Location-level processes drive the establishment of alien bird populations worldwide.

Human-mediated translocation of species to areas beyond their natural distribution (which results in 'alien' populations1) is a key signature of the Anthropocene2, and is a primary global driver of biodiversity loss and environmental change3. Stemming the tide of invasions requires understanding why some species fail to establish alien populations, and others succeed. To achieve this, we need to integrate the effects of features of the introduction site, the species introduced and the specific introduction event. Determining which, if any, location-level factors affect the success of establishment has proven difficult, owing to the multiple spatial, temporal and phylogenetic axes along which environmental variation may influence population survival. Here we apply Bayesian hierarchical regression analysis to a global spatially and temporally explicit database of introduction events of alien birds4 to show that environmental conditions at the introduction location, notably climatic suitability and the presence of other groups of alien species, are the primary determinants of successful establishment. Species-level traits and the size of the founding population (propagule pressure) exert secondary, but important, effects on success. Thus, current trajectories of anthropogenic environmental change will most probably facilitate future incursions by alien species, but predicting future invasions will require the integration of multiple location-, species- and event-level characteristics.

Accelerated shifts in terrestrial life zones under rapid climate change.

Rapid climate change is impacting biodiversity, ecosystem function, and human well-being. Though the magnitude and trajectory of climate change are becoming clearer, our understanding of how these changes reshape terrestrial life zones-distinct biogeographic units characterized by biotemperature, precipitation, and aridity representing broad-scale ecosystem types-is limited. To address this gap, we used high-resolution historical climatologies and climate projections to determine the global distribution of historical (1901-1920), contemporary (1979-2013), and future (2061-2080) life zones. Comparing the historical and contemporary distributions shows that changes from one life zone to another during the 20th century impacted 27 million km2 (18.3% of land), with consequences for social and ecological systems. Such changes took place in all biomes, most notably in Boreal Forests, Temperate Coniferous Forests, and Tropical Coniferous Forests. Comparing the contemporary and future life zone distributions shows the pace of life zone changes accelerating rapidly in the 21st century. By 2070, such changes would impact an additional 62 million km2 (42.6% of land) under "business-as-usual" (RCP8.5) emissions scenarios. Accelerated rates of change are observed in hundreds of ecoregions across all biomes except Tropical Coniferous Forests. While only 30 ecoregions (3.5%) had over half of their areas change to a different life zone during the 20th century, by 2070 this number is projected to climb to 111 ecoregions (13.1%) under RCP4.5 and 281 ecoregions (33.2%) under RCP8.5. We identified weak correlations between life zone change and threatened vertebrate richness, levels of vertebrate endemism, cropland extent, and human population densities within ecoregions, illustrating the ubiquitous risks of life zone changes to diverse social-ecological systems. The accelerated pace of life zone changes will increasingly challenge adaptive conservation and sustainable development strategies that incorrectly assume current ecological patterns and livelihood provisioning systems will persist.

Multinational coordination required for conservation of over 90% of marine species.

Marine species are declining at an unprecedented rate, catalyzing many nations to adopt conservation and management targets within their jurisdictions. However, marine species and the biophysical processes that sustain them are naive to international borders. An understanding of the prevalence of cross-border species distributions is important for informing high-level conservation strategies, such as bilateral or regional agreements. Here, we examined 28,252 distribution maps to determine the number and locations of transboundary marine plants and animals. More than 90% of species have ranges spanning at least two jurisdictions, with 58% covering more than 10 jurisdictions. All jurisdictions have at least one transboundary species, with the highest concentrations of transboundary species in the USA, Australia, Indonesia, and the Areas Beyond National Jurisdiction. Distributions of mapped biodiversity indicate that overcoming the challenges of multinational governance is critical for a much wider suite of species than migratory megavertebrates and commercially exploited fish stocks-the groups that have received the vast majority of multinational management attention. To effectively protect marine biodiversity, international governance mechanisms (particularly those related to the Convention on Biological Diversity, the Convention on Migratory Species, and Regional Seas Organizations) must be expanded to promote multinational conservation planning, and complimented by a holistic governance framework for biodiversity beyond national jurisdiction.

The Global Distribution and Drivers of Alien Bird Species Richness.

Alien species are a major component of human-induced environmental change. Variation in the numbers of alien species found in different areas is likely to depend on a combination of anthropogenic and environmental factors, with anthropogenic factors affecting the number of species introduced to new locations, and when, and environmental factors influencing how many species are able to persist there. However, global spatial and temporal variation in the drivers of alien introduction and species richness remain poorly understood. Here, we analyse an extensive new database of alien birds to explore what determines the global distribution of alien species richness for an entire taxonomic class. We demonstrate that the locations of origin and introduction of alien birds, and their identities, were initially driven largely by European (mainly British) colonialism. However, recent introductions are a wider phenomenon, involving more species and countries, and driven in part by increasing economic activity. We find that, globally, alien bird species richness is currently highest at midlatitudes and is strongly determined by anthropogenic effects, most notably the number of species introduced (i.e., "colonisation pressure"). Nevertheless, environmental drivers are also important, with native and alien species richness being strongly and consistently positively associated. Our results demonstrate that colonisation pressure is key to understanding alien species richness, show that areas of high native species richness are not resistant to colonisation by alien species at the global scale, and emphasise the likely ongoing threats to global environments from introductions of species.

Global assessment of marine biodiversity potentially threatened by offshore hydrocarbon activities.

Increasing global energy demands have led to the ongoing intensification of hydrocarbon extraction from marine areas. Hydrocarbon extractive activities pose threats to native marine biodiversity, such as noise, light, and chemical pollution, physical changes to the sea floor, invasive species, and greenhouse gas emissions. Here, we assessed at a global scale the spatial overlap between offshore hydrocarbon activities and marine biodiversity (>25,000 species, nine major ecosystems, and marine protected areas), and quantify the changes over time. We discovered that two-thirds of global offshore hydrocarbon activities occur in areas within the top 10% for species richness, range rarity, and proportional range rarity values globally. Thus, while hydrocarbon activities are undertaken in less than one percent of the ocean's area, they overlap with approximately 85% of all assessed species. Of conservation concern, 4% of species with the largest proportion of their range overlapping hydrocarbon activities are range restricted, potentially increasing their vulnerability to localized threats such as oil spills. While hydrocarbon activities have extended to greater depths since the mid-1990s, we found that the largest overlap is with coastal ecosystems, particularly estuaries, saltmarshes and mangroves. Furthermore, in most countries where offshore hydrocarbon exploration licensing blocks have been delineated, they do not overlap with marine protected areas (MPAs). Although this is positive in principle, many countries have far more licensing block areas than protected areas, and in some instances, MPA coverage is minimal. These findings suggest the need for marine spatial prioritization to help limit future spatial overlap between marine conservation priorities and hydrocarbon activities. Such prioritization can be informed by the spatial and quantitative baseline information provided here. In increasingly shared seascapes, prioritizing management actions that set both conservation and development targets could help minimize further declines of biodiversity and environmental changes at a global scale.

Nest-site competition between invasive and native cavity nesting birds and its implication for conservation.

Nesting cavities are often a limited resource that multiple species use. There is an ongoing discussion on whether invasive cavity nesting birds restrict the availability of this key limited resource. While the answer to this question has important conservation implications, little experimental work has been done to examine it. Here, we aimed to experimentally test whether alien cavity nesting birds affect the occupancy of cavities and the resulting breeding success of native cavity breeders in a large urban park located in Tel Aviv, Israel. Over three breeding seasons, we manipulated the entry size of nest boxes and compared the occupancy and breeding success of birds in nest boxes of two treatments. These included nest boxes with large-entrance and small-entrance holes. The large-entrance holes allowed access for both the native and invasive birds (the two main aliens in the park are the common mynas and rose-ringed parakeets). The smaller-entrance boxes, on the other hand, allowed only the smaller sized native cavity breeders (great tits and house sparrows) to enter the boxes but prevented the alien species from entering. We found that the large-entrance nest boxes were occupied by five different bird species, comprising three natives (great tit, house sparrow, Scops owl) and two invasive species (common myna, rose-ringed parakeet) while the small-entrance boxes were only occupied by the two native species. The alien common mynas and rose-ringed parakeets occupied 77.5% of the large-entrance nest boxes whereas native species, mainly great tits, occupied less than 9% of the large-entrance boxes and 36.5% of the small-entrance boxes. When examining the occupancy of those cavities that were not occupied by the aliens, natives occupied both the small and large-entrance nest boxes equally. Three quarters (78%) of the great tits breeding in the large-entrance boxes were usurped by common mynas during the breeding season and as a result breeding success was significantly lower for great tits breeding in the large-entrance boxes compared with the small-entrance boxes. The results of this study suggests that the invasive alien species can reduce the breeding potential of native cavity breeders both by exploiting the limited breeding resource (nest cavities) and by directly usurping cavities already occupied by the native species. Since the majority of large-entrance nest boxes were occupied by the larger alien birds, less native species bred in the limited number of unoccupied large-entrance nest boxes because of exploitation competition. We propose that for management purposes, nest-box programs that alter the entrance size of available natural cavities may be a practical approach, reducing the competition between native cavity breeders and alien invasive birds, and especially benefiting the smaller native cavity breeders.

Where have all the people gone? Enhancing global conservation using night lights and social media.

Conservation prioritization at large scales is complex, combining biological, environmental, and social factors. While conservation scientists now more often aim to incorporate human-related factors, a critical yet unquantified challenge remains: to identify which areas people use for recreation outside urban centers. To address this gap in applied ecology and conservation, we developed a novel approach for quantifying human presence beyond populated areas by combining social media "big data" and remote sensing tools. We used data from the Flickr photo-sharing website as a surrogate for identifying spatial variation in visitation globally, and complemented this estimate with spatially explicit information on stable night lights between 2004 and 2012, used as a proxy for identifying urban and industrial centers. Natural and seminatural areas attracting visitors were defined as areas both highly photographed and non-lit. The number of Flickr photographers within protected areas was found to be a reliable surrogate for estimating visitor numbers as confirmed by local authority censuses (r = 0.8). Half of all visitors' photos taken in protected areas originated from under 1% of all protected areas on Earth (250 of -27 000). The most photographed protected areas globally included Yosemite and Yellowstone National Parks (USA), and the Lake and Peak Districts (UK). Factors explaining the spatial variation in protected areas Flickr photo coverage included their type (e.g., UNESCO World Heritage sites have higher visitation) and accessibility to roads and trails. Using this approach, we identified photography hotspots, which draw many visitors and are also unlit (i.e., are located outside urban centers), but currently remain largely unprotected, such as Brazil's Pantanal and Bolivia's Salar de Uyuni. The integrated big data approach developed here demonstrates the benefits of combining remote sensing sources and novel geo-tagged and crowd-sourced information from social media in future efforts to identify spatial conservation gaps and pressures in real time, and their spatial and temporal variation globally.

Sensitivity analysis of conservation targets in systematic conservation planning.

Systematic conservation planning has rapidly advanced in the past decade and has been increasingly incorporated in multiple studies and conservation projects. One of its requirements is a quantitative definition of conservation targets. While the Convention on Biological Diversity aims to expand the world's protected area network to 17% of the land surface, in many cases such uniform policy-driven targets may not be appropriate for achieving persistence of various species. Targets are often set arbitrarily, often because information required for the persistence of each species is unavailable or unknown in the focal region. Conservation planners therefore need to establish complementary novel approaches to address the gaps in setting targets. Here, we develop and present a novel method that aims to help guide the selection of conservation targets, providing support for decision makers, planners, and managers. This is achieved by examining the overall flexibility of the conservation network resulting from conservation prioritization, and aiming for greater flexibility. To test this approach we applied the decision support tool Marxan to determine marine conservation priority areas in the eastern Mediterranean Sea as a case study. We assessed the flexibility of the conservation network by comparing 80 different scenarios in which conservation targets were gradually increased and assessed by a range of calculated metrics (e.g., the percentage of the total area selected, the overall connectivity). We discovered that when conservation targets were set too low (i.e., below 10% of the distribution range of each species), very few areas were identified as irreplaceable and the conservation network was not well defined. Interestingly, when conservation targets were set too high (over 50% of the species' range), too many conservation priority areas were selected as irreplaceable, an outcome which is realistically infeasible to implement. As a general guideline, we found that flexibility in a conservation network is adequate when ~10-20% of the study area is considered irreplaceable (selection frequency values over 90%). This approach offers a useful sensitivity analysis when applying target-based systematic conservation planning tools, ensuring that the resulting protected area conservation network offers more choices for managers and decision makers.

Emerging conservation challenges and prospects in an era of offshore hydrocarbon exploration and exploitation.

Globally, extensive marine areas important for biodiversity conservation and ecosystem functioning are undergoing exploration and extraction of oil and natural gas resources. Such operations are expanding to previously inaccessible deep waters and other frontier regions, while conservation-related legislation and planning is often lacking. Conservation challenges arising from offshore hydrocarbon development are wide-ranging. These challenges include threats to ecosystems and marine species from oil spills, negative impacts on native biodiversity from invasive species colonizing drilling infrastructure, and increased political conflicts that can delay conservation actions. With mounting offshore operations, conservationists need to urgently consider some possible opportunities that could be leveraged for conservation. Leveraging options, as part of multi-billion dollar marine hydrocarbon operations, include the use of facilities and costly equipment of the deep and ultra-deep hydrocarbon industry for deep-sea conservation research and monitoring and establishing new conservation research, practice, and monitoring funds and environmental offsetting schemes. The conservation community, including conservation scientists, should become more involved in the earliest planning and exploration phases and remain involved throughout the operations so as to influence decision making and promote continuous monitoring of biodiversity and ecosystems. A prompt response by conservation professionals to offshore oil and gas developments can mitigate impacts of future decisions and actions of the industry and governments. New environmental decision support tools can be used to explicitly incorporate the impacts of hydrocarbon operations on biodiversity into marine spatial and conservation plans and thus allow for optimum trade-offs among multiple objectives, costs, and risks.

Large-scale conservation planning in a multinational marine environment: cost matters.

Explicitly including cost in marine conservation planning is essential for achieving feasible and efficient conservation outcomes. Yet, spatial priorities for marine conservation are still often based solely on biodiversity hotspots, species richness, and/or cumulative threat maps. This study aims to provide an approach for including cost when planning large-scale Marine Protected Area (MPA) networks that span multiple countries. Here, we explore the incorporation of cost in the complex setting of the Mediterranean Sea. In order to include cost in conservation prioritization, we developed surrogates that account for revenue from multiple marine sectors: commercial fishing, noncommercial fishing, and aquaculture. Such revenue can translate into an opportunity cost for the implementation of an MPA network. Using the software Marxan, we set conservation targets to protect 10% of the distribution of 77 threatened marine species in the Mediterranean Sea. We compared nine scenarios of opportunity cost by calculating the area and cost required to meet our targets. We further compared our spatial priorities with those that are considered consensus areas by several proposed prioritization schemes in the Mediterranean Sea, none of which explicitly considers cost. We found that for less than 10% of the Sea's area, our conservation targets can be achieved while incurring opportunity costs of less than 1%. In marine systems, we reveal that area is a poor cost surrogate and that the most effective surrogates are those that account for multiple sectors or stakeholders. Furthermore, our results indicate that including cost can greatly influence the selection of spatial priorities for marine conservation of threatened species. Although there are known limitations in multinational large-scale planning, attempting to devise more systematic and rigorous planning methods is especially critical given that collaborative conservation action is on the rise and global financial crisis restricts conservation investments.

Disentangling the role of environmental and human pressures on biological invasions across Europe.

The accelerating rates of international trade, travel, and transport in the latter half of the twentieth century have led to the progressive mixing of biota from across the world and the number of species introduced to new regions continues to increase. The importance of biogeographic, climatic, economic, and demographic factors as drivers of this trend is increasingly being realized but as yet there is no consensus regarding their relative importance. Whereas little may be done to mitigate the effects of geography and climate on invasions, a wider range of options may exist to moderate the impacts of economic and demographic drivers. Here we use the most recent data available from Europe to partition between macroecological, economic, and demographic variables the variation in alien species richness of bryophytes, fungi, vascular plants, terrestrial insects, aquatic invertebrates, fish, amphibians, reptiles, birds, and mammals. Only national wealth and human population density were statistically significant predictors in the majority of models when analyzed jointly with climate, geography, and land cover. The economic and demographic variables reflect the intensity of human activities and integrate the effect of factors that directly determine the outcome of invasion such as propagule pressure, pathways of introduction, eutrophication, and the intensity of anthropogenic disturbance. The strong influence of economic and demographic variables on the levels of invasion by alien species demonstrates that future solutions to the problem of biological invasions at a national scale lie in mitigating the negative environmental consequences of human activities that generate wealth and by promoting more sustainable population growth.

Backyard conservation in traditionally owned lands.

Backyard conservation is a community-based approach aiming to weave traditional knowledge, management practices, customary land ownership, and Indigenous-driven solutions with other scientific knowledge to enhance conservation. The islands of Papua New Guinea (PNG) provide an important example towards understanding the role of backyard conservation in culturally diverse and complex societies.

Alien vs. Predator: Impacts of Invasive Species and Native Predators on Urban Nest Box Use by Native Birds.

Many bird species in Australia require tree hollows for breeding. However, assessing the benefits of urban nest boxes to native birds requires frequent monitoring that allows to assess nesting success. To better understand the benefits of nest boxes for native birds, we examined the impact of local habitat characteristics, invasive species (common myna, Acridotheres tristis), and native mammalian predators on urban nest box use and nesting success of native birds. We installed 216 nest boxes across nine locations in southeastern Australia (S.E. Queensland and northern New South Wales) in both long-invaded sites (invaded before 1970) and more recently invaded sites (after 1990). We monitored all boxes weekly over two breeding seasons. We recorded seven bird species and three mammal species using the nest boxes. Weekly box occupancy by all species averaged 8% of all boxes, with the species most frequently recorded in the nest boxes being the common brushtail possum (Trichosurus vulpecula), a native cavity user and nest predator. We recorded 137 nesting attempts in the boxes across all bird species. The most frequent nesting species were the invasive alien common mynas (72 nesting attempts). We recorded an average nesting failure rate of 53.3% for all bird species. We did not record any common mynas evicting other nesting birds, and found that several native species used the same box after the common myna completed its nesting. We recorded native possums in 92% of the boxes, and possum occupancy of boxes per site was negatively correlated with bird nesting success (p = 0.021). These results suggest that when boxes are accessible to invasive species and native predators, they are unlikely to significantly improve nesting opportunities for native birds. To ensure efficient use of limited conservation resources, nest boxes should be designed to target species of high conservation importance and limit other species of both predators and competitors.

Between-country collaboration and consideration of costs increase conservation planning efficiency in the Mediterranean Basin.

The importance of global and regional coordination in conservation is growing, although currently, the majority of conservation programs are applied at national and subnational scales. Nevertheless, multinational programs incur transaction costs and resources beyond what is required in national programs. Given the need to maximize returns on investment within limited conservation budgets, it is crucial to quantify how much more biodiversity can be protected by coordinating multinational conservation efforts when resources are fungible. Previous studies that compared different scales of conservation decision-making mostly ignored spatial variability in biodiversity threats and the cost of actions. Here, we developed a simple integrating metric, taking into account both the cost of conservation and threats to biodiversity. We examined the Mediterranean Basin biodiversity hotspot, which encompasses over 20 countries. We discovered that for vertebrates to achieve similar conservation benefits, one would need substantially more money and area if each country were to act independently as compared to fully coordinated action across the Basin. A fully coordinated conservation plan is expected to save approximately US$67 billion, 45% of total cost, compared with the uncoordinated plan; and if implemented over a 10-year period, the plan would cost approximately 0.1% of the gross national income of all European Union (EU) countries annually. The initiative declared in the recent Paris Summit for the Mediterranean provides a political basis for such complex coordination. Surprisingly, because many conservation priority areas selected are located in EU countries, a partly coordinated solution incorporating only EU-Mediterranean countries is almost as efficient as the fully coordinated scenario.

The COVID-19 pandemic is intricately linked to biodiversity loss and ecosystem health.

The ongoing COVID-19 pandemic, caused by zoonotic SARS-CoV-2, has important links to biodiversity loss and ecosystem health. These links range from anthropogenic activities driving zoonotic disease emergence and extend to the pandemic affecting biodiversity conservation, environmental policy, ecosystem services, and multiple conservation facets. Crucially, such effects can exacerbate the initial drivers, resulting in feedback loops that are likely to promote future zoonotic disease outbreaks. We explore these feedback loops and relationships, highlighting known and potential zoonotic disease emergence drivers (eg, land-use change, intensive livestock production, wildlife trade, and climate change), and discuss direct and indirect effects of the ongoing pandemic on biodiversity loss and ecosystem health. We stress that responses to COVID-19 must include actions aimed at safeguarding biodiversity and ecosystems, in order to avoid future emergence of zoonoses and prevent their wide-ranging effects on human health, economies, and society. Such responses would benefit from adopting a One Health approach, enhancing cross-sector, transboundary communication, as well as from collaboration among multiple actors, promoting planetary and human health.

Motivations for conserving urban biodiversity.

In a time of increasing urbanization, the fundamental value of conserving urban biodiversity remains controversial. How much of a fixed budget should be spent on conservation in urban versus nonurban landscapes? The answer should depend on the goals that drive our conservation actions, yet proponents of urban conservation often fail to specify the motivation for protecting urban biodiversity. This is an important shortcoming on several fronts, including a missed opportunity to make a stronger appeal to those who believe conservation biology should focus exclusively on more natural, wilder landscapes. We argue that urban areas do offer an important venue for conservation biology, but that we must become better at choosing and articulating our goals. We explored seven possible motivations for urban biodiversity conservation: preserving local biodiversity, creating stepping stones to nonurban habitat, understanding and facilitating responses to environmental change, conducting environmental education, providing ecosystem services, fulfilling ethical responsibilities, and improving human well-being. To attain all these goals, challenges must be faced that are common to the urban environment, such as localized pollution, disruption of ecosystem structure, and limited availability of land. There are, however, also challenges specific only to particular goals, meaning that different goals will require different approaches and actions. This highlights the importance of specifying the motivations behind urban biodiversity conservation. If the goals are unknown, progress cannot be assessed.

Marine protected areas for demersal elasmobranchs in highly exploited Mediterranean ecosystems.

Marine ecosystems are complex socio-ecological systems where sustainable solutions can be best gained by satisfying both conservation and socioeconomic demands. Concretely, the Mediterranean Sea is facing a huge demand of resources and marine activities while hosting abundant and unique biodiversity. It is considered an important elasmobranch hotspot where seventy-two elasmobranch species are present in the basin. Despite the recognised importance of elasmobranchs as umbrella species, to date only a small number of marine protected areas have been designated towards their protection. The paucity of spatially-explicit abundance data on elasmobranchs often precludes the designation of these areas to protect these marine predators. Here, we aimed to identify marine areas to protect elasmobranch species by means of a systematic spatial planning approach. We first estimated the spatial distribution of five elasmobranch species (three sharks and two rays) in the western Mediterranean Sea and then applied Marxan decision support tools to find priority marine conservation areas. We found that the five elasmobranchs are distributed in coastal and slope areas of the southern waters of the study area while in the northern region they are abundant in the continental slope and towards offshore waters. Conservation priority areas were identified in the southern part of the western Mediterranean. Adding more complex cost layers and zoning to the analysis did not alter conservation priority areas, confirming such areas are highly consistent and highly important for elasmobranch protection. The marine conservation priority areas identified here can contribute to designate a proactive area-based protection strategy towards elasmobranch conservation, related species and the habitats that they depend in the western Mediterranean Sea.

Ecological variables for developing a global deep-ocean monitoring and conservation strategy.

The deep sea (>200 m depth) encompasses >95% of the world's ocean volume and represents the largest and least explored biome on Earth (<0.0001% of ocean surface), yet is increasingly under threat from multiple direct and indirect anthropogenic pressures. Our ability to preserve both benthic and pelagic deep-sea ecosystems depends upon effective ecosystem-based management strategies and monitoring based on widely agreed deep-sea ecological variables. Here, we identify a set of deep-sea essential ecological variables among five scientific areas of the deep ocean: (1) biodiversity; (2) ecosystem functions; (3) impacts and risk assessment; (4) climate change, adaptation and evolution; and (5) ecosystem conservation. Conducting an expert elicitation (1,155 deep-sea scientists consulted and 112 respondents), our analysis indicates a wide consensus amongst deep-sea experts that monitoring should prioritize large organisms (that is, macro- and megafauna) living in deep waters and in benthic habitats, whereas monitoring of ecosystem functioning should focus on trophic structure and biomass production. Habitat degradation and recovery rates are identified as crucial features for monitoring deep-sea ecosystem health, while global climate change will likely shift bathymetric distributions and cause local extinction in deep-sea species. Finally, deep-sea conservation efforts should focus primarily on vulnerable marine ecosystems and habitat-forming species. Deep-sea observation efforts that prioritize these variables will help to support the implementation of effective management strategies on a global scale.

Human-related processes drive the richness of exotic birds in Europe.

Both human-related and natural factors can affect the establishment and distribution of exotic species. Understanding the relative role of the different factors has important scientific and applied implications. Here, we examined the relative effect of human-related and natural factors in determining the richness of exotic bird species established across Europe. Using hierarchical partitioning, which controls for covariation among factors, we show that the most important factor is the human-related community-level propagule pressure (the number of exotic species introduced), which is often not included in invasion studies due to the lack of information for this early stage in the invasion process. Another, though less important, factor was the human footprint (an index that includes human population size, land use and infrastructure). Biotic and abiotic factors of the environment were of minor importance in shaping the number of established birds when tested at a European extent using 50 x 50 km2 grid squares. We provide, to our knowledge, the first map of the distribution of exotic bird richness in Europe. The richest hotspot of established exotic birds is located in southeastern England, followed by areas in Belgium and The Netherlands. Community-level propagule pressure remains the major factor shaping the distribution of exotic birds also when tested for the UK separately. Thus, studies examining the patterns of establishment should aim at collecting the crucial and hard-to-find information on community-level propagule pressure or develop reliable surrogates for estimating this factor. Allowing future introductions of exotic birds into Europe should be reconsidered carefully, as the number of introduced species is basically the main factor that determines the number established.