Dracula's ménagerie: A multispecies occupancy analysis of lynx, wildcat, and wolf in the Romanian Carpathians.

The recovery of terrestrial carnivores in Europe is a conservation success story. Initiatives focused on restoring top predators require information on how resident species may interact with the re-introduced species as their interactions have the potential to alter food webs, yet such data are scarce for Europe.In this study, we assessed patterns of occupancy and interactions between three carnivore species in the Romanian Carpathians. Romania houses one of the few intact carnivore guilds in Europe, making it an ideal system to assess intraguild interactions and serve as a guide for reintroductions elsewhere.We used camera trap data from two seasons in Transylvanian forests to assess occupancy and co-occurrence of carnivores using multispecies occupancy models.Mean occupancy in the study area was highest for lynx (Ψwinter = 0.76 95% CI: 0.42-0.92; Ψautumn = 0.71 CI: 0.38-0.84) and wolf (Ψwinter = 0.60 CI: 0.34-0.78; Ψautumn = 0.81 CI: 0.25-0.95) and lowest for wildcat (Ψwinter = 0.40 CI: 0.19-0.63; Ψautumn = 0.52 CI: 0.17-0.78)We found that marginal occupancy predictors for carnivores varied between seasons. We also found differences in predictors of co-occurrence between seasons for both lynx-wolf and wildcat-wolf co-occurrence. For both seasons, we found that conditional occupancy probabilities of all three species were higher when another species was present.Our results indicate that while there are seasonal differences in predictors of occupancy and co-occurrence of the three species, co-occurrence in our study area is high.Terrestrial carnivore recovery efforts are ongoing worldwide. Insights into interspecific relations between carnivore species are critical when considering the depauperate communities they are introduced in. Our work showcases that apex carnivore coexistence is possible, but dependent on protection afforded to forest habitats and their prey base.

Measuring protected-area effectiveness using vertebrate distributions from leech iDNA.

Protected areas are key to meeting biodiversity conservation goals, but direct measures of effectiveness have proven difficult to obtain. We address this challenge by using environmental DNA from leech-ingested bloodmeals to estimate spatially-resolved vertebrate occupancies across the 677 km2 Ailaoshan reserve in Yunnan, China. From 30,468 leeches collected by 163 park rangers across 172 patrol areas, we identify 86 vertebrate species, including amphibians, mammals, birds and squamates. Multi-species occupancy modelling shows that species richness increases with elevation and distance to reserve edge. Most large mammals (e.g. sambar, black bear, serow, tufted deer) follow this pattern; the exceptions are the three domestic mammal species (cows, sheep, goats) and muntjak deer, which are more common at lower elevations. Vertebrate occupancies are a direct measure of conservation outcomes that can help guide protected-area management and improve the contributions that protected areas make towards global biodiversity goals. Here, we show the feasibility of using invertebrate-derived DNA to estimate spatially-resolved vertebrate occupancies across entire protected areas.

Limited reciprocal surrogacy of bird and habitat diversity and inconsistencies in their representation in Romanian protected areas.

Because it is impossible to comprehensively characterize biodiversity at all levels of organization, conservation prioritization efforts need to rely on surrogates. As species distribution maps of relished groups as well as high-resolution remotely sensed data increasingly become available, both types of surrogates are commonly used. A good surrogate should represent as much of biodiversity as possible, but it often remains unclear to what extent this is the case. Here, we aimed to address this question by assessing how well bird species and habitat diversity represent one another. We conducted our study in Romania, a species-rich country with high landscape heterogeneity where bird species distribution data have only recently started to become available. First, we prioritized areas for conservation based on either 137 breeding bird species or 36 habitat classes, and then evaluated their reciprocal surrogacy performance. Second, we examined how well these features are represented in already existing protected areas. Finally, we identified target regions of high conservation value for the potential expansion of the current network of reserves (as planned under the new EU Biodiversity Strategy for 2030). We found a limited reciprocal surrogacy performance, with bird species performing slightly better as a conservation surrogate for habitat diversity than vice versa. We could also show that areas with a high conservation value based on habitat diversity were represented better in already existing protected areas than areas based on bird species, which varied considerably between species. Our results highlight that taxonomic and environmental (i.e., habitat types) data may perform rather poorly as reciprocal surrogates, and multiple sources of data are required for a full evaluation of protected areas expansion.

Habitat connectivity and resource selection in an expanding bobcat (Lynx rufus) population.

Terrestrial carnivores are among the most imperiled species worldwide, yet some species are resilient and are recovering in human-dominated landscapes after decades or centuries of absence. Bobcat (Lynx rufus) populations were extirpated from much of Midwestern US in the mid-1800's, and are currently expanding and recolonizing their former range. In this study, we investigated multi-scale habitat selection for Ohio's expanding bobcat population, and examined habitat connectivity in order to evaluate the conduits for dispersal statewide. We used citizen observations collected between 1978 and 2019 and logistic regression to evaluate population-level habitat selection, and GPS telemetry data for 20 individuals collected between 2012 and 2014 and a distribution-weighted exponential Resource Selection Function to evaluate individual-level habitat selection within home ranges. At the population level, bobcats selected for higher amounts of forest and pasture (at a 50 km2 scale) and herbaceous vegetation (at 15-50 50 km2 scales), thus overall heterogeneous forested habitat. At individual (home range) level, bobcats selected for forested habitats with low road density and farther away from high traffic roads; they also showed weak selection for open habitat at the home range level. Male home ranges were significantly greater than female home ranges. Lastly, we used the population-level spatial outputs (i.e. habitat suitability map) to parameterize habitat connectivity models using circuit theory in the program Circuitscape. We tested three relationships between habitat suitability and resistance to movement and used a subset of data on potential dispersing individuals to evaluate which relationship performed best. All three relationships performed almost equally well, and we calculated a weighted averaged connectivity map as our final map. Habitat was highly permeable to movements between core areas of two genetically distinct subpopulations located in southeastern Ohio. We also identified potential dispersal corridors from the core areas to other regions of Ohio dominated by agriculture and suburban development via forested riparian corridors. Overall, our analysis offers new information on habitat selection and connectivity in a rebounding felid population and offers important ecological information for wildlife management strategies. We recommend that the suitability and connectivity models should be periodically updated until the population reaches an equilibrium, and be integrated with data from neighboring states for a comprehensive assessment of a conservation success story.

Sublethal, Behavioral, and Developmental Effects of the Neonicotinoid Pesticide Imidacloprid on Larval Wood Frogs (Rana sylvatica).

Imidacloprid, a neonicotinoid pesticide, is used to prevent the spread of the hemlock woolly adelgid, currently affecting Eastern Hemlock trees across North America. When the pesticide is sprayed directly onto soil around infested trees (soil drenching), it can run off into aquatic systems, with potential negative effects on biota. Simultaneously, climate change may lead to faster pool drying, which acts as an additional stressor for sensitive species such as amphibians. We evaluated the sublethal effects of imidacloprid (10 ppb), and interaction with shorter hydroperiods on the larval behavior, growth, and survival of a model organism, the wood frog (Rana sylvatica). We performed 3 behavioral experiments evaluating swimming speed, time spent swimming, and distance the larvae swam. We found that larvae raised in 10 ppb imidacloprid or shorter hydroperiod did not differ in their swimming time, distance, and speed from nonexposed larvae. Naïve larvae exposed for 20 min to 10- to 500-ppb concentrations also showed similar performance to nonexposed larvae. However, when we applied a stimulus halfway through each experiment, we found that larvae exposed to 10 ppb imidacloprid (short and long term) swam shorter distances and spent less time swimming, suggesting that imidacloprid exposure may slow reaction time, potentially increasing the risk of predation. To minimize impacts on pool-breeding amphibians, imidacloprid application to combat the invasive hemlock woolly adelgid should use trunk injection and avoid soil drenching. Environ Toxicol Chem 2021;40:1840-1849. © 2021 SETAC.

Complex hydroperiod induced carryover responses for survival, growth, and endurance of a pond-breeding amphibian.

Assessing carryover effects from the aquatic to the terrestrial stage of pond-breeding amphibians is critical as temperature and hydrologic regimes of temporary ponds continue to be altered as a result of climate change and other stressors. We evaluated carryover effects of hydroperiod length (50-62 days) on amphibian survival, developmental rates, and locomotor performance using a model organism, the wood frog (Rana sylvatica), through aquatic and terrestrial mesocosm experiments with individual tests of locomotor performance. We found that shorter hydroperiods (50 days) had low larval survival (0.44 ± 0.03) compared to the 62-day hydroperiod (0.91 ± 0.09) and increased developmental rates, resulting in smaller sizes at metamorphosis. We did not find evidence of carryover effects on terrestrial survival three months post-metamorphosis with all hydroperiod treatments showing high terrestrial survival (0.88 ± 0.07). However, post-metamorphic frogs from the longer hydroperiod treatments grew faster and larger compared to individuals from shortest hydroperiods and performed significantly better during endurance trials at 18 °C. Disentangling complex carryover effects across multiple life stages in species with high phenotypic plasticity can shed light on the physiological capacity of species to respond to changing environments and inform mechanistic predictions of persistence in the face of anthropogenic stressors.

Identification of areas of very high biodiversity value to achieve the EU Biodiversity Strategy for 2030 key commitments.

BACKGROUND: The European Union strives to increase protected areas of the EU terrestrial surface to 30% by year 2030, of which one third should be strictly protected. Designation of the Natura 2000 network, the backbone of nature protection in the EU, was mostly an expert-opinion process with little systematic conservation planning. The designation of the Natura 2000 network in Romania followed the same non-systematic approach, resulting in a suboptimal representation of invertebrates and plants. To help identify areas with very high biodiversity without repeating past planning missteps, we present a reproducible example of spatial prioritization using Romania's current terrestrial Natura 2000 network and coarse-scale terrestrial species occurrence. METHODS: We used 371 terrestrial Natura 2000 Sites of Community Importance (Natura 2000 SCI), designated to protect 164 terrestrial species listed under Annex II of Habitats Directive in Romania in our spatial prioritization analyses (marine Natura 2000 sites and species were excluded). Species occurrences in terrestrial Natura 2000 sites were aggregated at a Universal Traverse Mercator spatial resolution of 1 km2. To identify priority terrestrial Natura 2000 sites for species conservation, and to explore if the Romanian Natura 2000 network sufficiently represents species included in Annex II of Habitats Directive, we used Zonation v4, a decision support software tool for spatial conservation planning. We carried out the analyses nationwide (all Natura 2000 sites) as well as separately for each biogeographic region (i.e., Alpine, Continental, Pannonian, Steppic and Black Sea). RESULTS: The results of spatial prioritization of terrestrial Natura 2000 vary greatly by planning scenario. The performance of national-level planning of top priorities is minimal. On average, when 33% of the landscape of Natura 2000 sites is protected, only 20% of the distribution of species listed in Annex II of Habitats Directive are protected. As a consequence, the representation of species by priority terrestrial Natura 2000 sites is lessened when compared to the initial set of species. When planning by taxonomic group, the top-priority areas include only 10% of invertebrate distribution in Natura 2000. When selecting top-priority areas by biogeographical region, there are significantly fewer gap species than in the national level and by taxa scenarios; thusly, the scenario outperforms the national-level prioritization. The designation of strictly protected areas as required by the EU Biodiversity Strategy for 2030 should be followed by setting clear objectives, including a good representation of species and habitats at the biogeographical region level.

Quantifying biodiversity trade-offs in the face of widespread renewable and unconventional energy development.

The challenge of balancing biodiversity protection with economic growth is epitomized by the development of renewable and unconventional energy, whose adoption is aimed at stemming the impacts of global climate change, yet has outpaced our understanding of biodiversity impacts. We evaluated the potential conflict between biodiversity protection and future electricity generation from renewable (wind farms, run-of-river hydro) and non-renewable (shale gas) sources in British Columbia (BC), Canada using three metrics: greenhouse gas (GHG) emissions, electricity cost, and overlap between future development and conservation priorities for several fish and wildlife groups - small-bodied vertebrates, large mammals, freshwater fish - and undisturbed landscapes. Sharp trade-offs in global versus regional biodiversity conservation exist for all energy technologies, and in BC they are currently smallest for wind energy: low GHG emissions, low-moderate overlap with top conservation priorities, and competitive energy cost. GHG emissions from shale gas are 1000 times higher than those from renewable sources, and run-of-river hydro has high overlap with conservation priorities for small-bodied vertebrates. When all species groups were considered simultaneously, run-of-river hydro had moderate overlap (0.56), while shale gas and onshore wind had low overlap with top conservation priorities (0.23 and 0.24, respectively). The unintended cost of distributed energy sources for regional biodiversity suggest that trade-offs based on more diverse metrics must be incorporated into energy planning.

Author Correction: Roadkill and space use data predict vehicle-strike hotspots and mortality rates in a recovering bobcat (Lynx rufus) population.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Roadkill and space use data predict vehicle-strike hotspots and mortality rates in a recovering bobcat (Lynx rufus) population.

Roadways pose challenges for conserving wide-ranging animal species. As bobcat (Lynx rufus) populations recover in Ohio, an accurate evaluation of population metrics is critical to understanding future population trajectories. In this study, we integrated multiple datasets to examine overall road mortality rates in Ohio. First, we utilized a long-term vehicle-strike dataset (1978-2017) to determine landscape and local predictors of road mortality. We found that bobcats were killed at higher rates on interstates regardless of surrounding landscape composition, but that landscape variables were useful at predicting mortality on lower-traffic roads. To explore road avoidance behaviors, we used GPS telemetry data from 18 individuals to compare road crossings along trajectory paths with random road crossings simulated using Correlated Random Walks. Bobcats exhibited avoidance of certain route types (county, municipal, and US routes). Finally, by integrating traffic volume data, road crossing behavior, and accounting for the proportion of each route type present in the study area, we estimated that a minimum of 6% and up to 18% of the bobcat population in Ohio is lost to vehicle-strikes annually. To fully understand the population level impacts of this mortality, we recommend further monitoring of age structure and sex of roadkill animals. Our results identify potential areas for mitigation of vehicle-strikes and emphasize the importance of accounting for road mortality when making management decisions for Ohio's recovering bobcat population.

Empirical analysis and modeling of Argos Doppler location errors in Romania.

BACKGROUND: Advances in wildlife tracking technology have allowed researchers to understand the spatial ecology of many terrestrial and aquatic animal species. Argos Doppler is a technology that is widely used for wildlife tracking owing to the small size and low weight of the Argos transmitters. This allows them to be fitted to small-bodied species. The longer lifespan of the Argos units in comparison to units outfitted with miniaturized global positioning system (GPS) technology has also recommended their use. In practice, large Argos location errors often occur due to communication conditions such as transmitter settings, local environment, and the behavior of the tracked individual. METHODS: Considering the geographic specificity of errors and the lack of benchmark studies in Eastern Europe, the research objectives were: (1) to evaluate the accuracy of Argos Doppler technology under various environmental conditions in Romania, (2) to investigate the effectiveness of straightforward destructive filters for improving Argos Doppler data quality, and (3) to provide guidelines for processing Argos Doppler wildlife monitoring data. The errors associated with Argos locations in four geographic locations in Romania were assessed during static, low-speed and high-speed tests. The effectiveness of the Douglas Argos distance angle filter algorithm was then evaluated to ascertain its effect on the minimization of localization errors. RESULTS: Argos locations received in the tests had larger associated horizontal errors than those indicated by the operator of the Argos system, including under ideal reception conditions. Positional errors were similar to those obtained in other studies outside of Europe. The errors were anisotropic, with larger longitudinal errors for the vast majority of the data. Errors were mostly related to speed of the Argos transmitter at the time of reception, but other factors such as topographical conditions and orientation of antenna at the time of the transmission also contributed to receiving low-quality data. The Douglas Argos filter successfully excluded the largest errors while retaining a large amount of data when the threshold was set to the local scale (two km). DISCUSSION: Filter selection requires knowledge about the movement patterns and behavior of the species of interest, and the parametrization of the selected filter typically requires a trial and error approach. Selecting the proper filter reduces the errors while retaining a large amount of data. However, the post-processed data typically includes large positional errors; thus, we recommend incorporating Argos error metrics (e.g., error ellipse) or use complex modeling approaches when working with filtered data.

Navigating protected areas networks for improving diffusion of conservation practices.

The Natura 2000 protected area network is the cornerstone of European Union's biodiversity conservation strategy. These protected areas range across multiple biogeographic regions, and they include a diversity of species assemblages along with a diversity of managing organizations, altogether making difficult to pool relevant sites to facilitate the flow of knowledge significant to their management. Here we introduce an approach to navigating protected area networks that has the potential to foster systematic identification of key sites for facilitating the exchange of knowledge and diffusion of information within the network. To demonstrate our approach, we abstractly represented Romanian Natura 2000 network as a co-occurrence network, with individual sites as nodes and shared species as edges, further combining into our analysis network topology, community detection, and network reduction methods. We identified most representative Natura 2000 sites that may increase the transfer of information within the national network of protected areas, detected clusters of sites and key sites for maintaining network cohesiveness, and highlighted the subsample of sites that retain the characteristics of the entire network. Our analysis provides implications for protected area prioritization by proposing a network perspective approach to collaboration rooted in ecological principles.

Conservation priorities for terrestrial mammals in Dobrogea Region, Romania.

Based on species occurrence records of museum collections, published literature, and unpublished records shared by mammalian experts, we compiled a distribution database for 59 terrestrial mammals populating the extensively protected Dobrogea Region of Romania. The spatial patterns of mammal distribution and diversity was evaluated and systematic conservation planning applied to identify priority areas for their conservation. The spatial analyses revealed that intensive sampling was not directly correlated to mammal diversity but rather to accessibility for inventory. The spatial prioritisation analysis indicated a relatively aggregated pattern of areas with a high or low conservation value with virtually no connecting corridors between them. The significant overlap between Natura 2000 sites and national protected areas induced an over-optimistic vision of the effectiveness and representativeness of existing Natura 2000 network for species found in Annexes II and IV of the Habitats Directive. These results represent a key step in identifying core areas for the protection of mammal diversity and dispersal corridors for improved connectivity, and to guide future conservation efforts in increasing the effectiveness of the existing protected areas in the context of environmental changes.

Power, influence and structure in Natura 2000 governance networks. A comparative analysis of two protected areas in Romania.

Successful management of complex social-ecological landscapes overlapping Natura 2000 sites requires collaboration between various actors such as law enforcement agencies, NGOs and enterprises. Natura 2000 governance is stimulated by central actors (e.g., Natura 2000 administrators), with successes and failures of management activities depending on the capacity of the network leader to implement a collaborative approach to environmental governance. By using social network analysis, we analysed the cooperation, information flow and capacity for collective action within Natura 2000 governance networks within two Romanian protected areas: Lower Siret Floodplain and Iron Gates Natural Park. The two networks represent protected areas managed by different types of organisations (i.e., Lower Siret Floodplain - by an NGO, Iron Gates Natural Park - by a public entity). Taking into consideration that NGOs may favour an adaptive co-management, while the public bodies may take a top-down management approach, we hypothesize that Lower Siret Floodplain will have a more cohesive and collaborating network compared to Iron Gates Natural Park, and that there will be a greater representation of private and NGO sector in the network coordinated by Lower Siret Floodplain. Contrary to our expectations, the results show that collaboration patterns are similar in the two networks, although they are governed by two different types of institutions, both being less participative than expected, with low involvement of NGOs and private stakeholders. Furthermore, Lower Siret Floodplain network is surprisingly more centralized around a small number of public authorities, and the pre-existing power of public bodies likely inhibit the capacity of the NGO to collaborate with private stakeholders. We also found lower collaboration levels between actors in the network periphery with other organisations from the same cluster, denoting a clear top-down approach of the management in both networks. Our findings suggest that delegating the protected areas administration to NGOs, a solution to increase the use of co-management in protected areas, does not solve the poor representation of private stakeholders.

Integrating sign surveys and telemetry data for estimating brown bear (Ursus arctos) density in the Romanian Carpathians.

Accurate population size estimates are important information for sustainable wildlife management. The Romanian Carpathians harbor the largest brown bear (Ursus arctos) population in Europe, yet current management relies on estimates of density that lack statistical oversight and ignore uncertainty deriving from track surveys. In this study, we investigate an alternative approach to estimate brown bear density using sign surveys along transects within a novel integration of occupancy models and home range methods. We performed repeated surveys along 2-km segments of forest roads during three distinct seasons: spring 2011, fall-winter 2011, and spring 2012, within three game management units and a Natura 2000 site. We estimated bears abundances along transects using the number of unique tracks observed per survey occasion via N-mixture hierarchical models, which account for imperfect detection. To obtain brown bear densities, we combined these abundances with the effective sampling area of the transects, that is, estimated as a function of the median (± bootstrapped SE) of the core home range (5.58 ± 1.08 km2) based on telemetry data from 17 bears tracked for 1-month periods overlapping our surveys windows. Our analyses yielded average brown bear densities (and 95% confidence intervals) for the three seasons of: 11.5 (7.8-15.3), 11.3 (7.4-15.2), and 12.4 (8.6-16.3) individuals/100 km2. Across game management units, mean densities ranged between 7.5 and 14.8 individuals/100 km2. Our method incorporates multiple sources of uncertainty (e.g., effective sampling area, imperfect detection) to estimate brown bear density, but the inference fundamentally relies on unmarked individuals only. While useful as a temporary approach to monitor brown bears, we urge implementing DNA capture-recapture methods regionally to inform brown bear management and recommend increasing resources for GPS collars to improve estimates of effective sampling area.

Collaboration Networks in Applied Conservation Projects across Europe.

The main funding instrument for implementing EU policies on nature conservation and supporting environmental and climate action is the LIFE Nature programme, established by the European Commission in 1992. LIFE Nature projects (>1400 awarded) are applied conservation projects in which partnerships between institutions are critical for successful conservation outcomes, yet little is known about the structure of collaborative networks within and between EU countries. The aim of our study is to understand the nature of collaboration in LIFE Nature projects using a novel application of social network theory at two levels: (1) collaboration between countries, and (2) collaboration within countries using six case studies: Western Europe (United Kingdom and Netherlands), Eastern Europe (Romania and Latvia) and Southern Europe (Greece and Portugal). Using data on 1261 projects financed between 1996 and 2013, we found that Italy was the most successful country not only in terms of awarded number of projects, but also in terms of overall influence being by far the most influent country in the European LIFE Nature network, having the highest eigenvector (0.989) and degree centrality (0.177). Another key player in the network is Netherlands, which ensures a fast communication flow with other network members (closeness-0.318) by staying connected with the most active countries. Although Western European countries have higher centrality scores than most of the Eastern European countries, our results showed that overall there is a lower tendency to create partnerships between different organization categories. Also, the comparisons of the six case studies indicates significant differences in regards to the pattern of creating partnerships, providing valuable information on collaboration on EU nature conservation. This study represents a starting point in predicting the formation of future partnerships within LIFE Nature programme, suggesting ways to improve transnational cooperation and communication.

Higher fungal diversity is correlated with lower CO2 emissions from dead wood in a natural forest.

Wood decomposition releases almost as much CO2 to the atmosphere as does fossil-fuel combustion, so the factors regulating wood decomposition can affect global carbon cycling. We used metabarcoding to estimate the fungal species diversities of naturally colonized decomposing wood in subtropical China and, for the first time, compared them to concurrent measures of CO2 emissions. Wood hosting more diverse fungal communities emitted less CO2, with Shannon diversity explaining 26 to 44% of emissions variation. Community analysis supports a 'pure diversity' effect of fungi on decomposition rates and thus suggests that interference competition is an underlying mechanism. Our findings extend the results of published experiments using low-diversity, laboratory-inoculated wood to a high-diversity, natural system. We hypothesize that high levels of saprotrophic fungal biodiversity could be providing globally important ecosystem services by maintaining dead-wood habitats and by slowing the atmospheric contribution of CO2 from the world's stock of decomposing wood. However, large-scale surveys and controlled experimental tests in natural settings will be needed to test this hypothesis.

Species, habitats, society: an evaluation of research supporting EU's Natura 2000 network.

The Natura 2000 network is regarded as one of the conservation success stories in the global effort to protect biodiversity. However, significant challenges remain in Natura 2000 implementation, owing to its rapid expansion, and lack of a coherent vision for its future. Scientific research is critical for identifying conservation priorities, setting management goals, and reconciling biodiversity protection and society in the complex political European landscape. Thus, there is an urgent need for a comprehensive evaluation of published Natura 2000 research to highlight prevalent research themes, disciplinary approaches, and spatial entities. We conducted a systematic review of 572 scientific articles and conference proceedings focused on Natura 2000 research, published between 1996 and 2014. We grouped these articles into 'ecological' and 'social and policy' categories. Using a novel application of network analysis of article keywords, we found that Natura 2000 research forms a cohesive small-world network, owing to the emphasis on ecological research (79% of studies, with a strong focus on spatial conservation planning), and the underrepresentation of studies addressing 'social and policy' issues (typically focused on environmental impact assessment, multi-level governance, agri-environment policy, and ecosystem services valuation). 'Ecological' and 'social and policy' research shared only general concepts (e.g., Natura 2000, Habitats Directive) suggesting a disconnection between these disciplines. The UK and the Mediterranean basin countries dominated Natura 2000 research, and there was a weak correlation between number of studies and proportion of national territory protected. Approximately 40% of 'social and policy' research and 26% of 'ecological' studies highlighted negative implications of Natura 2000, while 21% of studies found positive social and biodiversity effects. We emphasize the need for designing inter- and transdisciplinary research in order to promote a social-ecological understanding of Natura 2000, and advance EU conservation policies.