Land-use changes interact with geology to facilitate dispersal of the rock hyrax (Procavia capensis) and leishmaniasis across Israel and the West Bank.

Geology plays a fundamental role in establishing species' habitats, determining both physical (e.g., landscape morphology, soil texture) and chemical (e.g., mineral composition, water availability) properties. In the current Anthropocene epoch, human activity is transforming Earth's geology and ecosystems. Yet to date, there have been almost no studies incorporating geology when examining the effect of such land-use changes on species distribution. This study seeks to uncover how specific land-use changes interact with geology, in order to explain the recent and rapid expansion of the rock hyrax (Procavia capensis) across the mountains of central Israel and the West Bank. Hyraxes are dependent on rock mounds for their habitat, and their expansion seems to be correlated with increasing infrastructure construction. However, their expansion patterns differ among locations, even when the human land-use is similar. To explain the patterns of hyrax distribution observed over the past 46 years, we converted geological data into ecological data, which present the probability of the local bedrock breaking into boulders, whether due to either natural or anthropogenic weathering processes. We applied species distribution models (SDMs) and found that the expansion of rock hyrax populations was facilitated by means of the interaction of specific geological units with land-use practices (e.g., roads and construction), which resulted in the accumulation of large boulders, creating novel habitats and stepping stones in previously unsuitable areas for hyraxes. Since rock hyraxes are major hosts of the leishmaniasis pathogen, the findings from this study offer important insights into the progression and potential outbreaks of the disease in human populations. Understanding the role that geology plays in shaping a species' niche is expected to prove useful in studying the distribution of other wildlife species and is fundamental in studies seeking to predict the potential ecological impacts on local biodiversity associated with land-use change.

Climate change maladaptation for health: Agricultural practice against shifting seasonal rainfall affects snakebite risk for farmers in the tropics.

Snakebite affects more than 1.8 million people annually. Factors explaining snakebite variability include farmers' behaviors, snake ecology and climate. One unstudied issue is how farmers' adaptation to novel climates affect their health. Here we examined potential impacts of adaptation on snakebite using individual-based simulations, focusing on strategies meant to counteract major crop yield decline because of changing rainfall in Sri Lanka. For rubber cropping, adaptation led to a 33% increase in snakebite incidence per farmer work hour because of work during risky months, but a 17% decrease in total annual snakebites because of decreased labor in plantations overall. Rice farming adaptation decreased snakebites by 16%, because of shifting labor towards safer months, whereas tea adaptation led to a general increase. These results indicate that adaptation could have both a positive and negative effect, potentially intensified by ENSO. Our research highlights the need for assessing adaptation strategies for potential health maladaptations.

Future temperature extremes threaten land vertebrates.

The frequency, duration, and intensity of extreme thermal events are increasing and are projected to further increase by the end of the century1,2. Despite the considerable consequences of temperature extremes on biological systems3-8, we do not know which species and locations are most exposed worldwide. Here we provide a global assessment of land vertebrates' exposures to future extreme thermal events. We use daily maximum temperature data from 1950 to 2099 to quantify future exposure to high frequency, duration, and intensity of extreme thermal events to land vertebrates. Under a high greenhouse gas emission scenario (Shared Socioeconomic Pathway 5-8.5 (SSP5-8.5); 4.4 °C warmer world), 41.0% of all land vertebrates (31.1% mammals, 25.8% birds, 55.5% amphibians and 51.0% reptiles) will be exposed to extreme thermal events beyond their historical levels in at least half their distribution by 2099. Under intermediate-high (SSP3-7.0; 3.6 °C warmer world) and intermediate (SSP2-4.5; 2.7 °C warmer world) emission scenarios, estimates for all vertebrates are 28.8% and 15.1%, respectively. Importantly, a low-emission future (SSP1-2.6, 1.8 °C warmer world) will greatly reduce the overall exposure of vertebrates (6.1% of species) and can fully prevent exposure in many species assemblages. Mid-latitude assemblages (desert, shrubland, and grassland biomes), rather than tropics9,10, will face the most severe exposure to future extreme thermal events. By 2099, under SSP5-8.5, on average 3,773 species of land vertebrates (11.2%) will face extreme thermal events for more than half a year period. Overall, future extreme thermal events will force many species and assemblages into constant severe thermal stress. Deep greenhouse gas emissions cuts are urgently needed to limit species' exposure to thermal extremes.

Social-ecological cascade effects of land use on vertebrate pest dynamics in arid agricultural communities.

Extensive land conversion to agriculture in drylands and associated resource use have wide-ranging impacts on desert ecosystems globally. Incorporating the impacts of human-social aspects is thus imperative in examining ecological interactions. The provision of agricultural inputs in these resource-scarce regions supports invasive and pest species, negatively impacting both agricultural productivity and native desert ecosystems. Understanding the spatial dynamics of invasive and pest species requires analyzing both bottom-up resource availability factors underlying animal distributions and top-down biological controls. Here, we evaluate the social-ecological cascading effects of dryland agriculture on vertebrate pest communities in dryland agricultural communities of Israel. Our study region is characterized by 18 agricultural cooperatives with distinct crop regimes due to contrasting social decision-making and resource allocation schemes (i.e., communal kibbutzim vs. privatized moshavim). Crop choices further affect land management (e.g., enclosed vs. open farm systems) and resource intensity. This system is ideal for studying trophic mechanisms underlying animal assemblages between agricultural regimes. We examine the role of agricultural land-use practices on pest spatial distributions based on multiyear vertebrate pest observations with agricultural data sets. We use structural equation modeling (SEM) to quantify the relative importance of added agricultural resources underlying bottom-up and top-down trophic processes regulating vertebrate pest assemblages. Results reveal that crop choices determine pest distributions through bottom-up processes directly, while simultaneously driving pest competitive interactions through indirect top-down cascades impacting pest communities. For example, due to the indirect negative effect of wolves on mesopredators (foxes and jackals) mediated by livestock, the total positive effect of livestock on the abundance of mesopredators is reduced. Our study illustrates the social-ecological cascading effects of agricultural regimes on pest community assemblages mediated by contrasting agricultural land-use practices. Considering the expansion of dryland agroecological systems globally, understanding the intricate cascading pathways of predator- and prey-pest communities has important implications for agricultural management, biological invasions in drylands, and fragile desert environments.

Exploring the risks and benefits of flexibility in biodiversity offset location in a case study of migratory shorebirds.

Biodiversity offsets aim to counterbalance the residual impacts of development on species and ecosystems. Guidance documents explicitly recommend that biodiversity offset actions be located close to the location of impact because of higher potential for similar ecological conditions, but allowing greater spatial flexibility has been proposed. We examined the circumstances under which offsets distant from the impact location could be more likely to achieve no net loss or provide better ecological outcomes than offsets close to the impact area. We applied a graphical model for migratory shorebirds in the East Asian-Australasian Flyway as a case study to explore the problems that arise when incorporating spatial flexibility into offset planning. Spatially flexible offsets may alleviate impacts more effectively than local offsets; however, the risks involved can be substantial. For our case study, there were inadequate data to make robust conclusions about the effectiveness and equivalence of distant habitat-based offsets for migratory shorebirds. Decisions around offset placement should be driven by the potential to achieve equivalent ecological outcomes; however, when considering more distant offsets, there is a need to evaluate the likely increased risks alongside the potential benefits. Although spatially flexible offsets have the potential to provide more cost-effective biodiversity outcomes and more cobenefits, our case study showed the difficulty of demonstrating these benefits in practice and the potential risks that need to be considered to ensure effective offset placement.

Estudio de los riesgos y beneficios de la flexibilidad en la ubicación de compensación de la biodiversidad en el estudio de caso de aves costeras migratorias Resumen Las compensaciones de la biodiversidad buscan contrabalancear el impacto residual que tiene el desarrollo sobre las especies y los ecosistemas. Los documentos guía recomiendan explícitamente que las acciones de estas compensaciones estén ubicadas cerca del lugar del impacto debido al potencial elevado de que haya condiciones ecológicas similares, aunque ya hay propuestas de una mayor flexibilidad espacial. Analizamos las circunstancias bajo las cuales las compensaciones alejadas del lugar de impacto tendrían mayor probabilidad de lograr pérdidas netas nulas o de proporcionar mejores resultados ecológicos que las compensaciones cercanas al área de impacto. Aplicamos un modelo gráfico para las aves costeras migratorias en el corredor aéreo asiático-australasiático del este como estudio de caso para estudiar los problemas que surgen cuando se incorpora la flexibilidad espacial a la planeación de las compensaciones. Las compensaciones espacialmente flexibles pueden mitigar los impactos más efectivamente que las compensaciones locales; sin embargo, los riesgos que esto involucra pueden ser considerables. En nuestro estudio de caso hubo datos insuficientes para concluir contundentemente sobre la efectividad y equivalencia de las compensaciones basadas en los hábitats distantes para las aves costeras migratorias. Las decisiones en torno a la ubicación de las compensaciones deberían estar impulsadas por el potencial para obtener resultados ecológicos equivalentes; sin embargo, al considerar compensaciones más alejadas, existe la necesidad de evaluar el incremento probable de riesgos junto a los beneficios potenciales. Aunque las compensaciones espacialmente flexibles tienen el potencial para proporcionar resultados más rentables y más beneficios colaterales, nuestro estudio de caso mostró la dificultad para demostrar estos beneficios en la práctica y los riesgos potenciales que necesitan considerarse para asegurar una ubicación efectiva de las compensaciones.

A mechanistic model of snakebite as a zoonosis: Envenoming incidence is driven by snake ecology, socioeconomics and its impacts on snakes.

Snakebite is the only WHO-listed, not infectious neglected tropical disease (NTD), although its eco-epidemiology is similar to that of zoonotic infections: envenoming occurs after a vertebrate host contacts a human. Accordingly, snakebite risk represents the interaction between snake and human factors, but their quantification has been limited by data availability. Models of infectious disease transmission are instrumental for the mitigation of NTDs and zoonoses. Here, we represented snake-human interactions with disease transmission models to approximate geospatial estimates of snakebite incidence in Sri Lanka, a global hotspot. Snakebites and envenomings are described by the product of snake and human abundance, mirroring directly transmitted zoonoses. We found that human-snake contact rates vary according to land cover (surrogate of occupation and socioeconomic status), the impacts of humans and climate on snake abundance, and by snake species. Our findings show that modelling snakebite as zoonosis provides a mechanistic eco-epidemiological basis to understand snakebites, and the possible implications of global environmental and demographic change for the burden of snakebite.

Implications of global environmental change for the burden of snakebite.

Snakebite envenoming is a set of intoxication diseases that disproportionately affect people of poor socioeconomic backgrounds in tropical countries. As it is highly dependent on the environment its burden is expected to shift spatially with global anthropogenic environmental (climate, land use) and demographic change. The mechanisms underlying the changes to snakebite epidemiology are related to factors of snakes and humans. The distribution and abundance of snakes are expected to change with global warming via their thermal tolerance, while rainfall may affect the timing of key activities like feeding and reproduction. Human population growth is the primary cause of land-use change, which may impact snakes at smaller spatial scales than climate via habitat and biodiversity loss (e.g. prey availability). Human populations, on the other hand, could experience novel patterns and morbidity of snakebite envenoming, both as a result of snake responses to environmental change and due to the development of agricultural adaptations to climate change, socioeconomic and cultural changes, development and availability of better antivenoms, personal protective equipment, and mechanization of agriculture that mediate risk of encounters with snakes and their outcomes. The likely global effects of environmental and demographic change are thus context-dependent and could encompass both increasing and or snakebite burden (incidence, number of cases or morbidity), exposing new populations to snakes in temperate areas due to "tropicalization", or by land use change-induced snake biodiversity loss, respectively. Tackling global change requires drastic measures to ensure large-scale ecosystem functionality. However, as ecosystems represent the main source of venomous snakes their conservation should be accompanied by comprehensive public health campaigns. The challenges associated with the joint efforts of biodiversity conservation and public health professionals should be considered in the global sustainability agenda in a wider context that applies to neglected tropical and zoonotic and emerging diseases.

Evaluating spatiotemporal dynamics of snakebite in Sri Lanka: Monthly incidence mapping from a national representative survey sample.

BACKGROUND: Snakebite incidence shows both spatial and temporal variation. However, no study has evaluated spatiotemporal patterns of snakebites across a country or region in detail. We used a nationally representative population sample to evaluate spatiotemporal patterns of snakebite in Sri Lanka. METHODOLOGY: We conducted a community-based cross-sectional survey representing all nine provinces of Sri Lanka. We interviewed 165 665 people (0.8% of the national population), and snakebite events reported by the respondents were recorded. Sri Lanka is an agricultural country; its central, southern and western parts receive rain mainly from Southwest monsoon (May to September) and northern and eastern parts receive rain mainly from Northeast monsoon (November to February). We developed spatiotemporal models using multivariate Poisson process modelling to explain monthly snakebite and envenoming incidences in the country. These models were developed at the provincial level to explain local spatiotemporal patterns. PRINCIPAL FINDINGS: Snakebites and envenomings showed clear spatiotemporal patterns. Snakebite hotspots were found in North-Central, North-West, South-West and Eastern Sri Lanka. They exhibited biannual seasonal patterns except in South-Western inlands, which showed triannual seasonality. Envenoming hotspots were confined to North-Central, East and South-West parts of the country. Hotspots in North-Central regions showed triannual seasonal patterns and South-West regions had annual patterns. Hotspots remained persistent throughout the year in Eastern regions. The overall monthly snakebite and envenoming incidences in Sri Lanka were 39 (95%CI: 38-40) and 19 (95%CI: 13-30) per 100 000, respectively, translating into 110 000 (95%CI: 107 500-112 500) snakebites and 45 000 (95%CI: 32 000-73 000) envenomings in a calendar year. CONCLUSIONS/SIGNIFICANCE: This study provides information on community-based monthly incidence of snakebites and envenomings over the whole country. Thus, it provides useful insights into healthcare decision-making, such as, prioritizing locations to establish specialized centres for snakebite management and allocating resources based on risk assessments which take into account both location and season.

Limited flexibility in departure timing of migratory passerines at the East-Mediterranean flyway.

The rapid pace of current global warming lead to the advancement of spring migration in the majority of long-distance migratory bird species. While data on arrival timing to breeding grounds in Europe is plentiful, information from the African departure sites are scarce. Here we analysed changes in arrival timing at a stopover site in Israel and any links to Enhanced Vegetation Index (EVI) on the species-specific African non-breeding range in three migratory passerines between 2000-2017. Differences in wing length between early and late arriving individuals were also examined as a proxy for migration distance. We found that male redstart, but not females, advanced arrival to stopover site, but interestingly, not as a response to EVI phenology. Blackcap and barred warbler did not shift arrival timing significantly, although the arrival of blackcap was dependent on EVI. Barred warbler from the early arrival phase had longer wings, suggesting different populations. Our study further supports the existence species-specific migration decisions and inter-sexual differences, which may be triggered by both exogenous (local vegetation condition) and endogenous cues. Given rapid rate of changes in environmental conditions at higher latitudes, some migrants may experience difficulty in the race to match global changes to ensure their survival.

Integrating human behavior and snake ecology with agent-based models to predict snakebite in high risk landscapes.

Snakebite causes more than 1.8 million envenoming cases annually and is a major cause of death in the tropics especially for poor farmers. While both social and ecological factors influence the chance encounter between snakes and people, the spatio-temporal processes underlying snakebites remain poorly explored. Previous research has focused on statistical correlates between snakebites and ecological, sociological, or environmental factors, but the human and snake behavioral patterns that drive the spatio-temporal process have not yet been integrated into a single model. Here we use a bottom-up simulation approach using agent-based modelling (ABM) parameterized with datasets from Sri Lanka, a snakebite hotspot, to characterise the mechanisms of snakebite and identify risk factors. Spatio-temporal dynamics of snakebite risks are examined through the model incorporating six snake species and three farmer types (rice, tea, and rubber). We find that snakebites are mainly climatically driven, but the risks also depend on farmer types due to working schedules as well as species present in landscapes. Snake species are differentiated by both distribution and by habitat preference, and farmers are differentiated by working patterns that are climatically driven, and the combination of these factors leads to unique encounter rates for different landcover types as well as locations. Validation using epidemiological studies demonstrated that our model can explain observed patterns, including temporal patterns of snakebite incidence, and relative contribution of bites by each snake species. Our predictions can be used to generate hypotheses and inform future studies and decision makers. Additionally, our model is transferable to other locations with high snakebite burden as well.

Shelter availability and human attitudes as drivers of rock hyrax (Procavia capensis) expansion along a rural-urban gradient.

While anthropogenic land-use changes threaten wildlife globally, some species take advantage of such changes and disperse into urban areas. The wildlife in urban areas often promotes conflicts with humans, notably when the animals are associated with the spread of zoonotic diseases. In Israel, current urban invasion of rock hyraxes (Procavia capensis) draws public attention, since the species is a reservoir host of cutaneous leishmaniasis, a serious skin disease. The rock hyrax, however, has seldom been studied in densely populated areas, and the drivers for its urban expansion, as well as its abilities to live and spread in core urban areas, are relatively unknown. Here, we explore the rock hyrax expansion to urban areas process by examining the availability, characteristics and use of shelter along an urban gradient. Our findings suggest that a series of factors determines shelter availability and quality for the rock hyrax, which facilitates its dispersion across the urban gradient. We found that rock hyraxes from the Judean Desert expand to the peri-urban region of Jerusalem by colonizing new rocky shelters formed as by-products of urban development. With their populations reaching extreme densities in this area and saturating the available shelters, there is some spill over to the adjacent core urban areas where they colonize littered sites, which are made available due to the local socio-economic conditions and cultural norms of waste disposal and illegal placement of temporary structures. Our work emphasizes the significance of the urban gradient approach for studying the mechanisms promoting wildlife expansion to cities. Our findings suggest that changes in shelter availability and quality due to urban development, and cultural norms promote shifts of the hyrax population by pushing from the already established areas and pulling into new environment across the urban gradient.

Accelerating invasion potential of disease vector Aedes aegypti under climate change.

Vector-borne diseases remain a major contributor to the global burden of disease, while climate change is expected to exacerbate their risk. Characterising vector development rate and its spatio-temporal variation under climate change is central to assessing the changing basis of human disease risk. We develop a mechanistic phenology model and apply it to Aedes aegypti, an invasive mosquito vector for arboviruses (e.g. dengue, zika and yellow fever). The model predicts the number of life-cycle completions (LCC) for a given location per unit time based on empirically derived biophysical responses to environmental conditions. Results suggest that the world became ~1.5% more suitable per decade for the development of Ae. aegypti during 1950-2000, while this trend is predicted to accelerate to 3.2-4.4% per decade by 2050. Invasion fronts in North America and China are projected to accelerate from ~2 to 6 km/yr by 2050. An increase in peak LCC combined with extended periods suitable for mosquito development is simulated to accelerate the vector's global invasion potential.

Adjusting for spatial variation when assessing individual-level risk: A case-study in the epidemiology of snake-bite in Sri Lanka.

BACKGROUND: Health outcomes and causality are usually assessed with individual level sociodemographic variables. Studies that consider only individual-level variables can suffer from residual confounding. This can result in individual variables that are unrelated to risk behaving as proxies for uncaptured information. There is a scarcity of literature on risk factors for snakebite. In this study, we evaluate the individual-level risk factors of snakebite in Sri Lanka and highlight the impact of spatial confounding on determining the individual-level risk effects. METHODS: Data was obtained from the National Snakebite Survey of Sri Lanka. This was an Island-wide community-based survey. The survey sampled 165,665 individuals from all 25 districts of the country. We used generalized linear models to identify individual-level factors that contribute to an individual's risk of experiencing a snakebite event. We fitted separate models to assess risk factors with and without considering spatial variation in snakebite incidence in the country. RESULTS: Both spatially adjusted and non-adjusted models revealed that middle-aged people, males, field workers and individuals with low level of education have high risk of snakebites. The model without spatial adjustment showed an interaction between ethnicity and income levels. When the model included a spatial adjustment for the overall snakebite incidence, this interaction disappeared and income level appeared as an independent risk factor. Both models showed similar effect sizes for gender and age. HEmployment and education showed lower effect sizes in the spatially adjusted model. CONCLUSIONS: Both individual-level characteristics and local snakebite incidence are important to determine snakebite risk at a given location. Individual level variables could act as proxies for underling residual spatial variation when environmental information is not considered. This can lead to misinterpretation of risk factors and biased estimates of effect sizes. Both individual-level and environmental variables are important in assessing causality in epidemiological studies.

Adapting environmental management to uncertain but inevitable change.

Implementation of adaptation actions to protect biodiversity is limited by uncertainty about the future. One reason for this is the fear of making the wrong decisions caused by the myriad future scenarios presented to decision-makers. We propose an adaptive management (AM) method for optimally managing a population under uncertain and changing habitat conditions. Our approach incorporates multiple future scenarios and continually learns the best management strategy from observations, even as conditions change. We demonstrate the performance of our AM approach by applying it to the spatial management of migratory shorebird habitats on the East Asian-Australasian flyway, predicted to be severely impacted by future sea-level rise. By accounting for non-stationary dynamics, our solution protects 25,000 more birds per year than the current best stationary approach. Our approach can be applied to many ecological systems that require efficient adaptation strategies for an uncertain future.

Optimal management of a multispecies shorebird flyway under sea-level rise.

Every year, millions of migratory shorebirds fly through the East Asian-Australasian Flyway between their arctic breeding grounds and Australasia. This flyway includes numerous coastal wetlands in Asia and the Pacific that are used as stopover sites where birds rest and feed. Loss of a few important stopover sites through sea-level rise (SLR) could cause sudden population declines. We formulated and solved mathematically the problem of how to identify the most important stopover sites to minimize losses of bird populations across flyways by conserving land that facilitates upshore shifts of tidal flats in response to SLR. To guide conservation investment that minimizes losses of migratory bird populations during migration, we developed a spatially explicit flyway model coupled with a maximum flow algorithm. Migratory routes of 10 shorebird taxa were modeled in a graph theoretic framework by representing clusters of important wetlands as nodes and the number of birds flying between 2 nodes as edges. We also evaluated several resource allocation algorithms that required only partial information on flyway connectivity (node strategy, based on the impacts of SLR at nodes; habitat strategy, based on habitat change at sites; population strategy, based on population change at sites; and random investment). The resource allocation algorithms based on flyway information performed on average 15% better than simpler allocations based on patterns of habitat loss or local bird counts. The Yellow Sea region stood out as the most important priority for effective conservation of migratory shorebirds, but investment in this area alone will not ensure the persistence of species across the flyway. The spatial distribution of conservation investments differed enormously according to the severity of SLR and whether information about flyway connectivity was used to guide the prioritizations. With the rapid ongoing loss of coastal wetlands globally, our method provides insight into efficient conservation planning for migratory species.

Targeting global protected area expansion for imperiled biodiversity.

Governments have agreed to expand the global protected area network from 13% to 17% of the world's land surface by 2020 (Aichi target 11) and to prevent the further loss of known threatened species (Aichi target 12). These targets are interdependent, as protected areas can stem biodiversity loss when strategically located and effectively managed. However, the global protected area estate is currently biased toward locations that are cheap to protect and away from important areas for biodiversity. Here we use data on the distribution of protected areas and threatened terrestrial birds, mammals, and amphibians to assess current and possible future coverage of these species under the convention. We discover that 17% of the 4,118 threatened vertebrates are not found in a single protected area and that fully 85% are not adequately covered (i.e., to a level consistent with their likely persistence). Using systematic conservation planning, we show that expanding protected areas to reach 17% coverage by protecting the cheapest land, even if ecoregionally representative, would increase the number of threatened vertebrates covered by only 6%. However, the nonlinear relationship between the cost of acquiring land and species coverage means that fivefold more threatened vertebrates could be adequately covered for only 1.5 times the cost of the cheapest solution, if cost efficiency and threatened vertebrates are both incorporated into protected area decision making. These results are robust to known errors in the vertebrate range maps. The Convention on Biological Diversity targets may stimulate major expansion of the global protected area estate. If this expansion is to secure a future for imperiled species, new protected areas must be sited more strategically than is presently the case.

Migratory connectivity magnifies the consequences of habitat loss from sea-level rise for shorebird populations.

Sea-level rise (SLR) will greatly alter littoral ecosystems, causing habitat change and loss for coastal species. Habitat loss is widely used as a measurement of the risk of extinction, but because many coastal species are migratory, the impact of habitat loss will depend not only on its extent, but also on where it occurs. Here, we develop a novel graph-theoretic approach to measure the vulnerability of a migratory network to the impact of habitat loss from SLR based on population flow through the network. We show that reductions in population flow far exceed the proportion of habitat lost for 10 long-distance migrant shorebirds using the East Asian-Australasian Flyway. We estimate that SLR will inundate 23-40% of intertidal habitat area along their migration routes, but cause a reduction in population flow of up to 72 per cent across the taxa. This magnifying effect was particularly strong for taxa whose migration routes contain bottlenecks-sites through which a large fraction of the population travels. We develop the bottleneck index, a new network metric that positively correlates with the predicted impacts of habitat loss on overall population flow. Our results indicate that migratory species are at greater risk than previously realized.