On how much biodiversity is covered in Europe by national protected areas and by the Natura 2000 network: insights from terrestrial vertebrates

The European Union has made extensive biodiversity conservation efforts with the Habitats and Birds Directives and with the establishment of the Natura 2000 network of protected areas, one of the largest networks of conservation areas worldwide. We performed a gap analysis of the entire Natura 2000 system plus national protected areas and all terrestrial vertebrates (freshwater fish excluded). We also evaluated the level of connectivity of both systems, providing therefore a first estimate of the functionality of the Natura 2000 system as an effective network of protected areas. Together national protected areas and the Natura 2000 network covered more than one‐third of the European Union. National protected areas did not offer protection to 13 total gap species (i.e., species not covered by any protected area) or to almost 300 partial gap species (i.e., species whose representation target is not met). Together the Natura 2000 network and national protected areas left 1 total gap species and 121 partial gap species unprotected. The terrestrial vertebrates listed in the Habitats and Birds Directives were relatively well covered (especially birds), and overall connectivity was improved considerably by Natura 2000 sites that act as stepping stones between national protected areas. Overall, we found that the Natura 2000 network represents at continental level an important network of protected areas that acts as a good complement to existing national protected areas. However, a number of problems remain that are mainly linked to the criteria used to list the species in the Habitats and Birds Directives. The European Commission initiated in 2014 a process aimed at assessing the importance of the Birds and Habitats Directives for biodiversity conservation. Our results contribute to this assessment and suggest the system is largely effective for terrestrial vertebrates but would benefit from further updating of the species lists and field management.     

Terrestrial Reserve Networks Do Not Adequately Represent Aquatic Ecosystems

Abstract: Protected areas are a cornerstone of conservation and have been designed largely around terrestrial features. Freshwater species and ecosystems are highly imperiled, but the effectiveness of existing protected areas in representing freshwater features is poorly known. Using the inland waters of Michigan as a test case, we quantified the coverage of four key freshwater features (wetlands, riparian zones, groundwater recharge, rare species) within conservation lands and compared these with representation of terrestrial features. Wetlands were included within protected areas more often than expected by chance, but riparian zones were underrepresented across all (GAP 1–3) protected lands, particularly for headwater streams and large rivers. Nevertheless, within strictly protected lands (GAP 1–2), riparian zones were highly represented because of the contribution of the national Wild and Scenic Rivers Program. Representation of areas of groundwater recharge was generally proportional to area of the reserve network within watersheds, although a recharge hotspot associated with some of Michigan's most valued rivers is almost entirely unprotected. Species representation in protected areas differed significantly among obligate aquatic, wetland, and terrestrial species, with representation generally highest for terrestrial species and lowest for aquatic species. Our results illustrate the need to further evaluate and address the representation of freshwater features within protected areas and the value of broadening gap analysis and other protected‐areas assessments to include key ecosystem processes that are requisite to long‐term conservation of species and ecosystems. We conclude that terrestrially oriented protected‐area networks provide a weak safety net for aquatic features, which means complementary planning and management for both freshwater and terrestrial conservation targets is needed.     

Relationships between Plant and Animal Species Richness at a Regional Scale in China

Abstract:  Important questions in conservation biology and ecology include whether species diversities of different groups of organisms are correlated and, in particular, whether plant diversity influences animal diversity. I used correlation and partial regression analyses to examine the relationships between species richness of vascular plants and four major groups of terrestrial vertebrates (mammals, amphibians, reptiles, and birds) in 28 provinces in China. Species richness data were obtained from the literature. Environmental variables included normalized difference vegetation index, mean January temperature, mean annual temperature, annual precipitation, May through August precipitation, actual evapotranspiration, potential evapotranspiration, and elevation range. Species richness was strongly and positively correlated among the five groups of organisms. Plant richness was correlated with animal richness more strongly than the richness of different animal groups correlated with each other except for reptile richness, which had a slightly higher correlation with amphibian richness than with plant richness. Plant richness uniquely explained 41 times more variance in the species richness of the four vertebrate groups combined than environmental variables uniquely did, suggesting that plant richness influences terrestrial vertebrate richness at the regional scale examined. Because of strong correlations between the diversity of vascular plants and vertebrates, the diversity of vascular plants may be used as a surrogate for the diversity of terrestrial animals in China. My results have implications for selection of areas to be protected at both regional and local scales.     

Key Role of European Rabbits in the Conservation of the Western Mediterranean Basin Hotspot

Abstract: The Mediterranean Basin is a global hotspot of biodiversity. Hotspots are said to be experiencing a major loss of habitat, but an added risk could be the decline of some species having a special role in ecological relationships of the system. We reviewed the role of European rabbits (Oryctolagus cuniculus) as a keystone species in the Iberian Peninsula portion of the Mediterranean hotspot. Rabbits conspicuously alter plant species composition and vegetation structure through grazing and seed dispersal, which creates open areas and preserves plant species diversity. Moreover, rabbit latrines have a demonstrable effect on soil fertility and plant growth and provide new feeding resources for many invertebrate species. Rabbit burrows provide nest sites and shelter for vertebrates and invertebrates. In addition, rabbits serve as prey for a number of predators, including the critically endangered Iberian lynx (Lynx pardinus) and Spanish Imperial Eagle (Aquila adalberti). Thus, the Mediterranean ecosystem of the Iberian Peninsula should be termed “the rabbit's ecosystem.” To our knowledge, this is the first empirical support for existence of a multifunctional keystone species in a global hotspot of biodiversity. Rabbit populations have declined drastically on the Iberian Peninsula, with potential cascading effects and serious ecological and economic consequences. From this perspective, rabbit recovery is one of the biggest challenges for conservation of the Mediterranean Basin hotspot.     

Incorporating connectivity into conservation planning for the optimal representation of multiple species and ecosystem services

Conservation planning tends to focus on protecting species’ ranges or landscape connectivity but seldom both—particularly in the case of diverse taxonomic assemblages and multiple planning goals. Therefore, information on potential trade-offs between maintaining landscape connectivity and achieving other conservation objectives is lacking. We developed an optimization approach to prioritize the maximal protection of species’ ranges, ecosystem types, and forest carbon stocks, while also including habitat connectivity for range-shifting species and dispersal corridors to link protected area. We applied our approach to Sabah, Malaysia, where the state government mandated an increase in protected-area coverage of approximately 305,000 ha but did not specify where new protected areas should be. Compared with a conservation planning approach that did not incorporate the 2 connectivity features, our approach increased the protection of dispersal corridors and elevational connectivity by 13% and 21%, respectively. Coverage of vertebrate and plant species’ ranges and forest types were the same whether connectivity was included or excluded. Our approach protected 2% less forest carbon and 3% less butterfly range than when connectivity features were not included. Hence, the inclusion of connectivity into conservation planning can generate large increases in the protection of landscape connectivity with minimal loss of representation of other conservation targets.     

Past and Present Effectiveness of Protected Areas for Conservation of Naturally and Anthropogenically Rare Plant Species

Abstract:  The Global Strategy of Plant Conservation states that at least 60% of threatened plant species should be within protected areas. This goal has been met in some regions with long traditions of plant protection. We used gap analysis to explore how particular groups of species of conservation interest, representing different types of natural or anthropogenic rarity, have been covered by protected areas on a national scale in Estonia during the last 100 years. Species-accumulation curves indicated that plant species that are naturally rare (restricted global or local distribution, always small populations, or very rare habitat requirements) needed almost twice as many protected areas to reach the 60% target as plant species that are rare owing to lack of suitable management (species depending on grassland management, moderate forest disturbances, extensive traditional agriculture, or species potentially threatened by collecting). Temporal analysis of the establishment of protected areas suggested that grouping plant species according to the predominant cause of rarity accurately reflected the history of conservation decision making. Species found in very rare habitats have previously received special conservation attention; species dependent on traditional extensive agriculture have been largely ignored until recently. Legislative initiative and new nature-protection schemes (e.g., Natura 2000, network of protected areas in the European Union) have had a positive influence on all species groups. Consequently, the species groups needing similar action for their conservation are sensitive indicators of the effectiveness of protected-area networks. Different species groups, however, may not be uniformly conserved within protected areas, and all species groups should fulfill the target of 60% coverage within protected areas.     

Global gap analysis of cactus species and priority sites for their conservation

Knowing how much biodiversity is captured by protected areas (PAs) is important to meeting country commitments to international conservation agreements, such as the Convention on Biological Diversity, and analyzing gaps in species coverage by PAs contributes greatly to improved locating of new PAs and conservation of species. Regardless of their importance, global gap analyses have been conducted only for a few taxonomic groups (e.g., mangroves, corals, amphibians, birds, mammals). We conducted the first global gap analysis for a complete specious plant group, the highly threatened Cactaceae. Using geographic distribution data of 1438 cactus species, we assessed how well the current PA network represents them. We also systematically identified priority areas for conservation of cactus species that met and failed to meet conservation targets accounting for their conservation status. There were 261 species with no coverage by PAs (gap species). A greater percentage of cacti species (18%) lacked protection than mammals (9.7%) and birds (5.6%), and also a greater percentage of threatened cacti species (32%) were outside protected areas than amphibians (26.5%), birds (19.9%), or mammals (16%). The top 17% of the landscape that best captured covered species represented on average 52.9% of species ranges. The priority areas for gap species and the unprotected portion of the ranges of species that only partially met their conservation target (i.e., partial gap) captured on average 75.2% of their ranges, of which 100 were threatened gap species. These findings and knowledge of the threats affecting species provide information that can be used to improve planning for cacti conservation and highlight the importance of assessing the representation of major groups, such as plants, in PAs to determining the performance of the current PA network.     

Using abiotic variables to predict importance of sites for species representation

In systematic conservation planning, species distribution data for all sites in a planning area are used to prioritize each site in terms of the site's importance toward meeting the goal of species representation. But comprehensive species data are not available in most planning areas and would be expensive to acquire. As a shortcut, ecologists use surrogates, such as occurrences of birds or another well‐surveyed taxon, or land types defined from remotely sensed data, in the hope that sites that represent the surrogates also represent biodiversity. Unfortunately, surrogates have not performed reliably. We propose a new type of surrogate, predicted importance, that can be developed from species data for a q% subset of sites. With species data from this subset of sites, importance can be modeled as a function of abiotic variables available at no charge for all terrestrial areas on Earth. Predicted importance can then be used as a surrogate to prioritize all sites. We tested this surrogate with 8 sets of species data. For each data set, we used a q% subset of sites to model importance as a function of abiotic variables, used the resulting function to predict importance for all sites, and evaluated the number of species in the sites with highest predicted importance. Sites with the highest predicted importance represented species efficiently for all data sets when q = 25% and for 7 of 8 data sets when q = 20%. Predicted importance requires less survey effort than direct selection for species representation and meets representation goals well compared with other surrogates currently in use. This less expensive surrogate may be useful in those areas of the world that need it most, namely tropical regions with the highest biodiversity, greatest biodiversity loss, most severe lack of inventory data, and poorly developed protected area networks.     

Biogeographical differences between the two Capra pyrenaica subspecies, C. p. victoriae and C. p. hispanica, inhabiting the Iberian Peninsula: Implications for conservation

The Iberian wild goat (Capra pyrenaica), an endemic species to the Iberian Peninsula, was represented in the past by four subspecies. Currently, the conservation status of the recognized subspecies - C. p. hispanica and C. p. victoriae - is quite different, being the former broadly distributed while the latter inhabits fewer and smaller areas. In this global context, our specific aims were to assess the biogeographical relationships, similarities and differences between the Iberian wild goat subspecies, and to evaluate their conservation status from a biogeographical perspective. We modelled the ecogeographical favourability for each subspecies, and also for the whole species, and we comparatively analyzed the spatial relationships among models in order to detect differential biogeographical patterns. Our results showed a differential pattern for each Iberian wild goat subspecies, and thus each one should be independently considered when designing management strategies. The distribution of C. p. victoriae was closely related to climatic conditions, thus it is highly sensitive to variations in climatic characteristics. Finally, from the perspective of metapopulation theory, and according to the geographical structure of their favourable areas, C. p. victoriae is at higher risk than C. p. hispanica. We discuss the implications of our results for species/subspecies conservation, and highlight the importance of conservation biogeography.     

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.     

Conservation Development Practices,Extent, and Land‐Use Effects in the United States

Abstract: Conservation development projects combine real‐estate development with conservation of land and other natural resources. Thousands of such projects have been conducted in the United States and other countries through the involvement of private developers, landowners, land trusts, and government agencies. Previous research has demonstrated the potential value of conservation development for conserving species, ecological functions, and other resource values on private lands, especially when traditional sources of conservation funding are not available. Nevertheless, the aggregate extent and effects of conservation development were previously unknown. To address this gap, we estimated the extent and trends of conservation development in the United States and characterized its key attributes to understand its aggregate contribution to land‐conservation and growth‐management objectives. We interviewed representatives from land trusts, planning agencies, and development companies, searched the Internet for conservation development projects and programs, and compiled existing databases of conservation development projects. We collected data on 3884 projects encompassing 1.38 million ha. About 43% of the projects targeted the conservation of specific plant or animal species or ecological communities of conservation concern; 84% targeted the protection of native ecosystems representative of the project area; and 42% provided buffers to existing protected areas. The percentage of protected land in conservation development projects ranged from <40% to >99%, and the effects of these projects on natural resources differed widely. We estimate that conservation development projects have protected roughly 4 million ha of land in the United States and account for about 25% of private‐land conservation activity nationwide.     

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 Errors and Gaps in Spatial Data Sets on Assessment of Conservation Progress

Data on the location and extent of protected areas, ecosystems, and species’ distributions are essential for determining gaps in biodiversity protection and identifying future conservation priorities. However, these data sets always come with errors in the maps and associated metadata. Errors are often overlooked in conservation studies, despite their potential negative effects on the reported extent of protection of species and ecosystems. We used 3 case studies to illustrate the implications of 3 sources of errors in reporting progress toward conservation objectives: protected areas with unknown boundaries that are replaced by buffered centroids, propagation of multiple errors in spatial data, and incomplete protected‐area data sets. As of 2010, the frequency of protected areas with unknown boundaries in the World Database on Protected Areas (WDPA) caused the estimated extent of protection of 37.1% of the terrestrial Neotropical mammals to be overestimated by an average 402.8% and of 62.6% of species to be underestimated by an average 10.9%. Estimated level of protection of the world's coral reefs was 25% higher when using recent finer‐resolution data on coral reefs as opposed to globally available coarse‐resolution data. Accounting for additional data sets not yet incorporated into WDPA contributed up to 6.7% of additional protection to marine ecosystems in the Philippines. We suggest ways for data providers to reduce the errors in spatial and ancillary data and ways for data users to mitigate the effects of these errors on biodiversity assessments. Efectos de Errores y Vacíos en Conjuntos de Datos Espaciales sobre la Evaluación del Progreso de la Conservación     

Priority areas for conservation of and research focused on terrestrial vertebrates

Effective conservation policies require comprehensive knowledge of biodiversity. However, knowledge shortfalls still remain, hindering possibilities to improve decision making and built such policies. During the last 2 decades, conservationists have made great efforts to allocate resources as efficiently as possible but have rarely considered the idea that if research investments are also strategically allocated, it would likely fill knowledge gaps while simultaneously improving conservation actions. Therefore, prioritizing areas where both conservation and research actions could be conducted becomes a critical endeavor that can further maximize return on investment. We used Zonation, a conservation planning tool and geographical distributions of amphibians, birds, mammals, and reptiles to suggest and compare priority areas for conservation and research of terrestrial vertebrates worldwide. We also evaluated the degree of human disturbance in both types of priority areas by describing the value of the human footprint index within such areas. The spatial concordance between priority conservation and research areas was low: 0.36% of the world's land area. In these areas, we found it would be possible to protect almost half of the currently threatened species and to gather information on nearly 42% of data-deficient (DD) species. We also found that 6199 protected areas worldwide are located in such places, although only 35% of them have strict conservation purposes. Areas of consensus between conservation and research areas represent an opportunity for simultaneously conserving and acquiring knowledge of threatened and DD species of vertebrates. Although the picture is not the most encouraging, joint conservation and research efforts are possible and should be fostered to save vertebrate species from our own ignorance and extinction.     

Effectiveness of protected areas for vertebrates based on taxonomic and phylogenetic diversity

Establishing protected areas is the primary goal and tool for preventing irreversible biodiversity loss. However, the effectiveness of protected areas that target specific species has been questioned for some time because targeting key species for conservation may impair the integral regional pool of species diversity and phylogenetic and functional diversity are seldom considered. We assessed the efficacy of protected areas in China for the conservation of phylogenetic diversity based on the ranges and phylogenies of 2279 terrestrial vertebrates. Phylogenetic and taxonomic diversity were strongly and positively correlated, and only 12.1–43.8% of priority conservation areas are currently protected. However, the patterns and coverage of phylogenetic diversity were affected when weighted by species richness. These results indicated that in China, protected areas targeting high species richness protected phylogenetic diversity well overall but failed to do so in some regions with more unique or threatened communities (e.g., coastal areas of eastern China, where severely threatened avian communities were less protected). Our results suggest that the current distribution of protected areas could be improved, although most protected areas protect both taxonomic and phylogenetic diversity.     

Protected area planning to conserve biodiversity in an uncertain future

Protected areas are a key instrument for conservation. Despite this, they are vulnerable to risks associated with weak governance, land-use intensification, and climate change. We used a novel hierarchical optimization approach to identify priority areas for expanding the global protected area system that explicitly accounted for such risks while maximizing protection of all known terrestrial vertebrate species. To incorporate risk categories, we built on the minimum set problem, where the objective is to reach species distribution protection targets while accounting for 1 constraint, such as land cost or area. We expanded this approach to include multiple objectives accounting for risk in the problem formulation by treating each risk layer as a separate objective in the problem formulation. Reducing exposure to these risks required expanding the area of the global protected area system by 1.6% while still meeting conservation targets. Incorporating risks from weak governance drove the greatest changes in spatial priorities for protection, and incorporating risks from climate change required the largest increase (2.52%) in global protected area. Conserving wide-ranging species required countries with relatively strong governance to protect more land when they bordered nations with comparatively weak governance. Our results underscore the need for cross-jurisdictional coordination and demonstrate how risk can be efficiently incorporated into conservation planning. Planeación de las áreas protegidas para conservar la biodiversidad en un futuro incierto     

Conservation planning on China's borders with Myanmar,Laos, and Vietnam

Transboundary conservation is playing an increasingly important role in maintaining ecosystem integrity and halting biodiversity loss caused by anthropogenic activities. However, lack of information on species distributions in transboundary regions and understanding of the threats in these areas impairs conservation. We developed a spatial conservation plan for the transboundary areas between Yunnan province, southwestern China, and neighboring Myanmar, Laos, and Vietnam in the Indo-Burma biodiversity hotspot. To identify priority areas for conservation and restoration, we determined species distribution patterns and recent land-use changes and examined the spatiotemporal dynamics of the connected natural forest, which supports most species. We assessed connectivity with equivalent connected area (ECA), which is the amount of reachable habitat for a species. An ECA incorporates the presence of habitat in a patch and the amount of habitat in other patches within dispersal distance. We analyzed 197,845 locality records from specimen collections and monographs for 21,004 plant and vertebrate species. The region of Yunnan immediately adjacent to the international borders had the highest species richness, with 61% of recorded species and 56% of threatened vertebrates, which suggests high conservation value. Satellite imagery showed the area of natural forest in the border zone declined by 5.2% (13,255 km²) from 1995 to 2018 and monoculture plantations increased 92.4%, shrubland 10.1%, and other cropland 6.2%. The resulting decline in connected natural forest reduced the amount of habitat, especially for forest specialists with limited dispersal abilities. The most severe decline in connectivity was along the Sino-Vietnamese border. Many priority areas straddle international boundaries, indicating demand and potential for establishing transboundary protected areas. Our results illustrate the importance of bi- and multilateral cooperation to protect biodiversity in this region and provide guidance for future conservation planning and practice.     

Conservation Prioritization Using GAP Data

Data collected by the Gap Analysis Program in the state of Idaho (U.S.A.) are used to prioritize the selection of locations for conservation action and research. Set coverage and integer programming algorithms provide a sequence of localities that maximize the number of species or vegetation classes represented at each step. Richness maps of vegetation cover class diversity, terrestrial vertebrate species diversity ("hot spot analysis"), endangered, threatened, and candidate species diversity, and unprotected vertebrate species diversity ("gap analysis"), when prioritized, show a rapid accumulation of species as more localities are chosen for terrestrial vertebrates and unprotected vertebrates. Gap analysis identifies four target areas ("gaps") that include 79 of the 83 vertebrate species not currently protected. Accumulation of vegetation cover classes and endangered, threatened, and candidate species is much slower. Sweep analysis is used to determine how well prioritizing on one component of diversity accumulates other components. Endangered, threatened, and candidate species do not sweep total vertebrates as well as unprotected vertebrates do, but are better than vegetation classes. Total vertebrates sweep endangered, threatened, and candidate species better than unprotected vertebrates do, which in turn are better than vegetation classes. We emphasize that prioritization must be part of conservation efforts at multiple scales and that prioritization points out important localities where more detailed work mast be undertaken.     

Alarming decline of freshwater trigger species in western Mediterranean key biodiversity areas

Theidentification of key biodiversity areas (KBA) was initiated by the International Union for Conservation of Nature in 2004 to overcome taxonomic biases in the selection of important areas for conservation, including freshwater ecosystems. Since then, several KBAs have been identified mainly based on the presence of trigger species (i.e., species that trigger either the vulnerability and or the irreplaceability criterion and thus identify a site as a KBA). However, to our knowledge, many of these KBAs have not been validated. Therefore, classical surveys of the taxa used to identify freshwater KBAs (fishes, molluscs, odonates, and aquatic plants) were conducted in Douro (Iberian Peninsula) and Sebou (Morocco) River basins in the Mediterranean Biodiversity Hotspot. Environmental DNA analyses were undertaken in the Moroccan KBAs. There was a mismatch between the supposed and actual presence of trigger species. None of the trigger species were found in 43% and 50% of all KBAs surveyed in the Douro and Sebou basins, respectively. Shortcomings of freshwater KBA identification relate to flawed or lack of distribution data for trigger species. This situation results from a misleading initial identification of KBAs based on poor (or even inaccurate) ecological information or due to increased human disturbance between initial KBA identification and the present. To improve identification of future freshwater KBAs, we suggest selecting trigger species with a more conservative approach; use of local expert knowledge and digital data (to assess habitat quality, species distribution, and potential threats); consideration of the subcatchment when delineating KBAs boundaries; thoughtful consideration of terrestrial special areas for conservation limits; and periodic field validation.