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Overhauling ocean spatial planning to improve marine megafauna conservation

Tracking data have led to evidence-based conservation of marine megafauna, but a disconnect remains between the many thousands of individual animals that have been tracked and the use of these data in conservation and management actions. Furthermore, the focus of most conservation efforts is within Exclusive Economic Zones despite the ability of these species to move thousands of kilometres across multiple national jurisdictions. To assist the goal of the United Nations General Assembly’s recent effort to negotiate a global treaty to conserve biodiversity on the high seas, we propose the development of a new frontier in dynamic marine spatial management. We argue that a global approach combining tracked movements of marine megafauna and human activities at-sea, and using existing and emerging technologies (e.g., through new tracking devices and big data approaches) can be applied to deliver near real-time diagnostics on existing risks and threats to mitigate global risks for marine megafauna. With technology developments over the next decade expected to catalyse the potential to survey marine animals and human activities in ever more detail and at global scales, the development of dynamic predictive tools based on near real-time tracking and environmental data will become crucial to address increasing risks. Such global tools for dynamic spatial and temporal management will, however, require extensive synoptic data updates and will be dependent on a shift to a culture of data sharing and open access. We propose a global mechanism to store and make such data available in near real-time, enabling a holistic view of space use by marine megafauna and humans that would significantly accelerate efforts to mitigate impacts and improve conservation and management of marine megafauna.

Frontiers in Marine Science

From models to management: Implementing the climate-change refugia conservation cycle in national parks of the northeastern United States

Managing climate-change refugia is a commonly recommended strategy for conserving biodiversity. However, few efforts have moved beyond the modeling phase of refugia science to management. Here, we present two case studies that move beyond modeling to testing models and management. In the first case study, we model refugia for two plant species (three-toothed cinquefoil [ Sibbaldia tridentata ] and black crowberry [ Empetrum nigrum ]) in Acadia National Park, Maine, United States, and use greenhouse experiments, common-garden experiments, and participatory science to evaluate the output of those models. Our results suggest that three-toothed cinquefoil growth and survival are reduced under increased temperatures as models predict. However, other variables (e.g., soil moisture and salinity) might also be important to modeling and managing refugia for both species. National Park Service staff and partners have been directing restoration for both species to refugia, but are also exploring other adaptation strategies. In the second case study, we demonstrate that existing prioritization processes for habitat restoration in national parks of the northeastern United States rarely incorporate refugia for two indicator species: Jefferson salamander ( Ambystoma jeffersonianum ) and grasshopper sparrow ( Ammodramus savannarum ). Our work demonstrates that moving beyond modeling can improve models and lead to new management insights.

Northeast United States

Genetic tagging in the Anthropocene: Scaling ecology from alleles to ecosystems

The Anthropocene is an era of marked human impact on the world. Quantifying these impacts 51 has become central to understanding the dynamics of coupled human-natural systems, resource52 dependent livelihoods, and biodiversity conservation. Ecologists are facing growing pressure to 53 quantify the size, distribution, and trajectory of wild populations in a cost-effective and socially54 acceptable manner. Genetic tagging, combined with modern computational and genetic analyses, 55 is an under-utilized tool to meet this demand, especially for wide-ranging, elusive, sensitive, and 56 low-density species. Genetic tagging studies are now revealing unprecedented insight into the 57 mechanisms that control the density, trajectory, connectivity and human-wildlife conflict for 58 populations over vast spatial scales. Here we outline the application of, and ecological inferences 59 from, new analytical techniques applied to genetically-tagged individuals, contrast this approach 60 with conventional methods, and describe how genetic tagging can be better applied to address 61 outstanding questions in ecology. We provide example analyses using a long-term genetic 62 tagging dataset of grizzly bears in the Canadian Rockies. The genetic tagging toolbox is a 63 powerful and overlooked ensemble that ecologists and conservation biologists can leverage to 64 generate evidence and meet the challenges of the Anthropocene.

Ecological Applications

Restoration based on cost-benefit optimization: A grasslands pilot study

Ecological restoration is essential to meeting global biodiversity conservation goals. Given limited conservation budgets, deciding where to restore habitat is a key challenge for the coming decade. We developed a spatially explicit framework to optimize ecological restoration site selection by integrating land use history, species distributions, and economic costs. The framework includes the following steps: identify potential restoration area based on relevant environmental measures like land use; identify species of interest; calculate restoration benefits by modeling habitat and climate suitability and estimating reduced extinction risk associated with restoring a particular land parcel based on a modified species–area relationship; aggregate benefits across species; and compare to parcel-level land acquisition costs. We applied linear programming to maximize conservation benefit/restoration cost ratios to identify optimized restoration sites. We illustrate this approach using a case study for highly threatened grassland ecosystems in the Great Plains region of Kansas, USA. We selected five grassland animal species (greater prairie chickens [ Tympanuchus cupido ], lesser prairie chickens [ Tympanuchus pallidicinctus ], swift fox [ Vulpes velox ], pronghorn [ Antilocapra americana ], and regal fritillary [ Speyeria idalia ]) as indicators of restoration benefit across taxa. For the indicator species that we chose, shortgrass and mixed-grass prairies had the highest conservation benefit to cost ratio. Setting a minimum restoration threshold for each habitat type allowed us to identify high-priority tallgrass prairie sites. Despite increasing interest in ecological restoration, optimizing restoration site selection is challenging because one must consider habitat features that do not currently exist. The modeling approach described here is flexible and can be updated for different ecosystems, species, and conservation priorities. We outline potential alterations that can be made in future analyses, depending on desired restoration goals.

Kansas

Intraspecific functional diversity of common species enhances community stability

Common species are fundamental to the structure and function of their communities and may enhance community stability through intraspecific functional diversity (iFD). We measured among-habitat and within-habitat iFD (i.e., among- and within-plant community types) of two common small mammal species using stable isotopes and functional trait dendrograms, determined whether iFD was related to short-term population stability and small mammal community stability, and tested whether spatially explicit trait filters helped explain observed patterns of iFD. Southern red-backed voles ( Myodes gapperi ) had greater iFD than deer mice ( Peromyscus maniculatus ), both among habitats, and within the plant community in which they were most abundant (their “primary habitat”). Peromyscus maniculatus populations across habitats differed significantly between years and declined 78% in deciduous forests, their primary habitat, as did the overall deciduous forest small mammal community. Myodes gapperi populations were stable across habitats and within coniferous forest, their primary habitat, as was the coniferous forest small mammal community. Generalized linear models representing internal trait filters (e.g., competition), which increase within-habitat type iFD, best explained variation in M. gapperi diet, while models representing internal filters and external filters (e.g., climate), which suppress within-habitat iFD, best explained P. maniculatus diet. This supports the finding that M. gapperi had higher iFD than P. maniculatus and is consistent with the theory that internal trait filters are associated with higher iFD than external filters. Common species with high iFD can impart a stabilizing influence on their communities, information that can be important for conserving biodiversity under environmental change.

Ecology and Evolution

Fijian sea krait behavior relates to fine-scale environmental heterogeneity in old-growth coastal forest: The importance of integrated land–sea management for protecting amphibious animals

The importance of terrestrial coastal ecosystems for maintaining healthy coral reef ecosystems remains understudied. Sea kraits are amphibious snakes that require healthy coral reefs for foraging, but little is known about their requirements of terrestrial habitats, where they slough their skin, digest prey, and breed. Using concurrent microclimate measurements and behavior surveys, we show that a small, topographically flat atoll in Fiji with coastal forest provides many microhabitats that relate to the behaviors of Yellow Lipped Sea Kraits, Laticauda colubrina . Microclimates were significantly related to canopy cover, leaf litter depth, and distance from the high-water mark (HWM). Sea kraits were almost exclusively observed in coastal forest within 30 m of the HWM. Sloughing of skins only occurred within crevices of mature or dying trees. Resting L. colubrina were significantly more likely to occur at locations with higher mean diurnal temperatures, lower leaf litter depths, and shorter distances from the HWM. On Leleuvia, behavior of L. colubrina therefore relates to environmental heterogeneity created by old-growth coastal forests, particularly canopy cover and crevices in mature and dead tree trunks. The importance of healthy coastal habitats, both terrestrial and marine, for L. colubrina suggests it could be a good flagship species for advocating integrated land-sea management. Furthermore, our study highlights the importance of coastal forests and topographically flat atolls for biodiversity conservation. Effective conservation management of amphibious species that utilize land- and seascapes is therefore likely to require a holistic approach that incorporates connectivity among ecosystems and environmental heterogeneity at all relevant scales.

Ecology and Evolution

Models combining multiple scales of inference capture hydrologic and climatic drivers of riparian tree distributions

Predicting species geographic distributions is key to managing invasive species, conserving biodiversity, and understanding species' environmental requirements. Species distribution models (SDMs) commonly focus on climatic predictors, but other environmental factors can also be essential, particularly for species with specialized habitats defined by hydrologic, topographic, or edaphic conditions (e.g., riparian, wetland, alpine, coastal, serpentine). Here, we demonstrate a novel approach for capturing strong effects of both hydrologic and climatic predictors in SDMs for riparian plants, by merging analyses targeted at environmental drivers within riparian ecosystems and across the western USA (3.8 × 10 6 km 2 ). We developed presence-background SDMs from five algorithms for three invasive riparian trees ( Tamarix ramossisima / chinensis [saltcedar], Elaeagnus angustifolia [Russian olive], and Ulmus pumila [Siberian elm]) and three native Populus spp. (cottonwoods). We used separate background datasets to develop models with different spatial scales of inference: (1) spatially filtered random points to represent available habitat across the study area and (2) target-group points from Salix (willow) occurrences to represent available riparian habitat. Random-background models captured hydrologic drivers of riparian tree distributions relative to the largely upland western USA, whereas Salix -background models captured climatic drivers within the context of riparian ecosystems. Combining predictions from the two backgrounds identified hydrologically suitable habitats within climatically suitable regions, resulting in fewer false “absences” than either background alone, improving predictions over previous SDMs, and providing more complete information to guide management decisions. Surprisingly, the predicted habitat for U. pumila , a newly recognized riparian invader, was as or more extensive than Populus deltoides / fremontii , T. ramossisima / chinensis , and E. angustifolia , the most common riparian tree complexes in the western USA. Watersheds constituting 20% of U. pumila predicted habitat contained no occurrence records, indicating high risk of future and unrecognized invasions. Combining models from random and ecosystem-specific target-group backgrounds may improve SDMs for species from many specialized habitats, providing a method to link predicted distributions to localized geographic features while capturing broad-scale climatic requirements.

western United States

Integrating community science and agency-collected monitoring data to expand monitoring capacity at large spatial scales

Monitoring species to better understand their status, ecology, and management needs is a major expense for agencies tasked with biodiversity conservation. Community science data have the potential to improve monitoring for minimal cost, given appropriate analytical frameworks. We describe a framework for integrating data from the eBird community science platform with agency-collected monitoring data using a multistate occupancy model. Our model accounts for the structural differences across datasets and allows for estimation of both occupancy and breeding probabilities. The framework was applied to Common Loons ( Gavia immer ) in Washington State. A total of 766 sites had observation effort, of which 713 sites had only eBird effort, 26 sites had only Washington Department of Fish and Wildlife (WDFW) effort, and 27 sites had both. We predicted that the probability of occupancy was only 0.07 (95% Bayesian credible interval, BCI = 0.02–0.51) at the 2324 sites in our sampling frame, though the probability that Common Loons were breeding at occupied sites was 0.95 (95% BCI = 0.71–1.00). We found that probability of occupancy was positively related to waterbody size (probability of a positive effect = 0.88) and negatively related to an index of human influence (probability of a negative effect = 0.94). We found that probability of breeding at occupied sites was positively related to tree canopy cover (0.86), negatively related to elevation (0.99), and negatively related to barren, scrub/shrub, and herbaceous land cover (0.98). We found that state agency biologists were 16 times more likely to detect breeding Common Loons at a site than were eBird users (0.94, 95% BCI = 0.78–0.99 for agency biologists vs. 0.08, 95% BCI = 0.06–0.10 for eBird users). However, the amount of effort expended by eBird users meant that they confirmed Common Loons at 94 sites while agency biologists confirmed them at just 24 sites, although evidence of reproduction was only contributed by agency biologists. Our results provide a better understanding of the distribution of Common Loons in Washington, while further demonstrating that community science data can be a valuable complement to agency-collected data, if appropriate frameworks are developed to integrate these data sources.

Ecosphere

Oases of the future? Evaluating springs as potential hydrologic refugia in drying climates

Springs in water-limited landscapes are biodiversity hotspots and keystone ecosystems, disproportionately influencing surrounding landscapes despite their often small areas. Some springs served as evolutionary refugia during previous climate drying, supporting relict species in isolated habitats. Understanding whether springs will provide hydrologic refugia from future climate change is important to biodiversity conservation but complicated by hydrologic variability among springs, data limitations, and multiple non-climate threats to groundwater-dependent ecosystems. Here, we present a conceptual framework for categorizing springs as potentially stable, relative, or transient hydrologic refugia in a drying climate. Clues about refugial capacity of springs can be assembled from diverse approaches, including citizen-science-powered ecohydrologic monitoring, remote sensing, landowner interviews, and environmental tracer analysis. Managers can integrate multiple lines of evidence to predict which springs may become future refugia for species of concern, strengthening the long-term effectiveness of springs conservation and restoration and informing climate adaptation for terrestrial and freshwater species.

Frontiers in Ecology and the Environment

An integrated sampling and analysis approach for improved biodiversity monitoring

Successful biodiversity conservation requires high quality monitoring data and analyses to ensure scientifically defensible policy, legislation, and management. Although monitoring is a critical component in assessing population status and trends, many governmental and non-governmental organizations struggle to develop and implement effective sampling protocols and statistical analyses because of the magnitude and diversity of species in conservation concern. In this article we describe a practical and sophisticated data collection and analysis framework for developing a comprehensive wildlife monitoring program that includes multi-species inventory techniques and community-level hierarchical modeling. Compared to monitoring many species individually, the multi-species approach allows for improved estimates of individual species occurrences, including rare species, and an increased understanding of the aggregated response of a community to landscape and habitat heterogeneity. We demonstrate the benefits and practicality of this approach to address challenges associated with monitoring in the context of US state agencies that are legislatively required to monitor and protect species in greatest conservation need. We believe this approach will be useful to regional, national, and international organizations interested in assessing the status of both common and rare species.

Environmental Management

Application of science-based restoration planning to a desert river system

Persistence of many desert river species is threatened by a suite of impacts linked to water infrastructure projects that provide human water security where water is scarce. Many desert rivers have undergone regime shifts from spatially and temporally dynamic ecosystems to more stable systems dominated by homogenous physical habitat. Restoration of desert river systems could aid in biodiversity conservation, but poses formidable challenges due to multiple threats and the infeasibility of recovery to pre-development conditions. The challenges faced in restoring desert rivers can be addressed by incorporating scientific recommendations into restoration planning efforts at multiple stages, as demonstrated here through an example restoration project. In particular, use of a watershed-scale planning process can identify data gaps and irreversible constraints, which aid in developing achievable restoration goals and objectives. Site-prioritization focuses limited the resources for restoration on areas with the greatest potential to improve populations of target organisms. Investment in research to understand causes of degradation, coupled with adoption of a guiding vision is critical for identifying feasible restoration actions that can enhance river processes. Setting monitoring as a project goal, developing hypotheses for expected outcomes, and implementing restoration as an experimental design will facilitate adaptive management and learning from project implementation. Involvement of scientists and managers during all planning stages is critical for developing process-based restoration actions and an implementation plan to maximize learning. The planning process developed here provides a roadmap for use of scientific recommendations in future efforts to recover dynamic processes in imperiled riverine ecosystems.

San Rafael River watershed

Modeling and estimating co-occurrence between the invasive Shiny Cowbird and its Puerto Rican hosts

Invasive species threaten island biodiversity globally. For example, the Shiny Cowbird ( Molothrus bonariensis ) parasitizes many of Puerto Rico’s endemic species, particularly in the open forests in the island’s southwest. Less is known, however, about cowbird parasitism in the agro-ecological highlands, which contain a patchwork of forests, shaded-coffee plantations, and coffee farms without shade. In this paper, we estimated co-occurrence rates, a potential indicator of parasitism rates, between the cowbird and four host species across these three land uses, hypothesizing that cowbirds would most likely co-occur with their hosts in shaded-coffee farms. We also hypothesized that the presence of host species would increase the probability of cowbird occurrence. To investigate these hypotheses, we developed three Bayesian hierarchical occupancy models: one where the hosts and parasite occurred independently, one that used the latent host species richness as a predictor of cowbird occurrence, and one that used each latent host occurrence state as predictors. These methods addressed observation errors and appropriately propagated error to our predictions of co-occurrence rates. We selected the best performing model using WAIC, then used it to predict co-occurrence rates. While there was some evidence that host species richness increased the probability of cowbirds, the parsimonious model assumed no interaction. With this model, we found that cowbirds were more likely to overlap with certain hosts in shaded-coffee plantations. This may suggest increased parasitism at these plantations, potentially presenting challenges for managers who advocate for shade restoration to gain ecological services such as biodiversity conservation.

Biological Invasions

Population genetics reveals bidirectional fish movement across the Continental Divide via an interbasin water transfer

Interbasin water transfers are becoming an increasingly common tool to satisfy municipal and agricultural water demand, but their impacts on movement and gene flow of aquatic organisms are poorly understood. The Grand Ditch is an interbasin water transfer that diverts water from tributaries of the upper Colorado River on the west side of the Continental Divide to the upper Cache la Poudre River on the east side of the Continental Divide. We used single nucleotide polymorphisms to characterize population genetic structure in cutthroat trout ( Oncorhynchus clarkii ) and determine if fish utilize the Grand Ditch as a movement corridor. Samples were collected from two sites on the west side and three sites on the east side of the Continental Divide. We identified two or three genetic clusters, and relative migration rates and spatial distributions of admixed individuals indicated that the Grand Ditch facilitated bidirectional fish movement across the Continental Divide, a major biogeographic barrier. Previous studies have demonstrated ecological impacts of interbasin water transfers, but our study is one of the first to use genetics to understand how interbasin water transfers affect connectivity between previously isolated watersheds. We also discuss implications on native trout management and balancing water demand and biodiversity conservation.

Colorado

Integrated climate and land use change scenarios for California rangeland ecosystem services: wildlife habitat, soil carbon, and water supply

Context In addition to biodiversity conservation, California rangelands generate multiple ecosystem services including livestock production, drinking and irrigation water, and carbon sequestration. California rangeland ecosystems have experienced substantial conversion to residential land use and more intensive agriculture. Objectives To understand the potential impacts to rangeland ecosystem services, we developed six spatially explicit (250 m) climate/land use change scenarios for the Central Valley of California and surrounding foothills consistent with three Intergovernmental Panel on Climate Change emission scenario narratives. Methods We quantified baseline and projected change in wildlife habitat, soil organic carbon (SOC), and water supply (recharge and runoff). For six case study watersheds we quantified the interactions of future development and changing climate on recharge, runoff and streamflow, and precipitation thresholds where dominant watershed hydrological processes shift through analysis of covariance. Results The scenarios show that across the region, habitat loss is expected to occur predominantly in grasslands, primarily due to future development (up to a 37 % decline by 2100), however habitat loss in priority conservation errors will likely be due to cropland and hay/pasture expansion (up to 40 % by 2100). Grasslands in the region contain approximately 100 teragrams SOC in the top 20 cm, and up to 39 % of this SOC is subject to conversion by 2100. In dryer periods recharge processes typically dominate runoff. Future development lowers the precipitation value at which recharge processes dominate runoff, and combined with periods of drought, reduces the opportunity for recharge, especially on deep soils. Conclusion Results support the need for climate-smart land use planning that takes recharge areas into account, which will provide opportunities for water storage in dry years. Given projections for agriculture, more modeling is needed on feedbacks between agricultural expansion on rangelands and water supply.

California

Fire modulates climate change response of simulated aspen distribution across topoclimatic gradients in a semi-arid montane landscape

Content Changing aspen distribution in response to climate change and fire is a major focus of biodiversity conservation, yet little is known about the potential response of aspen to these two driving forces along topoclimatic gradients. Objective This study is set to evaluate how aspen distribution might shift in response to different climate-fire scenarios in a semi-arid montane landscape, and quantify the influence of fire regime along topoclimatic gradients. Methods We used a novel integration of a forest landscape succession and disturbance model (LANDIS-II) with a fine-scale climatic water deficit approach to simulate dynamics of aspen and associated conifer and shrub species over the next 150 years under various climate-fire scenarios. Results Simulations suggest that many aspen stands could persist without fire for centuries under current climate conditions. However, a simulated 2–5 °C increase in temperature caused a substantial reduction of aspen coverage at lower elevations and a modest increase at upper elevations, leading to an overall reduction of aspen range at the landscape level. Increasing fire activity may favor aspen increase at its upper elevation limits adjacent to coniferous forest, but may also favor reduction of aspen at lower elevation limits adjacent to xeric shrubland. Conclusions Our study highlights the importance of incorporating fine-scale terrain effects on climatic water deficit and ecohydrology when modeling species distribution response to climate change. This modeling study suggests that climate mitigation and adaptation strategies that use fire would benefit from consideration of spatial context at landscape scales.

Landscape Ecology

Effects of large-scale wetland loss on network connectivity of the Rainwater Basin, Nebraska

Context The Rainwater Basin region in south-central Nebraska supports a complex network of spatially-isolated wetlands that harbor diverse floral and faunal communities. Since European settlement, many wetlands have been lost from the network, which has increased distances among remaining wetlands. As a result, populations of wildlife species with limited dispersal capabilities may have become isolated and face greater local extinction risks. Objectives We compared the pre-European settlement and current extent of the Rainwater Basin network to assess the effects of wetland losses on network connectivity for a range of maximum dispersal distances. Methods We constructed network models for a range of maximum dispersal distances and calculated network metrics to assess changes in network connectivity and the relative importance of individual wetlands in regulating flow. Results Since European settlement, the number of wetlands in the Rainwater Basin has decreased by >90%. The average distance to the nearest neighboring wetland has increased by 150% to ~1.2 km, and the dispersal distance necessary to travel throughout the whole network has increased from 3.5 to 10.0 km. Last, relative importance of individual wetlands depended on the maximum dispersal distance. Which wetlands to preserve to maintain connectivity might therefore depend on the dispersal capabilities of the species or taxa of interest. Conclusions To preserve a broad range of biodiversity, conservation efforts should focus on preserving dense clusters of wetlands at fine spatial scales to maintain current levels of network connectivity, and restoring connections between clusters to facilitate long-range dispersal of species with limited dispersal capabilities.

Nebraska

Aquatic-macroinvertebrate communities of Prairie-Pothole wetlands and lakes under a changed climate

Understanding how aquatic-macroinvertebrate communities respond to changes in climate is important for biodiversity conservation in the Prairie Pothole Region and other wetland-rich landscapes. We sampled macroinvertebrate communities of 162 wetlands and lakes previously sampled from 1966 to 1976, a much drier period compared to our 2012–2013 sampling timeframe. To identify possible influences of a changed climate and predation pressures on macroinvertebrates, we compared two predictors of aquatic-macroinvertebrate communities: ponded-water dissolved-ion concentration and vertebrate-predator presence/abundance. Further, we make inferences of how macroinvertebrate communities were structured during the drier period when the range of dissolved-ion concentrations was much greater and fish occurrence in aquatic habitats was rare. We found that aquatic-macroinvertebrate community structure was influenced by dissolved-ion concentrations through a complex combination of direct and indirect relationships. Ion concentrations also influenced predator occurrence and abundance, which indirectly affected macroinvertebrate communities. It is important to consider both abiotic and biotic gradients when predicting how invertebrate communities will respond to climate change. Generally, in the wetlands and lakes we studied, freshening of ponded water resulted in more homogenous communities than occurred during a much drier period when salinity range among sites was greater.

Wetlands

Responses to water depth and clipping of twenty−three plant species in an Indian monsoonal wetland

Responses of species to disturbances give insights into how species might respond to future wetland changes. In this study, species of monsoonal wetlands belonging to various functional types (graminoid and non−graminoid emergents, submersed aquatic, floating−leaved aquatic) varied in their growth responses to water depth and harvesting. We tested the effects of water depth (moist soil, flooded) and clipping (unclipped, and clipped) on the biomass and longevity of twenty−three dominant plant species of monsoonal wetlands in the Keoladeo National Park, India in a controlled experiment. With respect to total biomass and survival, six species responded positively to flooding and twelve species responded negatively to clipping. Responses to flooding and clipping, however, sometimes interacted. Individualistic responses of species to water levels and clipping regimes were apparent; species within a functional group did not always respond similarly. Therefore, detailed information on the individualistic responses of species may be needed to predict the vegetation composition of post−disturbance wetlands. In particular, as demands for fresh water increase around the world, studies of life history constraints and responses to hydrological changes will aid wetland managers in developing strategies to conserve biodiversity.

Keoladeo National Park