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In addition to habitat loss and fragmentation, demographic processes—the vagaries of births, deaths and sex ratio fluctuations—pose substantial threats to wild giant panda populations. Additionally, climate change and plans for the Giant Panda National Park may influence (in opposing directions) the extinction risk for wild giant pandas. The Fourth National Giant Panda Census showed pandas living in 33 isolated populations. An estimated 259 animals live in 25 of these groups, ~14% of the total population. We used individual-based models to simulate time series of these small populations for 100 years. We analysed the spatial pattern of their risk of extinction under current conditions and multiple climate change models. Furthermore, we consider the impact of the proposed Giant Panda National Park. Results showed that 15 populations face a risk >90%, and for 3 other populations the risk is >50%. Of the 15 most at-risk populations, national parks can protect only 3. Under the Representative Concentration Pathway 8.5 climate change scenario, the 33 populations will probably further divide into 56 populations. Some 41 of them will face a risk >50% and 35 face a risk >90%. Although national parks will probably connect some fragmented habitats, 26 populations will be outside national park planning. Our study gives practical advice for conservation policies and management and has implications for the conservation of other species in the world that live in isolated, fragmented habitats. The authors model likely outcomes for the 33 isolated populations reported in the Fourth National Giant Panda Census under multiple RCP scenarios and with the provision of the planned Giant Panda National Park. They find that, although the National Park may connect some fragmented habitats, most of the populations with high extinction risk fall outside the current plans.

Conservation science is a new and evolving discipline, so it seems prudent to explore different approaches. That said, we should examine what we know and, vitally, what works to conserve biodiversity and what does not. Ecosystem processes determine the fate of many species, but many attempts to theorise about ecosystems have led to ever more fanciful descriptions of nature. All conservation is local. It will only succeed if we find ways to accommodate people and nature. That does not mean indigenous knowledge acquired over millennia will be sufficient to our ever more overcrowded planet. Observational and experimental studies of small populations of wild species, however, do provide practical insights into how to manage biodiversity across much larger geographical extents.

Accurate maps of species ranges are essential to inform conservation, but time-consuming to produce and update. Given the pace of change of knowledge about species distributions and shifts in ranges under climate change and land use, a need exists for timely mapping approaches that enable batch processing employing widely available data. We develop a systematic approach of batch-processing range maps and derived Area of Habitat maps for terrestrial bird species with published ranges below 125,000 km2 in Central and South America. (Area of Habitat is the habitat available to a species within its range.) We combine existing range maps with the rapidly expanding crowd-sourced eBird data of presences and absences from frequently surveyed locations, plus readily accessible, high resolution satellite data on forest cover and elevation to map the Area of Habitat available to each species. Users can interrogate the maps produced to see details of the observations that contributed to the ranges. Previous estimates of Areas of Habitat were constrained within the published ranges and thus were, by definition, smaller—typically about 30%. This reflects how little habitat within suitable elevation ranges exists within the published ranges. Our results show that on average, Areas of Habitat are 12% larger than published ranges, reflecting the often-considerable extent that eBird records expand the known distributions of species. Interestingly, there are substantial differences between threatened and non-threatened species. Some 40% of Critically Endangered, 43% of Endangered, and 55% of Vulnerable species have Areas of Habitat larger than their published ranges, compared with 31% for Near Threatened and Least Concern species. The important finding for conservation is that threatened species are generally more widespread than previously estimated.

Different models of community dynamics, such as the MacArthur–Wilson theory of island biogeography and Hubbell’s neutral theory, have given us useful insights into the workings of ecological communities. Here, we develop the niche-hypervolume concept of the community into a powerful model of community dynamics. We describe the community’s size through the volume of the hypercube and the dynamics of the populations in it through the fluctuations of the axes of the niche hypercube on different timescales. While the community’s size remains constant, the relative volumes of the niches within it change continuously, thus allowing the populations of different species to rise and fall in a zero-sum fashion. This dynamic hypercube model reproduces several key patterns in communities: lognormal species abundance distributions, 1/f-noise population abundance, multiscale patterns of extinction debt and logarithmic species-time curves. It also provides a powerful framework to explore significant ideas in ecology, such as the drift of ecological communities into evolutionary time.

Long-term studies to understand biodiversity changes remain scarce-especially so for tropical mountains. We examined changes from 1911 to 2016 in the bird community of the cloud forest of San Antonio, a mountain ridge in the Colombian Andes. We evaluated the effects of past land-use change and assessed species vulnerability to climate disruption. Forest cover decreased from 95% to 50% by 1959, and 33 forest species were extirpated. From 1959 to 1990, forest cover remained stable, and an additional 15 species were lost-a total of 29% of the forest bird community. Thereafter, forest cover increased by 26% and 17 species recolonized the area. The main cause of extirpations was the loss of connections to adjacent forests. Of the 31 (19%) extirpated birds, 25 have ranges peripheral to San Antonio, mostly in the lowlands. Most still occurred regionally, but broken forest connections limited their recolonization. Other causes of extirpation were hunting, wildlife trade, and water diversion. Bird community changes included a shift from predominantly common species to rare species; forest generalists replaced forest specialists that require old growth, and functional groups, such as large-body frugivores and nectarivores, declined disproportionally. All water-dependent birds were extirpated. Of the remaining 122 forest species, 19 are vulnerable to climate disruption, 10 have declined in abundance, and 4 are threatened. Our results show unequivocal species losses and changes in community structure and abundance at the local scale. We found species were extirpated after habitat loss and fragmentation, but forest recovery stopped extirpations and helped species repopulate. Land-use changes increased species vulnerability to climate change, and we suggest reversing landscape transformation may restore biodiversity and improve resistance to future threats.Extirpaciones de Aves y las Dinámicas Comunitarias en un Bosque Nuboso Andino durante más de Cien Años de Cambios de Uso de Suelo Resumen Los estudios a largo plazo para entender cambios en la biodiversidad todavía son escasos - especialmente en las montañas tropicales. Examinamos los cambios en la comunidad de aves del bosque de San Antonio en los Andes colombianos, desde 1911 hasta 2016. Evaluamos los efectos pasados del cambio en el uso del suelo y analizamos la vulnerabilidad de las especies ante la disrupción climática. La cobertura del bosque disminuyó del 95% al 50% para el año 1959, y 33 especies de bosque fueron extirpadas. Desde 1959 y hasta 1990, la cobertura permaneció estable y se perdieron 15 especies más - un total del 29% de la comunidad de aves de bosque. A partir de ahí la cobertura del bosque incrementó en un 26% y 17 especies recolonizaron el área. La principal causa de las extirpaciones fue la pérdida de conectividad con los bosques adyacentes. De las 31 (19%) especies de aves extirpadas, 25 especies tienen una distribución periférica a San Antonio, principalmente en las tierras bajas. La mayoría de las especies aún tenían presencia regional, pero las conexiones rotas entre los bosques limitaron su recolonización. Otras causas de las extirpaciones fueron la caza, el mercado de fauna y el desvío de cursos de agua. Los cambios en la comunidad de aves incluyeron una transición de especies predominantemente comunes a especies raras; las especies generalistas de bosque reemplazaron a las especies especialistas que requieren bosques maduros y los grupos funcionales, como los grandes frugívoros y nectarívoros, declinaron desproporcionadamente. Todas las aves dependientes del agua fueron extirpadas. De las 122 especies que permanecen en el bosque, 19 son vulnerables a la disrupción climática, diez han disminuido en abundancia y cuatro se encuentran amenazadas. Nuestros resultados muestran una pérdida inequívoca de especies y cambios en la estructura y abundancia de la comunidad de aves a escala local. En general, encontramos que las especies fueron extirpadas después de la pérdida y fragmentación del hábitat, pero la recuperación del bosque detuvo las extirpaciones y ayudó a las especies a recolonizar. Los cambios en el uso de suelo incrementaron la vulnerabilidad de las especies ante el cambio climático, por lo que sugerimos que revertir la transformación del paisaje podría restaurar la biodiversidad y aumentar la resistencia a futuras amenazas.目前, 关于生物多样性变化的长期研究还很少见, 特别是在热带山区。我们研究了哥伦比亚安第斯山脉圣安东尼奥地区一个云雾林在 1911-2016 年间的鸟类群落变化。本研究评估了过去土地利用变化的影响和物种对气候变化的脆弱性。 1911-1959 年, 该地区森林覆盖率由 95% 下降到 50%, 33 种森林物种局部灭绝。 1959-1990 年, 森林覆盖率保持稳定, 但仍有 15 种鸟类绝迹, 占森林鸟类群落总数的 29% 。此后, 森林覆盖率增加了 26%, 且有 17 个物种在该地区重新定殖。物种局部灭绝的主要原因是相邻森林之间的连接被破坏。在 31 种 (19%) 灭绝的鸟类中, 25 种在圣安东尼奥周围也有分布, 且主要在低地地区。这些鸟类大多仍有区域性分布, 但森林连接被破坏限制了它们重新定殖。局部灭绝的其它原因还包括狩猎、野生动物贸易和引水工程。鸟类群落的变化包括占优势的鸟种从常见种向稀有种转变; 森林中的广幅种取代了需要原始林的狭域种; 而包括大型食果动物和食蜜动物在内的功能群则不成比例地减少。所有依赖水体的鸟类都消失了。在残余的 122 种森林物种中, 19 种易受到气候变化的影响, 10 种种群数量已大量下降, 还有 4 种受到胁迫。我们的研究结果展现了局部尺度上明确的物种灭绝和种群结构及数量变化。一般来说, 栖息地丧失和破碎化会导致物种局部灭绝, 但森林恢复可以阻止灭绝, 并帮助物种重建种群。土地利用变化增加了物种对气候变化的脆弱性, 我们认为逆转景观改造有助于恢复生物多样性, 并提高物种对未来威胁的抵抗力。【翻译: 胡怡思; 审校: 聂永刚】.

The International Union for Conservation of Nature (IUCN) Red List of Threatened Species includes assessment of extinction risk for 98 512 species, plus documentation of their range, habitat, elevation, and other factors. These range, habitat and elevation data can be matched with terrestrial land cover and elevation datasets to map the species’ area of habitat (AOH; also known as extent of suitable habitat; ESH). This differs from the two spatial metrics used for assessing extinction risk in the IUCN Red List criteria: extent of occurrence (EOO) and area of occupancy (AOO). AOH can guide conservation, for example, through targeting areas for field surveys, assessing proportions of species’ habitat within protected areas, and monitoring habitat loss and fragmentation. We recommend that IUCN Red List assessments document AOH wherever practical., Trends in Ecology & Evolution, 34 (11), ISSN:0169-5347, ISSN:1872-8383

As a landscape becomes increasingly fragmented through habitat loss, the individual patches become smaller and more isolated and thus less likely to sustain a local population. Metapopulation theory is appropriate for analyzing fragmented landscapes because it combines empirical landscape features with species‐specific information to produce direct information on population extinction risks. This approach contrasts with descriptions of habitat fragments, which provide only indirect information on risk. Combining a spatially explicit metapopulation model with empirical data on endemic species’ ranges and maps of habitat cover, we calculated the metapopulation capacity—a measure of a landscape's ability to sustain a metapopulation. Mangroves provide an ideal model landscape because they are of conservation concern and their patch boundaries are easily delineated. For 2000–20015, we calculated global metapopulation capacity for 99 metapopulations of 32 different bird species endemic to mangroves. Northern Australia and Southeast Asia had the highest richness of mangrove endemic birds. The Caribbean, Pacific coast of Central America, Madagascar, Borneo, and isolated patches in Southeast Asia in Myanmar and Malaysia had the highest metapopulation losses. Regions with the highest loss of habitat area were not necessarily those with the highest loss of metapopulation capacity. Often, it was not a matter of how much, but how the habitat was lost. Our method can be used by managers to evaluate and prioritize a landscape for metapopulation persistence., Article impact statement: Application of metapopulation theory shows where area of habitat loss is highest; metapopulation capacity is not necessarily also the highest.

Evidence-based approaches are key for underpinning effective conservation practice, but major gaps in the evidence of the effectiveness of interventions limit their use. Conservation practitioners could make major contributions to filling these gaps but often lack the time, funding, or capacity to do so properly. Many funders target the delivery of conservation and can be reluctant to fund primary research. We analysed the literature testing the effectiveness of interventions. Of a sample of 1,265 publications published in 2019 that tested conservation interventions, 96% included academics. Only 21% included conservation practitioners, of which just under half were first or last author. A community of conservation funders and practitioners undertook a series of workshops to explore means of improving the quality and quantity of intervention testing. A survey of the suggested proportion of conservation grants that should be allocated to testing intervention effectiveness showed practitioners tended to prefer larger percentages (median 3-6%) than funders (median 1-3%), but the overlap was considerable. Funders can facilitate the testing of interventions through a range of measures, including welcoming applications that incorporate testing, allocating funds to testing, and providing training and support to deliver testing. The funders represented by the authors of this paper have committed to these actions. Practitioners can contribute by committing to routine testing, benefiting from funding allocated specifically to testing, and establishing processes for testing interventions. The organisations of the practitioner authors have committed to test at least one intervention per year and share findings, regardless of outcome. Currently, practitioners rarely lead the testing of conservation actions. We suggest processes by which both funders and practitioners can make this routine. This will not only improve the effectiveness and cost-efficiency of practice, but also make conservation more attractive to funders.

The cause of deaths of 350 elephants in 2020 in a relatively small unprotected area of northern Botswana is unknown, and may never be known. Media speculations about it ignore ecological realities. Worse, they make conjectures that can be detrimental to wildlife and sometimes discredit conservation incentives. A broader understanding of the ecological and conservation issues speaks to elephant management across the Kavango–Zambezi Transfrontier Conservation Area that extends across Botswana, Namibia, Angola, Zambia, and Zimbabwe. Our communication addresses these. Malicious poisoning and poaching are unlikely to have played a role. Other species were unaffected, and elephant carcases had their tusks intact. Restriction of freshwater supplies that force elephants to use pans as a water source possibly polluted by blue-green algae blooms is a possible cause, but as yet not supported by evidence. No other species were involved. A contagious disease is the more probable one. Fences and a deep channel of water confine these elephants’ dispersal. These factors explain the elephants’ relatively high population growth rate despite a spell of increased poaching during 2014–2018. While the deaths represent only ~2% of the area’s elephants, the additive effects of poaching and stress induced by people protecting their crops cause alarm. Confinement and relatively high densities probably explain why the die-off occurred only here. It suggests a re-alignment or removal of fences that restrict elephant movements and limits year-round access to freshwater.

International audience; Human actions challenge nature in many ways. Ecological responses are ineluctably complex, demanding measures that describe them succinctly. Collectively, these measures encapsulate the overall ‘stability’ of the system. Many international bodies, including the Intergovernmental Science‐Policy Platform on Biodiversity and Ecosystem Services, broadly aspire to maintain or enhance ecological stability. Such bodies frequently use terms pertaining to stability that lack clear definition. Consequently, we cannot measure them and so they disconnect from a large body of theoretical and empirical understanding. We assess the scientific and policy literature and show that this disconnect is one consequence of an inconsistent and one‐dimensional approach that ecologists have taken to both disturbances and stability. This has led to confused communication of the nature of stability and the level of our insight into it. Disturbances and stability are multidimensional. Our understanding of them is not. We have a remarkably poor understanding of the impacts on stability of the characteristics that define many, perhaps all, of the most important elements of global change. We provide recommendations for theoreticians, empiricists and policymakers on how to better integrate the multidimensional nature of ecological stability into their research, policies and actions.

International audience; The terms sustainability, resilience and others group under the heading of ‘stability’. Their ubiquity speaks to a vital need to characterize changes in complex social and environmental systems. In a bewildering array of terms, practical measurements are essential to permit comparisons and so untangle underlying relationships.

We analyze recently-collected feather tissues from two species of seabirds, the sooty tern (Onychoprion fuscatus) and brown noddy (Anous stolidus), in three ocean regions (North Atlantic, North Pacific, South Pacific) with different human impacts. The species are similar morphologically and are similar in the trophic levels from which they feed within each location. In contrast, we detect reliable differences in trophic position amongst the regions. Trophic position appears to decline as the intensity of commercial fishing increases, and is at its lowest in the Caribbean. The spatial gradient in trophic position we document in these regions exceeds those detected over specimens from the last 130 years in the Hawaiian Islands. Modeling suggests that climate velocity and human impacts on fish populations strongly align with these differences.

Just protecting Earth’s wilderness will not be sufficient to conserve most of its species; quality, not overall size, matters., It is theoretically possible to protect large fractions of species in relatively small regions. For plants, 85% of species occur entirely within just over a third of the Earth’s land surface, carefully optimized to maximize the species captured. Well-known vertebrate taxa show similar patterns. Protecting half of Earth might not be necessary, but would it be sufficient given the current trends of protection? The predilection of national governments is to protect areas that are “wild,” that is, typically remote, cold, or arid. Unfortunately, those areas often hold relatively few species. Wild places likely afford the easier opportunities for the future expansion of protected areas, with the expansion into human-dominated landscapes the greater challenge. We identify regions that are not currently protected, but that are wild, and consider which of them hold substantial numbers of especially small-ranged vertebrate species. We assess how successful the strategy of protecting the wilder half of Earth might be in conserving biodiversity. It is far from sufficient. (Protecting large wild places for reasons other than biodiversity protection, such as carbon sequestration and other ecosystem services, might still have importance.) Unexpectedly, we also show that, despite the bias in establishing large protected areas in wild places to date, numerous small protected areas are in biodiverse places. They at least partially protect significant fractions of especially small-ranged species. So, while a preoccupation with protecting large areas for the sake of getting half of Earth might achieve little for biodiversity, there is more progress in protecting high-biodiversity areas than currently appreciated. Continuing to prioritize the right parts of Earth, not just the total area protected, is what matters for biodiversity.

The notion of a ‘safe operating space for biodiversity’ is vague and encourages harmful policies. Attempts to fix it strip it of all meaningful content. Ecology is rapidly gaining insights into the connections between biodiversity and ecosystem stability. We have no option but to understand ecological complexity and act accordingly.

The giant panda attracts disproportionate conservation resources. How well does this emphasis protect other endemic species? Detailed data on geographical ranges are not available for plants or invertebrates, so we restrict our analyses to 3 vertebrate taxa: birds, mammals, and amphibians. There are gaps in their protection, and we recommend practical actions to fill them. We identified patterns of species richness, then identified which species are endemic to China, and then which, like the panda, live in forests. After refining each species' range by its known elevational range and remaining forest habitats as determined from remote sensing, we identified the top 5% richest areas as the centers of endemism. Southern mountains, especially the eastern Hengduan Mountains, were centers for all 3 taxa. Over 96% of the panda habitat overlapped the endemic centers. Thus, investing in almost any panda habitat will benefit many other endemics. Existing panda national nature reserves cover all but one of the endemic species that overlap with the panda's distribution. Of particular interest are 14 mammal, 20 bird, and 82 amphibian species that are inadequately protected. Most of these species the International Union for Conservation of Nature currently deems threatened. But 7 mammal, 3 bird, and 20 amphibian species are currently nonthreatened, yet their geographical ranges are

Technologies to identify individual animals, follow their movements, identify and locate animal and plant species, and assess the status of their habitats remotely have become better, faster, and cheaper as threats to the survival of species are increasing. New technologies alone do not save species, and new data create new problems. For example, improving technologies alone cannot prevent poaching: solutions require providing appropriate tools to the right people. Habitat loss is another driver: the challenge here is to connect existing sophisticated remote sensing with species occurrence data to predict where species remain. Other challenges include assembling a wider public to crowdsource data, managing the massive quantities of data generated, and developing solutions to rapidly emerging threats.

The global total of protected areas to conserve biodiversity is increasing steadily, while numerous studies show that they are broadly effective. That said, how will current conservation strategies work, given the current and expected changes to the global climate? The giant panda is a conservation icon and exceptional efforts protect its remaining habitats. It provides a unique case study to address this question. There are many studies on the projected loss of habitats as climate warms, but few consider the geographical arrangement of future habitats, current protected area, and species’ dispersal abilities. Most alarmingly, we expect much greater habitat fragmentation after climate change. Here, we combine long-term data on giant pandas with climate-change scenarios to predict future habitat loss and distribution in the Min Shan of Sichuan and Gansu, China. We employ metapopulation capacity as a mechanistic measure of a species’ response to habitat fragmentation. The results show that climate changes will lead to 16.3 ± 1.4 (%) losses of giant panda habitats. Alarmingly, 11.4% of the remaining habitat fragments would be smaller than the extinction threshold area as the extent of fragmentation increases nearly fourfold. The projected fragmentation of giant panda habitats predicts 9% lower effectiveness inside the protected area network compared with that outside of reserves. A 35% reduction will occur in future effectiveness of reserve networks. The results challenge the long-term effectiveness of protected areas in protecting the species’ persistence. They indicate a need for integrating both natural processes and dynamic threats over a simple reliance on individual static natural reserves.

The Eastern Arc Mountains of Tanzania and the Atlantic Forest of Brazil are two of the most fragmented biodiversity hotspots. Species–area relationships predict that their habitat fragments will experience a substantial loss of species. Most of these extinctions will occur over an extended time, and therefore, reconnecting fragments could prevent species losses and allow locally extinct species to recolonize former habitats. An empirical relaxation half-life vs. area relationship for tropical bird communities estimates the time that it takes to lose one-half of all species that will be eventually lost. We use it to estimate the increase in species persistence by regenerating a forest connection 1 km in width among the largest and closest fragments at 11 locations. In the Eastern Arc Mountains, regenerating 8,134 ha of forest would create >316,000 ha in total of restored contiguous forest. More importantly, it would increase the persistence time for species by a factor of 6.8 per location or ∼2,272 years, on average, relative to individual fragments. In the Atlantic Forest, regenerating 6,452 ha of forest would create >251,000 ha in total of restored contiguous forest and enhance species persistence by a factor of 13.0 per location or ∼5,102 years, on average, relative to individual fragments. Rapidly regenerating forest among fragments is important, because mean time to the first determined extinction across all fragments is 7 years. We estimate the cost of forest regeneration at $21–$49 million dollars. It could provide one of the highest returns on investment for biodiversity conservation worldwide.

Migratory seabirds face threats from climate change and a variety of anthropogenic disturbances. Although most seabird research has focused on the ecology of individuals at the colony, technological advances now allow researchers to track seabird movements at sea and during migration. We combined telemetry data on Onychoprion fuscatus (sooty terns) with a long-term capture-mark-recapture dataset from the Dry Tortugas National Park to map the movements at sea for this species, calculate estimates of mortality, and investigate the impact of hurricanes on a migratory seabird. Included in the latter analysis is information on the locations of recovered bands from deceased individuals wrecked by tropical storms. We present the first known map of sooty tern migration in the Atlantic Ocean. Our results indicate that the birds had minor overlaps with areas affected by the major 2010 oil spill and a major shrimp fishery. Indices of hurricane strength and occurrence are positively correlated with annual mortality and indices of numbers of wrecked birds. As climate change may lead to an increase in severity and frequency of major hurricanes, this may pose a long-term problem for this colony.

The conservation status of the iconic giant panda is a barometer of global conservation efforts. The IUCN Red List has downgraded the panda’s extinction risk from “endangered” to “vulnerable”. Newly obtained, detailed GIS and remotely sensed data applied consistently over the last four decades show that panda habitat covered less area and was more fragmented in 2013 than in 1988 when the species was listed as endangered. Despite recent IUCN downgrading of the giant panda’s conservation status from ‘endangered’ to ‘vulnerable’, new GIS and remote sensing data reveal panda habitats to cover less area and be more fragmented than previously.

A key measure of humanity's global impact is by how much it has increased species extinction rates. Familiar statements are that these are 100–1000 times pre-human or background extinction levels. Estimating recent rates is straightforward, but establishing a background rate for comparison is not. Previous researchers chose an approximate benchmark of 1 extinction per million species per year (E/MSY). We explored disparate lines of evidence that suggest a substantially lower estimate. Fossil data yield direct estimates of extinction rates, but they are temporally coarse, mostly limited to marine hard-bodied taxa, and generally involve genera not species. Based on these data, typical background loss is 0.01 genera per million genera per year. Molecular phylogenies are available for more taxa and ecosystems, but it is debated whether they can be used to estimate separately speciation and extinction rates. We selected data to address known concerns and used them to determine median extinction estimates from statistical distributions of probable values for terrestrial plants and animals. We then created simulations to explore effects of violating model assumptions. Finally, we compiled estimates of diversification—the difference between speciation and extinction rates for different taxa. Median estimates of extinction rates ranged from 0.023 to 0.135 E/MSY. Simulation results suggested over- and under-estimation of extinction from individual phylogenies partially canceled each other out when large sets of phylogenies were analyzed. There was no evidence for recent and widespread pre-human overall declines in diversity. This implies that average extinction rates are less than average diversification rates. Median diversification rates were 0.05–0.2 new species per million species per year. On the basis of these results, we concluded that typical rates of background extinction may be closer to 0.1 E/MSY. Thus, current extinction rates are 1,000 times higher than natural background rates of extinction and future rates are likely to be 10,000 times higher. Estimacion de la Tasa Normal de Extincion de Especies

We aspired to set conservation priorities in ways that lead to direct conservation actions. Very large-scale strategic mapping leads to familiar conservation priorities exemplified by biodiversity hotspots. In contrast, tactical conservation actions unfold on much smaller geographical extents and they need to reflect the habitat loss and fragmentation that have sharply restricted where species now live. Our aspirations for direct, practical actions were demanding. First, we identified the global, strategic conservation priorities and then downscaled to practical local actions within the selected priorities. In doing this, we recognized the limitations of incomplete information. We started such a process in Colombia and used the results presented here to implement reforestation of degraded land to prevent the isolation of a large area of cloud forest. We used existing range maps of 171 bird species to identify priority conservation areas that would conserve the greatest number of species at risk in Colombia. By at risk species, we mean those that are endemic and have small ranges. The Western Andes had the highest concentrations of such species—100 in total—but the lowest densities of national parks. We then adjusted the priorities for this region by refining these species ranges by selecting only areas of suitable elevation and remaining habitat. The estimated ranges of these species shrank by 18–100% after accounting for habitat and suitable elevation. Setting conservation priorities on the basis of currently available range maps excluded priority areas in the Western Andes and, by extension, likely elsewhere and for other taxa. By incorporating detailed maps of remaining natural habitats, we made practical recommendations for conservation actions. One recommendation was to restore forest connections to a patch of cloud forest about to become isolated from the main Andes. Establecimiento de Prioridades Practicas para la Conservacion de Aves en los Andes Occidentales de Colombia Resumen Intentamos fijar prioridades de conservacion de manera que conduzcan a acciones directas de conservacion. La elaboracion de mapas estrategicos de gran escala lleva a prioridades de conservacion ejemplificadas por los sitios importantes para la biodiversidad. En contraste, las acciones tacticas de conservacion se desarrollan en extensiones geograficas mucho menores y que necesitan reflejar la perdida de habitat y la fragmentacion que han limitado severamente el espacio en que viven actualmente las especies. Nuestras aspiraciones para acciones directas, practicas fueron demandantes. Primero, identificamos las prioridades de conservacion globales, estrategicas y luego redujimos la escala a acciones locales practicas en el contexto de las prioridades seleccionadas. Al hacerlo, reconocimos las limitaciones de la informacion incompleta. Iniciamos tal proceso en Colombia y utilizamos los resultados presentados aqui para implementar la reforestacion de terrenos degradados para prevenir el aislamiento de un area extensa de bosque de niebla. Utilizamos los mapas de distribucion de 171 especies de aves para identificar las areas prioritarias de conservacion que pudieran conservar el mayor numero de especies en riesgo en Colombia. Nos referimos como especies en riesgo a aquellas que son endemicas y tienen rangos de distribucion pequenos. Los Andes Occidentales tuvieron las concentraciones mayores de tales especies – 100 en total – pero la menor densidad de parques nacionales. Posteriormente ajustamos las prioridades para esta region al refinar los rangos de distribucion de estas especies seleccionando solo areas con la adecuada elevacion y remanentes de habitat. Los rangos de distribucion estimados para esas especies se redujo en 18-100% despues de considerar el habitat y elevacion adecuados. La definicion de prioridades con base en los mapas de distribucion disponibles actualmente excluyeron areas prioritarias en los Andes Occidentales y, por extension, probablemente en otras areas y taxa. Al incorporar mapas detallados de los habitats naturales remanentes, hicimos recomendaciones practicas para acciones de conservacion. Una de ellas fue restablecer la conectividad hacia el parche de bosque de niebla que esta a punto de quedar aislado de los cadena de los Andes.

Summary Most of the world's remaining habitats are split into small fragments that lose species quickly. Knowledge of this fact can guide practical actions to prevent extinctions.

Habitat loss is the principal threat to species. How much habitat remains-and how quickly it is shrinking-are implicitly included in the way the International Union for Conservation of Nature determines a species' risk of extinction. Many endangered species have habitats that are also fragmented to different extents. Thus, ideally, fragmentation should be quantified in a standard way in risk assessments. Although mapping fragmentation from satellite imagery is easy, efficient techniques for relating maps of remaining habitat to extinction risk are few. Purely spatial metrics from landscape ecology are hard to interpret and do not address extinction directly. Spatially explicit metapopulation models link fragmentation to extinction risk, but standard models work only at small scales. Counterintuitively, these models predict that a species in a large, contiguous habitat will fare worse than one in 2 tiny patches. This occurs because although the species in the large, contiguous habitat has a low probability of extinction, recolonization cannot occur if there are no other patches to provide colonists for a rescue effect. For 4 ecologically comparable bird species of the North Central American highland forests, we devised metapopulation models with area-weighted self-colonization terms; this reflected repopulation of a patch from a remnant of individuals that survived an adverse event. Use of this term gives extra weight to a patch in its own rescue effect. Species assigned least risk status were comparable in long-term extinction risk with those ranked as threatened. This finding suggests that fragmentation has had a substantial negative effect on them that is not accounted for in their Red List category.

S1 File. Additional methods and supporting results and references. This file contains Supporting Information Materials and Methods; Figure A (study schematic), Figure B (23 time series and best-fit population models); Figure C (partial residual plots of components of model averaged GLM to explain PIKE values from MIKE sites); Figure D (comparison of GAMs to explain extracted stable densities with and without influential points); Figure E (histogram of ecological benchmark estimates for each of 73 protected areas illustrating results of Monte Carlo simulation to incorporate uncertainty); Figure F (histogram of cumulative ecological benchmark across 73 protected areas recalculated for each of 1x10^6 runs from the Monte Carlo simulation to incorporate uncertainty); Table A (alternative candidate models to describe the population dynamics of 23 time-series populations); Table B (summary information on time series and extracted stable density and SE); Table C (summary information for 43 MIKE sites and the explanatory variables used to explain PIKE); Table D (Summary statistics and variables included in most likely quasi-binomial generalized linear models to explain PIKE for 43 MIKE sites across Africa and final predictive average model used to generate PIKE estimates for non-MIKE sites); Table E (selection parameters of candidate generalized additive models explaining variation in extracted stable population size for 18 populations); Table F (summary information, predicted stable density (give current PIKE), and ecological benchmark density (given zero PIKE), and comparisons between most recent density and population size estimates and ecological benchmark density and population size for 73 protected areas across 21 countries); and Supporting Information References.

The Red List classification of species’ risks of extinction has seriously underestimated the risks of many species., The IUCN (International Union for Conservation of Nature) Red List classifies species according to their risk of extinction, informing global to local conservation decisions. Unfortunately, important geospatial data do not explicitly or efficiently enter this process. Rapid growth in the availability of remotely sensed observations provides fine-scale data on elevation and increasingly sophisticated characterizations of land cover and its changes. These data readily show that species are likely not present within many areas within the overall envelopes of their distributions. Additionally, global databases on protected areas inform how extensively ranges are protected. We selected 586 endemic and threatened forest bird species from six of the world’s most biodiverse and threatened places (Atlantic Forest of Brazil, Central America, Western Andes of Colombia, Madagascar, Sumatra, and Southeast Asia). The Red List deems 18% of these species to be threatened (15 critically endangered, 29 endangered, and 64 vulnerable). Inevitably, after refining ranges by elevation and forest cover, ranges shrink. Do they do so consistently? For example, refined ranges of critically endangered species might reduce by (say) 50% but so might the ranges of endangered, vulnerable, and nonthreatened species. Critically, this is not the case. We find that 43% of species fall below the range threshold where comparable species are deemed threatened. Some 210 bird species belong in a higher-threat category than the current Red List placement, including 189 species that are currently deemed nonthreatened. Incorporating readily available spatial data substantially increases the numbers of species that should be considered at risk and alters priority areas for conservation.

A recent report assessing the world's plant species finds continuing progress in completing the taxonomic catalog. However, many undescribed species remain. The report concludes that, presently, 21% of known species risk extinction. We show this statement applies to the short-term and ignores the as-yet undescribed species, which are also likely at risk of extinction. Human actions will extirpate many more by the end of this century.

Migratory seabirds face threats from climate change and a variety of anthropogenic disturbances. Although most seabird research has focused on the ecology of individuals at the colony, technological advances now allow researchers to track seabird movements at sea and during migration. We combined telemetry data on Onychoprion fuscatus (sooty terns) with a long-term capture-mark-recapture dataset from the Dry Tortugas National Park to map the movements at sea for this species, calculate estimates of mortality, and investigate the impact of hurricanes on a migratory seabird. Included in the latter analysis is information on the locations of recovered bands from deceased individuals wrecked by tropical storms. We present the first known map of sooty tern migration in the Atlantic Ocean. Our results indicate that the birds had minor overlaps with areas affected by the major 2010 oil spill and a major shrimp fishery. Indices of hurricane strength and occurrence are positively correlated with annual mortality and indices of numbers of wrecked birds. As climate change may lead to an increase in severity and frequency of major hurricanes, this may pose a long-term problem for this colony.

Habitat loss and fragmentation has long been considered the primary cause for biodiversity loss and ecosystem degradation worldwide, and is a key research topic in landscape ecology (Wu 2013). Habitat fragmentation often refers to the reduction of continuous tracts of habitat to smaller, spatially distinct remnant patches, and habitat loss typically occurs concurrently with habitat fragmentation (Collinge 2009). Although some habitats are naturally patchy in terms of abiotic and biotic conditions (Wu and Loucks 1995), human actions have profoundly fragmented landscapes across the word (Haddad et al. 2015), altering the quality and connectivity of habitats. Therefore, understanding the causes and consequences of habitat fragmentation is critical to preserving biodiversity and ecosystem functioning. From May 4th to 10th, 2015, an International Workshop on Habitat Fragmentation and Biodiversity Conservation, held at the Thousand Island Lake, Zhejiang, China, discussed threats to biodiversity in fragmented landscapes and how fragmentation research can identify and help mitigate these threats. To meet these challenges, the Workshop had three goals. The first was to synthesize key findings in fragmentation science. Second was to identify important remaining research questions concerning the relationships between habitat fragmentation, biodiversity, and ecosystem functioning at local, regional, and global scales. Finally, we examined the unique roles of field-based fragmentation experiments in addressing these questions. The Workshop’s findings are relevant to the broader ecological community, and we present them here to stimulate research that will advance landscape ecology and conservation biology.

The World Bank is the largest international funder of biodiversity conservation. It invests in protected areas to conserve species and spaces, protect ecosystems, and provide food, shelter, and other ecosystem services to local communities. It spends on average, $275 million annually supporting parks in developing countries. We examined their protected areas investment portfolio from 1988 to 2008 to understand how they allocate these funds. We found that more money is allocated to countries with progressively larger GDPs. Many, but not all, of these investments correlate with consensus opinions of high biodiversity priorities. But the World Bank's investments are not proportional; poorer countries receive relatively more funds than richer ones, regardless of biodiversity importance. We suggest that these investments focus on supporting parks that provide benefits to local communities, particularly in poorer nations, rather than on biodiversity priorities in a vacuum. This mirrors their mission to work for a world without poverty.

We synthesize findings to date from the world’s largest and longest-running experimental study of habitat fragmentation, located in central Amazonia. Over the past 32 years, Amazonian forest fragments ranging from 1 to 100 ha have experienced a wide array of ecological changes. Edge effects have been a dominant driver of fragment dynamics, strongly affecting forest microclimate, tree mortality, carbon storage, fauna, and other aspects of fragment ecology. However, edge-effect intensity varies markedly in space and time, and is influenced by factors such as edge age, the number of nearby edges, and the adjoining matrix of modified vegetation surrounding fragments. In our study area, the matrix has changed markedly over the course of the study (evolving from large cattle pastures to mosaics of abandoned pasture and regrowth forest) and this in turn has strongly influenced fragment dynamics and faunal persistence. Rare weather events, especially windstorms and droughts, have further altered fragment ecology. In general, populations and communities of species in fragments are hyperdynamic relative to nearby intact forest. Some edge and fragment-isolation effects have declined with a partial recovery of secondary forests around fragments, but other changes, such as altered patterns of tree recruitment, are ongoing. Fragments are highly sensitive to external vicissitudes, and even small changes in local land-management practices may drive fragmented ecosystems in markedly different directions. The effects of fragmentation are likely to interact synergistically with other anthropogenic threats such as logging, hunting, and especially fire, creating an even greater peril for the Amazonian biota.

There really are more than two kinds of people in this world. It’s just that in addition to the two common ones, the third class is rare. There are those of us for whom slinging a hammock between two trees, draping first a mosquito net over it, then a rain fly, is as close to a definition of paradise as imaginable. There are those for whom this would be cruel and unusual torture. And, then there is Elizabeth Kolbert, who might have remained one of the latter, but has the pluck to sling her hammock and write about it.Walking through the forests of Panama, her guides point out the soldier ants, which leave their jaws in your legs after they’ve bitten. They look for, but do not find, the most venomous snakes, that “can really mess you up” and, eventually, the hammock site. “A slit in the bottom constituted the entryway, and … when I climbed into the thing, I felt as if I were lying in a coffin.” In this coffin, she has “vivid troubled dreams” of “bright yellow frogs”. Forest spirits do us — human actions are driving species to extinction at rates only seen five previous times in geological history.Kolbert is brave in another sense. Her main title is the same as the book by distinguished science writer Roger Lewin and famous anthropologist Richard Leakey, published in 1996. Kolbert is a staff writer at The New Yorker, a popular US magazine, and brings an essayist’s approach to the topic. Indeed, her book is a set of thirteen essays, with a three-page introduction and no summarising thoughts.About half the book is about geological extinctions — the five prior to the current one. These chapters cover familiar ground, with mastodons and ammonites, and familiar personalities — Cuvier, Lamarck, Lyell, Darwin, and more recently the father and son Alvarezes and their iridium layer.The remaining story of current extinctions is told well in essays on ocean acidification, fragmented forests, climate disruption, and the consequences of our carelessly moving plant and animal species to places where they do not belong.Kolbert captures the uniqueness of cloud forests. Her guide recommends that she pick out a leaf with an interesting shape. “You’ll see it for a few hundred metres and then it will be gone. That’s the tree’s entire range.”Kolbert is now getting used to fieldwork. Handed a handful of coca leaves and a pinch of baking soda, she finds they taste like “old books” — but the aches and pains of high altitude hiking in cloud forests soon vanish. (One of my perennial worries is to ensure that those I guide do not accidentally pack the coca tea bags from nightstands of good hotels in Quito when they return to the USA.) Spectacled bears have been through the campsite. Now, her visions are of Paddington Bear, the children’s story character, not of haunting frogs.The trees are moving uphill, too — or at least their descendants. The climate is warming fast enough that careful fieldwork readily documents their uphill dispersals. As they move, there is less room for each species and, perhaps in time, insufficient for their continued survival. Climate disruption — for it is more complex than mere warming — not only has the potential to exterminate species, but to eliminate species hitherto unaffected by the deforestation that disproportionately endangers lowland species.By the time Kolbert gets to the lowland Amazon, there is but the briefest mention of her hammock at the famous Camp 41 north of Manaus. (And she takes for granted the showers and toilets that may constitute my greatest contribution to tropical ecology: I chaired the committee that recommended their installation.) When noisy mating frogs wake her in the middle of the night, she enthusiastically steps out to look for them.The camp is the centre of the largest and most significant ecological experiment. As Kolbert explains, we live in a world where what habitats remain are usually in fragments, some small, some large. Nearly forty years ago, Tom Lovejoy recognised the global significance of fragmented landscapes and posed the critical questions: how many species will they lose and how fast will they lose them? Smaller fragments lose more species and do so more quickly. The experiments here, with their carefully designed fragment sizes, provide quantification that guides practical conservation actions elsewhere [1xConservation: forest fragments, facts, and fallacies. Pimm, S.L. and Brooks, T. Curr. Biol. 2013; 23: R1098–R1101Abstract | Full Text | Full Text PDF | PubMed | Scopus (4)See all References[1].Kolbert’s strength is in her essays. They are credible, well-selected and entertaining reads. Her weakness, alas, is the wider perspective. First, she does not attempt to justify why we should consider the present “the sixth extinction.” Bright yellow frogs may well haunt her dreams. Frogs have exceptional extinction rates — nearly 600 extinctions per million species per year. That’s some 6,000 times the background rate before human actions inflated it. How we derive these facts requires explanation and a story more remarkable than anything Kolbert presents.The International Union for Conservation of Nature compiles the Red List — the species that are in danger of becoming extinct or have already done so. It’s remarkable as the combined effort of hundreds of thousands of people, many of them amateurs, to compile data on the status of their favorite taxa. For amphibians, we know that there were 1437 species described by taxonomists before 1900. Of these, 18 are either extinct or presumed extinct. After 1900, taxonomists described 4967 species, for which 121 are extinct or presumed so. Following these newer discoveries from the time of their description allows an estimate of 587 extinctions per million species, per year. This contrasts to a background rate of about one extinction per ten million species, per year. These numbers attest to two other facts: first, the rate of species description is still increasing — new species (Figure 1Figure 1) are being found in places that were until recently inaccessible. Second, newly described species have very high rates of extinction and threat. They are rare — which is why we have only just found them — and in places where there is extensive habitat loss.Figure 1New species.Andinobate cassidyhornae is a recently described species of frog from the Western Andes of Colombia. It lives in forest protected by a land purchase made possible by SavingSpecies (www.savingspecies.org). (Photo: Luis Mazariegos.)View Large Image | View Hi-Res Image | Download PowerPoint SlideSecond, like too many others — such as diCaprio’s The 11th Hour or Call of Life: Facing the Mass Extinction — Kolbert details the problems, but none of the solutions. One wonders, were she to write about modern medicine, would she portray all physicians as telling us we are all going to die — which is true — but fail to notice that their profession greatly reduces mortality and disease? Tens of thousands of conservation professionals worldwide save species on the brink and are measurably reducing extinction rates. The fraction of the Earth’s surface now protected has increased steadily, while Brazil has massively reduced deforestation in the Amazon. From these and so many other stories, we surely conclude that the sixth extinction is not yet written. It’s a nightmare from which we can still escape.

Predicting whether the ranges of tropical species will shift to higher elevations in response to climate change requires models that incorporate data on topography and land use. We incorporated temperature gradients and land-cover data from the current ranges of species in a model of range shifts in response to climate change. We tested four possible scenarios of amphibian movement on a tropical mountain: movement upslope through and to land cover suitable for the species; movement upslope to land-cover types that will not sustain survival and reproduction; movement upslope to areas that previously were outside the species' range; and movement upslope to cooler areas within the current range. Areas in the final scenario will become isolated as climate continues to change. In our scenarios more than 30% of the range of 21 of 46 amphibian species in the tropical Sierra Nevada de Santa Marta is likely to become isolated as climate changes. More than 30% of the range of 13 amphibian species would shift to areas that currently are unlikely to sustain survival and reproduction. Combined, over 70% of the current range of seven species would become thermally isolated or shift to areas that currently are unlikely to support survival and reproduction. The constraints on species' movements to higher elevations in response to climate change can increase considerably the number of species threatened by climate change in tropical mountains.

We estimate the probable number of flowering plants. First, we apply a model that explicitly incorporates taxonomic effort over time to estimate the number of as-yet-unknown species. Second, we ask taxonomic experts their opinions on how many species are likely to be missing, on a family-by-family basis. The results are broadly comparable. We show that the current number of species should grow by between 10 and 20 per cent. There are, however, interesting discrepancies between expert and model estimates for some families, suggesting that our model does not always completely capture patterns of taxonomic activity. The as-yet-unknown species are probably similar to those taxonomists have described recently—overwhelmingly rare and local, and disproportionately in biodiversity hotspots, where there are high levels of habitat destruction.

Elephant are increasing across some areas of Africa leading to concerns that they may reduce woodlands through their feeding. Droughts may help limit elephant numbers, but they are generally both episodic and local. To explore more general impacts of rainfall, we examine how its annual variation influences elephant survival across ten sites. These sites span an almost coast-to-coast transect of southern Africa that holds the majority of the ~500,000 remaining savanna elephants. Elephants born in high rainfall years survive better than elephants born in low rainfall years. The relationship is generally weak, except at the two fenced sites, where rainfall greatly influenced juvenile survival. In these two sites, there are also extensive networks of artificial water. Rainfall likely affects elephant survival through its influence on food. The provision of artificial water opens new areas for elephants in the dry season, while fencing restricts their movements in the wet season. We conclude that the combination of these factors makes elephant survival more susceptible to reductions in rainfall. As a result, elephants living in enclosed reserves may be the first populations to feel the impacts of global warming which will decrease average rainfall and increase the frequency of droughts. A way to prevent these elephants from damaging the vegetation within these enclosed parks is for managers to reduce artificial water sources or, whenever practical, to remove fences.

Endangered species recovery plans are frustrated by small, spatially structured populations where understanding the influence of birth, death, and dispersal is difficult. Here we use a spatially explicit, long-term study to describe dispersal in the Cape Sable seaside sparrow (Ammodramus maritimus mirabilis). Since 1990, this species declined > 50%. It occurs as several geographically isolated subpopulations in the Florida Everglades. We characterize dispersal, recognizing that our sampling, as well as the species’ distribution, is spatially heterogeneous. The annual movements of juveniles and adults are statistically heavy-tailed. That is, while most individuals are recaptured locally, a significant portion exhibit long-distance dispersal. Individuals move between subpopulations to distances >30 km. Not accounting for the spatial heterogeneity of sampling or the species range itself underestimates dispersal and can lead to ineffective management decisions. Recovery focused on translocation will be less successful than strategies that protect habitat and increase breeding.

How many species are likely as-yet unknown to science? Even in relatively well-known groups, there may be substantial numbers of such species. It seems likely that these unknown species will be rare and threatened with extinction. Indeed, science may not discover them before they go extinct. We address these issues for a sample of endemic flowering plants and three vertebrate groups: amphibians, birds, and mammals, all from Brazil. We predict the likely numbers of missing species from models of the declining numbers of species described per five-year interval. The raw numbers increase over time, so we must scale these by the taxonomic effort. We show that while the catalogues of birds and mammals are nearly complete, the numbers of amphibians may increase by 15% and the numbers of endemic plants by ~10 to ~50% depending on region. These percentages may still seem encouragingly low, given the complexities of studying a country as large as Brazil, with its extraordinary diversity, and with many of its regions large and still poorly explored. What is more worrying is that these numbers of as-yet unknown species are broadly the same as the percentages of species that are presently considered threatened with extinction. That is, we know only half of the species in danger of extinction – and our knowledge of even those species has mostly been acquired in the last three decades.

The IUCN Redlist considers the African savannah elephant ( Loxodonta africana ) to be “vulnerable” despite it numbering in the 100,000s and having a large geographical range. This seeming paradox stems from how quickly human persecution can eliminate elephants across large areas and how quickly elephant numbers can increase when protected. Much elephant research concentrates on the extent and consequences of elephant persecution. Where elephants thrive, two other human interventions, the provision of artificial water and the construction of fences may have large, and perhaps unintended, impacts on elephant behavior. In general, successful management requires that we understand elephant movements and land-use choices across large areas and long periods. Here, we ask specifically how artificial water and fences might affect these movements. To do this, we first characterize how elephants move in different seasons and landscape types, in different years, and how these patterns change over a region that varies considerably in annual rainfall. We fitted 73 elephants with GPS collars across a large rainfall gradient spanning seven southern African countries over a period of 6 years. We analyzed remotely-sensed environmental data from four satellite borne sensors that measure daily rainfall, weekly temperature, bi-monthly greenness, and summarise human infrastructure. Elephants move approximately 6 km/day in dry landscapes, down to approximately 3 km/day in the wettest ones. Strong seasonal differences modulate geographic differences. Elephants move less, cover less area, and are more faithful to landscapes across years in the dry season than the wet. Water availability drives these seasonal patterns. Seasonal differences in the area covered are less pronounced in wet landscapes where permanent water is more dispersed. Within-day movements reveal that elephants are consistently crepuscular but more active at night than midday when temperatures are high. Direction-changes are centered at midnight when elephants are close to water indicating regular nighttime treks to water. By design, our analyses seek to find general patterns of elephant movements — something that one can achieve only across a large range of locations and ecological conditions — in order to understand the impact of human interventions. We show that both interventions reduce seasonal differences in elephant ranging patterns and increase local impacts of elephants on the vegetation. Artificial water sources allow more extensive dry season ranging, allowing elephants to use — and potentially overexploit — vegetation in areas that would have be otherwise inaccessible to them except in the wet season. Fences cause elephants to “bunch-up” against them during the wet season, again locally increasing the pressure elephants put on their resources.

Translocations are increasingly important tools for endangered species conservation, but their success is often uncertain. We analyzed 125 time series of grazing mammal translocations in South African protected areas. Some 94% of translocations succeeded (66% unambiguously) even though most populations began with