemic species). The accuracy of species‐distribution models was better for endemic and threatened species than it was for all species. Forty‐seven percent of the region has been deforested and additional conservation areas with 13.7% to 88.6% more native vegetation (76% to 96% of the areas are outside the current protected‐area system) are needed. There was overlap in 26 of the main selected areas in the conservation‐area network prioritized to preserve the target groups, and for all three target groups the proportion of vegetation types needed for their conservation was constant: 30% pine and oak forests, 22% tropical evergreen forest, 17% low deciduous forest, and 8% montane cloud forests. The fact that different groups of species require the same proportion of habitat types suggests that the pine and oak forests support the highest proportion of endemic and threatened species and should therefore be given priority over other types of vegetation for inclusion in the protected areas of southeastern Mexico.     

Land-cover change within and around protected areas in a biodiversity hotspot

The landscape surrounding protected areas influences their ability to maintain ecosystem functions and achieve conservation goals. As anthropogenic intensification continues, it is important to monitor land-use and land-cover change in and around protected areas. We measure land-cover change surrounding protected areas in the Maputaland-Pondoland-Albany Biodiversity hotspot from the 1980s to present. Using Landsat imagery, we classified land cover within and around each protected area. Agricultural land uses were increasing and often directly border protected area boundaries. Human settlements increased around every protected area, potentially increasing human activity along the edges of protected areas and threatening their ecological integrity. Urban expansion around protected areas varied but increased as much as 10%. Woody vegetation cover varied both within and around protected areas with possible evidence of deforestation and shrub encroachment throughout the hotspot. We recommend monitoring land cover across southeastern Africa to better understand regional trends in land-use impacts to protected areas.     

Bias in protected‐area location and its effects on long‐term aspirations of biodiversity conventions

To contribute to the aspirations of recent international biodiversity conventions, protected areas (PAs) must be strategically located and not simply established on economically marginal lands as they have in the past. With refined international commitments under the Convention on Biological Diversity to target protected areas in places of “importance to biodiversity,” perhaps they may now be. We analyzed location biases in PAs globally over historic (pre‐2004) and recent periods. Specifically, we examined whether the location of protected areas are more closely associated with high concentrations of threatened vertebrate species or with areas of low agricultural opportunity costs. We found that both old and new protected areas did not target places with high concentrations of threatened vertebrate species. Instead, they appeared to be established in locations that minimize conflict with agriculturally suitable lands. This entrenchment of past trends has substantial implications for the contributions these protected areas are making to international commitments to conserve biodiversity. If protected‐area growth from 2004 to 2014 had strategically targeted unrepresented threatened vertebrates, >30 times more species (3086 or 2553 potential vs. 85 actual new species represented) would have been protected for the same area or the same cost as the actual expansion. With the land available for conservation declining, nations must urgently focus new protection on places that provide for the conservation outcomes outlined in international treaties.     

Carbon dioxide and methane emission from subsurface peatlands and its controlling factors北大核心CSCD

Peatland is an efficient carbon dioxide (CO2) sink on the continent and plays an important role in global carbon cycle. Climate change and human activities, two of the notable global environmental issues, have accelerated the degradation of peatlands during recent years. Global warming will increase the rate of aerobic decomposition in the surface of peatlands. Carbon stored in the subsurface of peatlands will be metabolized if the climatic conditions become favorable for decomposition. This study reviewed the carbon circle of subsurface peatland in natural environment and in environments disturbed by human activity or climate change. Furthermore, the major factors (environmental and human factors) that affect the carbon cycle were also discussed. According to a previous study, subsurface peatland will rapidly participate in the carbon cycle when the peatland is degraded. Water level, vegetation, and temperature were the main natural factors affecting the carbon cycle, whereas drainage, farming, and grazing were the main anthropogenic factors. Further studies should focus on different soil layer carbon dynamics, inorganic carbon content, and conservation and restoration of peatlands. The study methods should be a combination of macro with micro scale and focus on developing deep peat research techniques. Most of the previous studies focused on greenhouse gas emission and their response factors in short-term experiments. Thus, the mechanism and process of subsurface carbon are not clear and needs further study. © 2018 Science Press. All rights reserved.     

Conservation Businesses and Conservation Planning in a Biological Diversity Hotspot

The allocation of land to biological diversity conservation competes with other land uses and the needs of society for development, food, and extraction of natural resources. Trade‐offs between biological diversity conservation and alternative land uses are unavoidable, given the realities of limited conservation resources and the competing demands of society. We developed a conservation‐planning assessment for the South African province of KwaZulu‐Natal, which forms the central component of the Maputaland–Pondoland–Albany biological diversity hotspot. Our objective was to enhance biological diversity protection while promoting sustainable development and providing spatial guidance in the resolution of potential policy conflicts over priority areas for conservation at risk of transformation. The conservation‐planning assessment combined spatial‐distribution models for 646 conservation features, spatial economic‐return models for 28 alternative land uses, and spatial maps for 4 threats. Nature‐based tourism businesses were competitive with other land uses and could provide revenues of >US$60 million/year to local stakeholders and simultaneously help meeting conservation goals for almost half the conservation features in the planning region. Accounting for opportunity costs substantially decreased conflicts between biological diversity, agricultural use, commercial forestry, and mining. Accounting for economic benefits arising from conservation and reducing potential policy conflicts with alternative plans for development can provide opportunities for successful strategies that combine conservation and sustainable development and facilitate conservation action. Negocios de Conservación y Planificación de la Conservación en un Sitio de Importancia para la Biodiversidad     

Use of Inverse Spatial Conservation Prioritization to Avoid Biological Diversity Loss Outside Protected Areas

Globally expanding human land use sets constantly increasing pressure for maintenance of biological diversity and functioning ecosystems. To fight the decline of biological diversity, conservation science has broken ground with methods such as the operational model of systematic conservation planning (SCP), which focuses on design and on‐the‐ground implementation of conservation areas. The most commonly used method in SCP is reserve selection that focuses on the spatial design of reserve networks and their expansion. We expanded these methods by introducing another form of spatial allocation of conservation effort relevant for land‐use zoning at the landscape scale that avoids negative ecological effects of human land use outside protected areas. We call our method inverse spatial conservation prioritization. It can be used to identify areas suitable for economic development while simultaneously limiting total ecological and environmental effects of that development at the landscape level by identifying areas with highest economic but lowest ecological value. Our method is not based on a priori targets, and as such it is applicable to cases where the effects of land use on, for example, individual species or ecosystem types are relatively small and would not lead to violation of regional or national conservation targets. We applied our method to land‐use allocation to peat mining. Our method identified a combination of profitable production areas that provides the needed area for peat production while retaining most of the landscape‐level ecological value of the ecosystem. The results of this inverse spatial conservation prioritization are being used in land‐use zoning in the province of Central Finland.     

The Biogeography of Globally Threatened Seabirds and Island Conservation Opportunities

Seabirds are the most threatened group of marine animals; 29% of species are at some risk of extinction. Significant threats to seabirds occur on islands where they breed, but in many cases, effective island conservation can mitigate these threats. To guide island‐based seabird conservation actions, we identified all islands with extant or extirpated populations of the 98 globally threatened seabird species, as recognized on the International Union for Conservation of Nature Red List, and quantified the presence of threatening invasive species, protected areas, and human populations. We matched these results with island attributes to highlight feasible island conservation opportunities. We identified 1362 threatened breeding seabird populations on 968 islands. On 803 (83%) of these islands, we identified threatening invasive species (20%), incomplete protected area coverage (23%), or both (40%). Most islands with threatened seabirds are amenable to island‐wide conservation action because they are small (57% were <1 km²), uninhabited (74%), and occur in high‐ or middle‐income countries (96%). Collectively these attributes make islands with threatened seabirds a rare opportunity for effective conservation at scale. La Biogeografía de Aves Marinas Amenazadas Globalmente y las Oportunidades de Conservación en Islas     

Effect of Land Cover and Ecosystem Mapping on Ecosystem‐Risk Assessment in the Little Karoo,South Africa

Extinction‐risk assessments aim to identify biological diversity features threatened with extinction. Although largely developed at the species level, these assessments have recently been applied at the ecosystem level. In South Africa, national legislation provides for the listing and protection of threatened ecosystems. We assessed how land‐cover mapping and the detail of ecosystem classification affected the results of risk assessments that were based on extent of habitat loss. We tested 3 ecosystem classifications and 4 land‐cover data sets of the Little Karoo region, South Africa. Degraded land (in particular, overgrazed areas) was successfully mapped in just one of the land‐cover data sets. From <3% to 25% of the Little Karoo was classified as threatened, depending on the land‐cover data set and ecosystem classification applied. The full suite of threatened ecosystems on a fine‐scale map was never completely represented within the spatial boundaries of a coarse‐scale map of threatened ecosystems. Our assessments highlight the importance of land‐degradation mapping for the listing of threatened ecosystems. On the basis of our results, we recommend that when budgets are constrained priority be given to generating more‐detailed land‐cover data sets rather than more‐detailed ecosystem classifications for the assessment of threatened ecosystems. El Efecto de la Cobertura Terrestre y el Mapeo de Ecosistemas en la Valoración de Riesgos en los Ecosistemas en Little Karoo, Sudáfrica     

The Capacity of Australia's Protected‐Area System to Represent Threatened Species

Abstract: The acquisition or designation of new protected areas is usually based on criteria for representation of different ecosystems or land‐cover classes, and it is unclear how well‐threatened species are conserved within protected‐area networks. Here, we assessed how Australia's terrestrial protected‐area system (89 million ha, 11.6% of the continent) overlaps with the geographic distributions of threatened species and compared this overlap against a model that randomly placed protected areas across the continent and a spatially efficient model that placed protected areas across the continent to maximize threatened species’ representation within the protected‐area estate. We defined the minimum area needed to conserve each species on the basis of the species’ range size. We found that although the current configuration of protected areas met targets for representation of a given percentage of species’ ranges better than a random selection of areas, 166 (12.6%) threatened species occurred entirely outside protected areas and target levels of protection were met for only 259 (19.6%) species. Critically endangered species were among those with the least protection; 12 (21.1%) species occurred entirely outside protected areas. Reptiles and plants were the most poorly represented taxonomic groups, and amphibians the best represented. Spatial prioritization analyses revealed that an efficient protected‐area system of the same size as the current protected‐area system (11.6% of the area of Australia) could meet representation targets for 1272 (93.3%) threatened species. Moreover, the results of these prioritization analyses showed that by protecting 17.8% of Australia, all threatened species could reach target levels of representation, assuming all current protected areas are retained. Although this amount of area theoretically could be protected, existing land uses and the finite resources available for conservation mean land acquisition may not be possible or even effective for the recovery of threatened species. The optimal use of resources must balance acquisition of new protected areas, where processes that threaten native species are mitigated by the change in ownership or on‐ground management jurisdiction, and management of threatened species inside and outside the existing protected‐area system.     

Protecting biodiversity and economic returns in resource-rich tropical forests

In pursuit of socioeconomic development, many countries are expanding oil and mineral extraction into tropical forests. These activities seed access to remote, biologically rich areas, thereby endangering global biodiversity. We examined how protection of biodiversity and economic revenues can be balanced in biologically valuable regions. Using spatial data on oil profits and predicted species and ecosystem extents, we optimized the protection of 741 terrestrial species and 20 ecosystems of the Ecuadorian Amazon across a range of opportunity costs (i.e., sacrifices of extractive profit). We also applied spatial statistics to remotely sensed, historic deforestation data to focus the optimization on areas most threatened by imminent forest loss. Giving up 5% of a year's oil profits (US$221 million) allowed for a protected area network that retained an average of 65% of the extent of each species and ecosystem. This performance far exceeded that of the network produced by simple optimization for land area (which required a sacrifice of approximately 40% of annual oil profits [US$1.7 billion]) and used only marginally less land to achieve equivalent levels of ecological protection. We identified what we call emergency conservation targets: regions that are essential components of a cost-effective conservation reserve network but at imminent risk of destruction, thus requiring urgent and effective protection. Governments can use our methods when evaluating extractive-led development options to responsibly manage the associated ecological and economic trade-offs and protect natural capital.     

Intact Faunal Assemblages in the Modern Era

Abstract: In light of limited conservation funding, global conservation initiatives are increasingly focused on regions of the planet that have been identified as valuable on the basis of their species diversity, the vulnerability of resident species to extinction, or the perceived pristine nature of their ecosystems. Regions that have been resilient to high rates of extinction have not yet been systematically considered in conservation efforts. We used published range maps for 392 vertebrate species to compare historical and current species ranges. We used the results of the comparison to identify regions of the globe in which no known vertebrate species has been extirpated in the past 200 years. In 17 regions, no detectable vertebrate extinctions occurred in the past 200 years. In 6 other regions, reintroductions of species restored the full historic complement of vertebrate species. The effects of humans on a landscape, as measured by the human‐footprint index, although useful, was not a singularly good predictor of faunal intactness because more than 20% of intact land area was in heavily affected areas (50% of Earth's land area), and several regions where humans have had very little effect did not have intact faunas. Only 22% of intact land area was within protected‐area networks. High‐latitude areas were particularly underrepresented; they made up 3 of the 4 least‐protected areas in our analyses. Our results indicate that although protected areas are in some cases associated with the prevention of extinctions, there are many regions in which human activity coexists with intact vertebrate assemblages. In addition, our new approach for assessing the value of global regions for conservation identifies several regions that are not represented in other prioritization metrics.     

Effects of payments for ecosystem services on wildlife habitat recovery

Conflicts between local people's livelihoods and conservation have led to many unsuccessful conservation efforts and have stimulated debates on policies that might simultaneously promote sustainable management of protected areas and improve the living conditions of local people. Many government‐sponsored payments‐for‐ecosystem‐services (PES) schemes have been implemented around the world. However, few empirical assessments of their effectiveness have been conducted, and even fewer assessments have directly measured their effects on ecosystem services. We conducted an empirical and spatially explicit assessment of the conservation effectiveness of one of the world's largest PES programs through the use of a long‐term empirical data set, a satellite‐based habitat model, and spatial autoregressive analyses on direct measures of change in an ecosystem service (i.e., the provision of wildlife species habitat). Giant panda (Ailuropoda melanoleuca) habitat improved in Wolong Nature Reserve of China after the implementation of the Natural Forest Conservation Program. The improvement was more pronounced in areas monitored by local residents than those monitored by the local government, but only when a higher payment was provided. Our results suggest that the effectiveness of a PES program depends on who receives the payment and on whether the payment provides sufficient incentives. As engagement of local residents has not been incorporated in many conservation strategies elsewhere in China or around the world, our results also suggest that using an incentive‐based strategy as a complement to command‐and‐control, community‐ and norm‐based strategies may help achieve greater conservation effectiveness and provide a potential solution for the park versus people conflict.     

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.